ICONES PLEUROTHALLIDINARUM
XII
i/, 5-7
SYSTEMATICS
OF
BRACHIONIDIUM
ADDENDA TO DRESSLERELLA, PLATYSTELE AND PORROGLOSSUM
(ORCHIDACEAE)
Missouri Botanical Garden
ICONES PLEUROTHALLIDINARUM
XII
SYSTEMATICS
J996
OF
BRACHIONIDIUM
ADDENDA TO DRESSLERELLA, PLATYSTELE AND PORROGLOSSUM
(ORCHIDACEAE)
Carlyle A. Luer
m
Missouri Botanical Garden
MONOGRAPHS IN SYSTEMATIC BOTANY
FROM THE MISSOURI BOTANICAL GARDEN
Volume 57
Published in December 1995 in an edition
of 500 copies.
ISSN 0161-1542
ISBN 0-915279-36-3
CariyleA.Luer
3222 Old Oak Drive
Sarasota, FL 34239
Publications Staff
Magdalen Larape
Robin Bruce
Bradenton, FL
Composed at 3222 Old Oak Drive. Sarasota, Florida
with WordStar 7.0
Typeset with a Texas Instruments ri
!
CONTENTS
Brachionidium uxorium .
Introduction to Brachionidium .
Key to the species.
List of the species.
The species.
References.
Index to the species.
Addendum to Dresslerella .
Addendum to Platystele .
Addendum to Porroglossum .
A Re-evaluation of the Pleurothallid Subgenera Satyria and Silenia..
.1-4
.5-9
.10-11
.12-137
.138
.139
.140
.142
.144
.146
e sea level, 14 Not.
1992, photo by Waller Teague.
SYSTEMATICS OF THE GENUS
BRACHIONIDIUM (ORCHIDACEAE)
white drawing. A distribution map is included for each species.
Brachionidium satyreum Luer. sp. nov.
Brachionidium stellare Luer & Hirtz, sp. nov.
Brachionidium symc-morrbii Luer, sp. nov.
Brachionidium uxorium Luer& V^uez. sp. nov.
Brachionidium zunagense Luer & Hirtz, sp. nov.
From a collection made by Jean Linden on his trip to New Granada to gather
plants for the trade, John Lindley described the first species of this genus in Restre-
pia in 1846. The specific name referred to the comparatively small leaves
as compared to those commonly seen in Restrepia. In 1859, Lindley erected the
genus Brachionidium to accommodate this exceptional Restrepia parvifolia along
with later collections of two other species. The name refers to the pair of tiny
“arms” of the stigma that protrude from either side of the anther. The epithet
parvifolium became a misnomer because the leaves have proved to be some of the
largest in the genus. Brachionidium parvifolium is one of the most robust and most
beautiful, and one of the most frequent and widely distributed species of the genus.
In none of the three original specimens attributed to Brachionidium was Lindley
able to find and describe the pollinia. The first reported number of pollinia in a
species of the genus was six by Rolfe when he described B. sherringii, the fourth
ICONES PLEUROTHALUDINARUM
species attributed to the genus, from the West Indian isle of Grenada in 1893.
Schlechter saw six in the Peruvian Brachionidium serratum in 1911. In 1919,
Hoehne found eight poUinia in a species from Brazil, and he described it in a new
genus, Yolanda, because, he thought, Brachionidium had only six pollinia. Garay
(1956) followed him, declaring that Brachionidium was distinguished by six polli¬
nia, and Yolanda, by eight pollinia. In the same publication he redescribed the rela¬
tively frequent B. tuberculatum, a species with eight pollinia, as B.floribundum.
Recently, ample fresh material of B. parvifolium, the type of the genus, has been
examined from several different localities in both Colombia and Ecuador. Eight
pollinia are consistently present. The poUinaria from numerous other species have
been examined, and either six or eight pollinia are present, there being no correla¬
tion with any morphological feature. Therefore, if two numbers of pollinia must be
acceptable in Brachionidium, two numbers of pollinia could be expected to occur in
other pleurothallid genera (i.e. Pleurothallis: P. ophiocephala Lindl. and P. kegellii
Rchb.f.). About 24 species have eight pollinia, and most of these are restricted to
South America. Brachionidium kuhniarum is the only species known in which two
sets of three pollinia are ncvmally joined at the clavate ends.
Today, the genus Brachionidium contains 64 known species widely distributed
in tropical America. Most of them are local and infrequent in wet, virgin forests.
Many grow terrestrially, erect or creeping, in shaded, loose, mossy or leafy litter,
but some species trail like a vine in tall trees, and flower only when sufficient light
is reached. Some species grow erect in relatively exposed Imbitats. Other species
grow fully exposed tenestrially or in stunted trees of cold paramos over 3000
meters above sea level.
Many plants flower infrequendy and the flowers are fragile, often lasting only a
day or two. Therefore, to our dismay, plants are too often without flowers when
found, and most are difficult or impossible to cultivate. At least two sterile collec¬
tions of undescribed species are known. Herbaria of the worid contain comparative¬
ly few collections. Wild plants are either neglected or unrecognized because many
collectors are unfamiliar with them. Many of the collections that are made are
either sterile or with poorly preserved flowers.
Some of the following illustrations were made from dried flowers removed from
herbarium sheets and reconstituted in concentrated ammonium hydroxide. Ob¬
viously, far superior drawings can be made from fresh or freshly pickled material,
but in some instances, only dried and pressed material has been available. Fresh
flowers dried without too much pressure rehydrate very well, while those flowers
crushed and dried tightly in a plant press are impossible to resurrect satisfactorily.
The species are too closely rela^ to each other to suggest any subgeneric divi¬
sion. All plants are char^terized by sheathed, stemlike, branching rhizomes that
may creep, ascend or stand erect, and produce at intervals a short, similarly
sheathed aerial stem (the ramicaul) and its leaf. The sheaths are usually long-
mucronate and glabrous, but in a few species the sheaths are scurfy with papilliform
scales that are often seen along the inferior margins.
The ephemeral flowers are solitary and usually non-resupinate with four free,
membranous flower parts: a middle sepal, a synsepal composed of the connate
blades of the lateral sepals, a large pair of petals mwe or less similar to the sepals,
plus the Up. The apex or the tip of the taU of the synsepal is entire, shortly bifid, or
SYSTEMATICS OF BRACHIONIDIUM
The lip (Figure 1, page 3; Figure 2, page 11) is small, thick and usually trans¬
verse with a central callus, reminiscent of the lip of a Stelis. There are four surfac¬
es. The lip is often variable, especially in widely distributed species such as B.
tuberculatum. The upper or top surface is held beneath the column, with some
modifications of a pubescent callus. The front surface is variably sculpted, more or
less concave and convex. A shiny, rounded structure, the glenion (commonly seen
in Platystele, Pleurothallis and Stelis), often extends forward from the callus onto
the front surface. The back surface is flat or modified by deflection of the base, or
claw, to the attachment to the obsolescent column-foot. The sides are more or less
triangular in outline. The anterior margin of the lip varies from entire, fringed, or
variously callous. A callus often runs parallel to the margin, sometimes with a
cavity between the two edges, thereby forming a bimarginate margin. Oxalate
The column is short and broad with an apical anther and a transverse, apical,
bilobulate stigma. The lobes appear to be separate, but beneath the rostellar flap
they are contiguous, as they are in Lepanthopsis, Platystele, most species of Pleur¬
othallis subgen. Pleurothallis and most species of Stelis. The tips of the stigmatic
processes, which prompted the generic name, project forward above the anterior
margin of the rostellum beneath the bed of the anther. The anther cap is thin and
deciduous. The pollinia are narrowly pyriform or clavate, entirely free or attached
to a small viscidium. Sometimes the pollinia vary in size. Sometimes all six or
eight pollinia are bound by a single viscid mass, and sometimes two sets of three or
four pollinia are bound by viscid masses.
narginate lip, columi
. Lower left: front SI
. Upper right: eight pollinia; oblique v
ICONES PLEUROTHALUDINARUM
Lectotype: Restrepia parv^olia Lindl., Orchid, lind., 4,1846. [i
LindlVFolia Orchid. Brachionidium 1. 1859] (Garay, Canad. J. Bot. 34:729,1956)
Syn.: Yolanda Hoehne, Arq. Mus. Nac. Rio de Janeiro 22:72,1919.
Type: Yolanda restrepioides Hoehne, Arq. Mus. Nac. Rio de Janeiro 22:72,1919.
[BrachionidUm restrepioides (Hoehne) Pabst, Orchid. Brasilienses 166,1975.
Ety.: Named in honor of Ydanda, daughter of F. C. Hoehne, Brazilian botanist.
Syn.: Brachionidium sect. Yolanda (Hoehne) Pabst, Bradea 1(24): 269,1972.
Type: Yolanda restrepioides Hoehne, Arq. Mus. Nac. Rio de Janeiro 22:72, 1919.
Contrary to one’s first impression, especially because two numbers of pollinia
are present, the species are too closely interrelated to define subgeneric categories.
The species are first identified in the key by the most obvious, gross morphological
features, and lasdy by more obscure or microscopic characters. The most obvious
dichotomous split seems to be the habit, whether it be erect or creeping, but both
habits may occur in some species. Therefore, a few species appear more than once
in the key. Similar species a^iear grouped together in the key. There is no obvious
morphological correlation between the number of pollinia and another physical
feature. The key is totally artificial.
SYSTEMATICS OF BRACHIONIDIUM
KEY TO THE SPECIES OF BRACHIONIDIUM
1 Habit more or less erect or erect-climbing, the roots mostly produced near the
base of the rhizome (if indefinite, try 48).2
r Habit scandent, more or less prostrate, repent or climbing, the roots produced
singly from nodes along the rhizome (if indefinite, try 2)
Habit erect, sheaths scurfy or with tuberculate margins
2 Sheaths of the rhizome and ramicauls scurfy, or with the margins crenulate or
2’ Sheaths of the rhizome and ramicauls neither scurfy, nor with crenulate or tuber¬
culate margms.10
3 Leaves rugose with vemicose, elevated veins; synsepal cucuUate. B. galeatum
3’ Leaves smooth, not rugose..4
4 Synsepal deeply concave, cucullate.5
4’ Synsepal concave, not cucullate.V
5 Synsepal with the tail as long as or longer than the blade. B. arethusa
5’ Synsepal with the tail much shorter than the blade.6
6 Leaves 10-14 mm wide; lip much broader than long. B. elegans
6’ Leaves 4-6 mm wide; lip about as broad as long. B.furfuraceum
7 Sepals more than 3 cm long. B. alpestre
T Sepals less than 2 cm long.8
8 Lateral angles of the lip obtuse, but more or less incurved. B. tuberculatum
8’ Lateral angles of the lip acute.9
9 Sepals prominently striped in purple . B. uxorium
9’ Sepals green to purple, not striped. B. cruzae
Habit erect, sheaths apiculate without epidermal appendages,
plants more than 10 centimeters tall
10 Mature plant more than 10 cm tall, the intemodes of the rhizome often longer
than the ramicauls.11
10’ Mature plant less than 10 cm tall, the intemodes of the rhizome often shorter
than the ramicauls. 21
11 Leaves rugose, with vemicose, elevated veins 12
11 ’ Leaves smooth, not rugose.13
BK
ICONES PLEUROTHALUDINARUM
12 Sepals and petals minutely ciliate; lip denticulate. B. dentatum
12’ Sepals and petals glabrous; lip not denticulate. B. rugosum
13 Leaves suboibicular or broadly elliptical with the apex rounded. B. loxense
13’ Leaves elliptical with the apex acute to obtuse.29
14 Pedicel thrice or more longer than the leaf.. B. vasquezii
14’ Pedicel shorter to about twice as long as the leaf. .15
15 Peduncle short, not longer than the ramicaul. B. dalstroemii
15’ Peduncle nearly as long to about twice as long as the leaf..16
16 Middle sepal 5- or more veined. 17
16 Middle sepal 3-veined_ 19
17 Synsepal deeply concave with the split tail shorter than the blade.
. B.fornicatwn
17’ Synsepal not deeply concave, with tails elongate.18
18 Sepals 5- to 7-veined, with tails more or less equaling the blades in length.
.... parvifolium
18 Sepals 11- to 15-veined, with tails much exceeding the blades in length.
. imperiale
19 Synsepal with a slender tail longer than the blade.
19’ Synsqtal tailless or with a tail shorter or as long as the blade..
20 Lip with prominent lateral angles..
..20
..21
...22
21 Lateral angles broadly rounded with the apex recurved.. B calypso
21’Uteral angles antrorse, narrow and fringed. b lucanoideum
Margin of the lip denticulate.
Margin of the lip entire.
CaUus extending down the back surface of the lip. b. satyreum
Callus not extending down the back of the lip. .5 hirtzii
Middle sepal abrupdy caudate; lip denticulate. b hateri
Middle sepal acuminate; lip not denticulate ’ 25
26 Leaves less than 2 cm long; sqials 6-10 mm long
26- Leaves more than 3 cm long; sepals ca. 15 mm long”'
.B. tetrapetalim
. B. andreettae
SYSTEMATICS OF BRACHIONIDIUM
Habit erect, plants less than 10 cm tall
7 Leaves rugose. B. operosum
T Leaves smooth.28
8 Synsepal tailless or shortly acuminate.29
8 Synsepal distinctly caudate, with the tail longer than the blade.35
9 Synsepal narrowly ovate, acute. B. pteroglossum
9’ Synsepal ovate to broadly ovate, obtuse to shortly acuminate.30
0 Lip longer than wide, subcordate-trilobed, ciliate-denticulate.31
0’ Lip transverse, broader than long, entire.32
1 Lip sagittate-triangular with basal lobes retrorse. B. brevicaudatum
r Lip trilobed with lateral lobes antrorse B ciliolatum
2 Sepals and petals glabrous.33
2’ Sepals and petals ciliate.34
3 Pollinia 8. B. restrepioides
34 Synsepal shortly acuminate. B. tetrapetalum
34’ Synsepal obtuse, not acuminate. B. simplex
35 Sepaline tails shorter than to about as long as the blade.36
35’ Sepaline tails twice or more longer than blade.41
36 Middle sepal with the tail less than 10 mm long.37
36’ Middle sepal with the tail more than 12 mm long.38
37 Sepals and petals glabrous; lip acutely deflexed near the middle with a broad
claw. B. peltarion
"iT Sepals and petals minutely ciliate; lip flat, not deflexed. B. parvum
38 Sepals and petals glabrous. B. portillae
38’ Sepals and petals minutely ciliate.39
39 Lip with a broad callus extending down the reflexed back of the lip.
. B. longicaudatum
39’ Lip with a small, central callus.40
40 Lip distinctly bimarginate. B. stellare
40’ Lip smooth to convex within the margins. B. kuhniarum
41 Lip with the margin denticulate.42
41’ Lip entire, not denticulate.45
ICONES PLEUROTHALUDINARUM
42 Peduncle about three times as long as the leaf; sepals with tails less than 20 mm
long. B. lehrmnnii
42’ Peduncle shorter to about as long as the leaf..43
43 Plant less than 2 cm tall; sepals with tails ca. 12 mm long. B. julianii
43’ Plant 4-7 cm tall; sepals with tails more than 25 mm long.44
44 Lip geniculate, with the base deflexed from near the middle, with the margin
denticulate. B.folsomii
44 Lip flat with the base shortly deflexed, with the margin minutely undulate.
45 Lip broadly ovate with a simple margin. B. dodsonii
45’ Lip transverse, broadly obtuse, bimarginate..46
46 Lip with narrow, acute lateral angles. B. syme-morrisii
46’ Lip with obtuse lateral angles..47
47 Peduncles 3-5 cm long; synsepal shallowly concave. B. sherringii
47’ Peduncles less tha 3 cm long; synsepal deeply concave. B. ballatrix
Habit repent
48 Leaves ellipdcal to suborbicular, less than 15 mm long..49
48’ Leaves elliptical to narrowly elliptical, more than 15 mm long.60
49 Petals sharply saw-toothed. B.serratum
49’ Petals not sharply senate...50
51 Peduncles about as long as the leaves; sepals less than 7 mm long.52
51 ’ Peduncles longer than the leaves; sepals 13-15 nun long.54
52 Petals eciliate; lip with broad, antrorse, lateral angles. B. minusculum
52’ Petals ciliate.53
53 Lip with acute, narrowly triangular, lateral angles. B. pusillim
53’ Lip with obtuse lateral angles. B. brachycladium
54 Leaves suborbicular, lip with simple margin, long-apiculate. B. capillare
54’ Leaves elliptical; lip bimarginate, minutely apiculate. B. jesupiae
55 Sepals broadly ovate, shorUy acuminate without tails. B. escobarii
55’ Sepals long-acuminate with slender tails.56
SYSTEMATICS OF BRACHIONIDIUM
56 Sepaline tails about as long as the blade; lip transverse.57
56’ Sepaline tails much exceeding the blades in length; lip ovate-triangular.58
57 Rhizome slender; leaves narrowly elliptical. B. ephemerum
57’ Rhizome stout; leaves broadly elliptical. B. polypodium
59 Leaves 8-10 mm long; sepals 10 mm long....
59’ Leaves 5-6 mm long; sepals 18 mm long.
61 Sepals acute to acuminate, without slender tails.
61’ Sepals contracted into slender tails longer than the blade...
62 Rhizome between ramicauls often as long a
62’ Rhizome between ramicauls mostly shorter than the leaves
63 Synsepal obtuse....,
63 Synsepal acute to a
67 Leaves narrow, 4-6 mm wide; sepals ca. 30 mm long including the tails .
67’ Leaves broad, usually 1-2 cm wide; sepals 20-25 mm long including tails.68
68 Synsepal with the tail entire; lip evenly bimarginate. B. hirtzii
68’ Synsepal with the tail bifid; lip bimarginate with the upper margin overhang¬
ing the lower margin. B. phalangiferwn
ICONES PLEUROTHALUDINARUM
Brachionidium alpestre Luer & >
ballatrix Luer & Hirtz.
I brachycladum Luer & Escobar..
Brachionidium calypso Luer.
Brachionidium capillare Luer & Hirtz
Brachionidium ciliolatum Garay.
Brachionidium concolor Lindl.
Brachionidium diaphanum Luer & V^uez..
Brachionidium dodsonii Luer.
Brachionidium dressleri I
r& Hirtz,
Brachionidium ephemerum Luer & I
Brachionidium escobarii Luer.
iLuer&Hirtz.
Brachionidiumfloribundum Garay =
Brachionidium fobomii Dressier.
Brachionidium fornicatum Luer & Hirtz...!
Brachionidium furfui
I Luer & Hirtz..
i Luer..
n hirtzii Luer.
s Luer & Escobar..,
Brachionidium jesupiae Luer.
Brachionidium julianiiCamevali & Ramirez....
Brachionidium muscosum Luer& V^uez
Brachionidium nebUnense Camevali & Ramirez
1 Luer & Hirtz..
.Plate 1.
.Plate!
.Plate 3.
.Plate 4.
.Plate 5.
.Plate 6.
.Plate 7.
....Plate 8.
.Plate 9.
...Plate 10.
...Plate 11.
...Plate 12.
...Plate 13.
...Plate 14.
...Plate 15.
.Plate 16.
....Plate 17.
.Plate 18.
....Plate 19.
.Plate 20.
.Plate 21.
.Plate 22.
....Plate 23.
.Plate 24.
.....Plate 25.
.Plate 26.
.Plate 27.
....Plate 28.
....Plate 29.
....Plate 30.
...Plate 31.
....Plate 32.
...Plate 33.
.Plate 34.
....Plate 35.
...Plate 36.
....Plate 37.
....Page 84.
...Plate 38.
....Plate 39.
...Plate 40.
SYSTEMATICS OF BRACHIONIDIUM
Brachionidium peltarion Luer.
Brachionidium phalangifenim Garay.
Brachionidium piuntzae Luer.
Brachionidium polypodium Luer.
Brachionidium portillae Luer.
Brachionidium pteroglossum Luer..
Brachionidium puraceense Luer..
Brachionidium pusillum Ames & Schweinf....
Brachionidium renzii Luer.
« restrepioides (Hoehne) Pabst..
sum Luer & Hirtz.
Brachionidium serratum Schltr.
Brachionidium sherringii Rolfe.
Brachionidium simplex Garay.
Brachionidium stellare Luer & Hirtz.
Brachionidium steyermarkii Foldats = Brachionidium tuberculatum
Brachionidium syme-morrisii Luer.
Brachionidium tetrapetalum (Lehm. & Krzl.) Schltr.
Brachionidium tuberculatum Lindl.
Brachionidium uxorium Luer & V^quez.
Brachionidium valerioi Ames & Schweinf....
Brachionidium vasquezii Luer.
Brachionidium zunagense Luer & Hirtz.
.Plate 41.
.Plate 42.
.Plate 43.
.Plate 44.
.Plate 45.
.Plate 46.
.Plate 47.
.Plate 48.
.Plate 49.
.Plate 50.
..Plate 51.
.Plate 52.
.Page 114.
.Plate 53.
.Plate 54.
.Plate 55.
.Plate 56.
.Plate 57.
....Plates 58., 59.
.Plate 60.
.Plate 61.
.Plate 62.
.Plate 63.
ICONES PLEUROTHALUDINARUM
i, flore grandi pulcherrimo, sepalo mediano
Plant large, epiphytic to terrestrial, erect, up to 25 cm tall, the t\
mg, 2-4 cm long between ramicauls, enclosed by 3 scurfy, tubular s
d; br^t inflated, acute, vermcose, 7 mm long, enclosing the pedicel and ovary: pedicel l
“ a filament 5-6 mm long; ovary 2 mm long; sepals yellow, minutely ciliate
3100 m. 21 Jan. 1988, C. Luer. J. Luer, R V^omz <t
J. Solomon 12888 (Htdotype: MO; Isotype; LPB);
tcrmtrial cast of t^ j^s, alt 2770 m, 21 Jan. 1988,
C Luer, J. Luer, R. Vdsquez & C. Luer, J. Luer, R.
Vdsquez A J. Solomon 12914 (MO); t.
This handsome species grows either
epiphytically in low shrubs or terrestrially
on road embankments on both sides of a
high, cold and wet mountain pass in the
Nor Yungas of Bolivia, where it is appar¬
ently endemic. The first known collection
was made in 1984 by A1 Gentry and James
Solomon.
Brachionidium alpestre is most similar
to the Ecuadorian B. elegans, from which
It is distinguished by the large, vermcose
floral bract; a long-acuminate, flat synse-
pal; petals held downward against the
middle sepal; and the broad, obtuse lip._
The sheaths of the stout, erect, rhizomes and ramicauls of both species are scurfy
rout, 1 mm long, pcdlinia 8.
\\>
\ 4 .
dm. is, Ute spaces am Covered wiU, minute, irregular; ceUu to MierBoi'
^flL^h Colombian B. parvifoUum which has stnooA sheaths
but from which they ate also easUy distinguished by the 3-veined middle senal and
5-m 7.veined and 8- „ mom-veinid.
SYSTEMATICS OF BRACHIONIDIUM
Plate 1. Brachionidiumalpestre hu&c&y^n&z
SYSTEMATICS OF BRACHIONIDIUM
Plate 2. Brachionidium andreettaeLim &B3itz
ICONES PLEUROTHALUDINARUM
Leaf erect, coriaceous, narrowly ovate, i
into a petiole 2-3 mm long, 8-1
e 3.5-4.5 a
:x of the ramicaul; floral bract inflated, acute, 5 mm long, «iclosing the pedicel and part of the ovary;
I l<Mig; ovary 3.5 mm long; sepals translucent green, minutely
nm long, the lateral sepals connate into a broadly ovate, acute,
n long, the blade 15-20 mm long, 13 mm wide unexpanded, 6-
tail ca. 20 mm long, the tips free for 5 mm; petals translucent
Q long, the blade 15 mm long, 6 n
wide, 3-veined. the tail 20 mm Icmg, the tip fr
mm long, 4 mm wide, the lateral angles
bade surface, the disc' ‘'
, obtuse, 2
type: K), C. Uerillustr. 14707.
This species, apparently endemic in
north-central Peru, was collected in a
bamboo thicket by the Rio Apisoncho
during the Cambridge Botanical Expedi¬
tion in 1965. No subsequent collection is
known.
Brachionidium arethusa is distin¬
guished by the scurfy cauline sheaths;
large, long-tailed flowers with a concave
synsepal much larger than the three-veined
middle sepal; and a transverse, obtuse lip
with thick margins reflexed onto the back
surface, but the reflection may be an arti¬
fact fixm rehydration.
Brachionidium arethusa is most simi-
lar to the Ecuadorian B. elegans, from
^ stinguished by the long, sepaline tails. Both species possess eight
SYSTEMATICS OF BRACHIONIDIUM
ICONES PLEUROTHALUDINARUM
branching, 3-5 nc
I, terrestrial, subeiect to erect, up to 8 cm tall, the rhizomes
mis, enclosed by 3 imbricating, long-mucronate sheaths; l___
ct, 3-5 mm long, enclosed by 2 mucronate sheaths. Leaf erect to suberect,
.u..»y conaceous, empiicai, acute, apiculate, 15-30 mm long including the petiole 2-3 mm long. 6-12
rm wide,7-vemed, cuneate below mto the petiole. Innoresccnce a solitary, yellow, resupinate flower
W by m en^ slender i^unde 25-30 mm long, with a mucronate bract near the middle and another
^r the base; floral bract long-mucronate. 6 mm long, enclosing the pedicel and much of the ovary;
Mtal 0.5 rom ^g. with a protrading filament 5-6 mm long; ovary 3 mm long; sepals eciliate, the
acule. loog^da^ 25 nm long indodmg laU 14 mn. long. 7
M inlo abroadly ovato-suboibicular. concave synsepal, 23 ran
tag ^adrag ta nnl 3 ran. tag. 8 ran. wide expanded, 4-v.toed; pelals ovara, oM^ne nibacrae,
...-b23 nun tag incloding U,e .ad 14 ran. tag. 6 ram wide. 3-veraed; Up
^ ig. 25 mm wide, with acuU
i, marginally callous, minutely apiculate, the disc w
m cloud foi^t nw Nambija. alt. 1300 m, Feb. 1!
Um (Holotype: MO), C. Luer illustr.
This little species is known only from
the original coUection by Alexander Hirtz
near the gold-mining community of
Nambija in the Cordillera del Condor in
southeastern Ecuador. It is a small,
erect species characterized by broadly
ovate sepals and petals abrupfly contracted
into slender tails. The sepals are eciliate-
the petals are minutely cUiate; and the lip
In habit, it is similar to the Central
American B.folsomii, from which it is
distinguished by the non-denticulate
Wgmate lip. It is also reminiscent oJ
B. shemngti from the Lesser Antilles
which It IS distinguished by shorter
peduncles and broader floral parts with a
more deeply concave dorsal se^.
SYSTEMATICS OF BRACHIONIDIUM
ICONES PLEUROTHALUDINARUM
SYSTEMATICS OF BRACHIONIDIUM
Plate 5. Brachionidiumbrachycladumhi^TScEscobar
ICONES PLEUROTHALUDINARUM
SYSTEMATICS OF BRACHIONIDIUM
Plate 6. Brachionidiumbrevicaudatum Rolfe
ICONES PLEUROTHALUDINARUM
SYSTEMATICS OF BRACHIONIDIUM
Plate 7. Brachionidium calypso Luer
26
ICONES PLEUROTHALUDINARUM
Brachionidium capillare Luer & Hirtz, sp. nov.
Ety.: From the Latin capillaris, “capiUaiy,” referring to the hairhke tails of the sepals and petab.
Plantaperparva repens rhizomate gracile elongate, pedunculo gracile foUo suborbiculato leviter
longiore, s^>ahs niveis eciliatis, sepalo impari ovato aoito longissime caudate, synsep^^guste tnM-
angulis lateralibus acutis, apice obtuse apiculato.
SYSTEMATICS OF BRACHIONIDIUM
Plate 8. Brachionidium capillarehu&r & Witz
ICONES PLEUROTHALUDINARUM
of fi. ciliolatum are minutely ciUolate and without taUs. The lip is longer than broad
with erect, ciliolate, obtuse lobes below the middle, and an acute, ciliolate middle
lobe. The disc is distinctly concave anterior to a central, horse shoe-shaped callus.
SYSTEMATICS OF BRACHIONIDIUM
Plate 9. Brachionidiumciliolatum Garay
PLEUROTHALLIDINARUM
SYSTEMATICS OF BRACHIONIDIUM
ICONES PLEUROTHALUDINARUM
Brachionidium dalstroemii Luer, Lindleyana 1: 170,1986.
Ety.: Named in honor of Stig Dalstrom of Falun, Sweden, who discovered this species.
ECUADOR: Prov. of Loja: epiphytic in cloud forest
between Loja and Saraguro, alt. 3150 m, 17 Mar.
1984. C. Luer, S. Dalstrom, T. Hoijer & J. Kmjt 9553
(Holotype: MO). Prov. of Morona-Santiago: in moist
forest between Churracos and Granadillas, eastern
slopes from Cuenca, alt. 2600-3000 m, F.C. Leh¬
mann 7056 (K); road constraction between Sigsig
and Gualaquiza. 9 Apr. 1968, G. Marling, G. Storm &
B. Strom 8129 (GB); Prov. of Morona-Santiago: east
C L* ^
Teague 13367 (MO). Without locality, F. C. Leh¬
mann 6512 QL).
PERU: Dept, of San Martin: Huallaga, Rio Api-
sondio, 30 Km above Jucusbamba, alt. 3100 m, 27
^ ^ Hamilton & P. M. Holligan 1283
This distinctive species was found by
Stig Dalstrdm in the dense, scrubby vege¬
tation at the pass north of Loja in southern
Ecuador. As is so often the case in this
genus, only one specimen was discovered.
Fortunately, a fine flower was present; too
often specimens of Brachionidium are
found without flowers, a frustrating event. - ■
Since most of these plants of high alUtodes refuse to adapt to cultivation positive
Utauftcauon nt Unpossible. Wore, a visit to the lociSit, is atteC^^to
This species, apparently infrequent and local at high altitudes in southern
Wtth long, slender tads. The lip is transvetse with the edge bintargS^
SYSTEMATICS OF BRACHIONIDIUM
ICONES PLEUROTHALLIDINARUM
Brachionidium dentatum Luer & Dressier, sp. nov.
By.: From the Latin dentatm, “toothed,” refemng to the labeUum.
PANAMA: Prov. of Chiriqui: above Ceiro Punta, alt
^ 2800 m, coUected and cultivated by A. Maduio,
flowered in cuUivalion at Cerro Punta. July 1979 R
(Holotype: MO), C. Luer iUustr
This species is apparenUy endemic in
the wet cloud forest of Cerro Punta in
western Panama where it was discovered
by Robert Dressier and Andres Maduro. It
is distinguished by the erect habit with
elliptical leaves with carinate veins, a
character found in very few species. The
flowers are comparatively small and dark
purple. The sepals and petals are minutely
cUiate and long-caudate. The unique lip is
more or less cordate-ovate and bimar-
ginate with two toothed crests, one within
the outCT margin.
SYSTEMATICS OF
Plate 12. Brachionidium dentatum Luer & Dressier
SYSTEMATICS OF BRACHIONIDIUM
Plate 13.
SYSTEMATICS OF BRACfflONIDIUM
Plate 14. BrachionidiumdodsoniilMeT
PLEUROTHALUDINARUM
SYSTEMATICS OF BRACHIONIDIUM
Plate 15. Brachionidiumdressleri Luer
PLEUROTHALUDINARUM
SYSTEMATICS OF BRACHIONIDIUM
Plate 16. Brachionidium ecuadorense Gdiay
ICONES PLEUROTHALUDINARUM
SYSTEMATICS OF BRACHIONIDIUM
Plate 17. BrachionidiumelegansLwr&i^
46
ICONES PLEUROTHALUDINARUM
, * -^-, prostrate, long-repent,
sheaths- roots solitanTfmU, , hctv^cea ramicauls. enclosed by 3 imbricating
nona bic 5 ZS’IS* ZZ “T '»’=■
scpais comiMe into aZme’ ZiZs sZ’lZ 3?Z”Z"i ” ““ '“‘S'
«™n long; pcdlinia 6. ^ ^
EV ADOR: Piw. of Morona-Santiago: epiphytic in
ToaoT*/ S‘g«g- alt- 2400 m. 13 Jan.
1989, C. Luer. J. Luer. P. Jesup, A. Jesup A Hirtz &
^Ort^aJJg<i6(Holotype:MO). Prov of Zamora-
CJinchipe: Nudo de SabaniUa, elfin forest at oass
^ve Valladolid, alt. 2800 m. 4 Feb iSrc £
®ast of Nudo de Cajanuma, alt
^ m. 31 Jan. 1989, J. E. Madsen 85750 (AAU,
Tliis fragile, little species is endemic in
eastern Ecuador at high altitudes. Plants
creep in the loose, wet humus at the base
of and on the trunks of large trees Sterile
specimens had been discovered several
l^imes before a usable flower was finally
ound. Common to similar species the
delicate flowers last but a short time.
Brachionidium ephemerum is distin¬
guished by the long-repent habit with
small, erect, elliptical leaves, and compar¬
atively short peduncles. The sepals and
petals are translucent, minutely ciliate and L____X.!\
Which
SYSTEMATICS OF BRACHIONIDIUM
Plate 18. BrachionidiumephemrtmlM^S lYMi
ICONES PLEUROTHALUDINARUM
Brachionidium escobarii Luer, Lindieyana 1:172,1986.
Ely.: Named in honor of Rodrigo Escobar R. erf Medellm, Cdombia, who discovered this species.
Plant very small, terrestrial, the rhizome repent, occasionally branching, up to ca. 8 cm long, 3-5
mm long with 2-3 pale, glabrous, loosely fitting sheaths between ramicauls; roots coarse, as thick as the
rhizome. Ramicauls 1.5-3 mm long, erect, mclosed by 2 similar sheaths. Leaf erect, thickly coriaceous,
suffused with purple, elliptical, acute, mucronate, 7-11 mm long including the petiole 0.5-1 mm long,
3.5-5 mm wide, the base cuneate into the petiole. Inflorescence a single, bright purple, non-resupinate
flower borne by a slender, purple peduncle 10-12 mm long, with 2 basal bracts, from near the apex of the
ramicaul; floral bract inflated, obhque, 2 mm long; pedicel thick, 1 mm long, with a protruding filament
2.5 mm long; ovary 2 mm long; sepals glabrous, the middle sepal ovate, acute, lightly acuminate, 5 mm
long, 3 mm wi^, the lateral sepals completely connate into an ovate lamina similar to the middle sepd,
transversely riiomboid or 3-lobed, 2.25 mm long, 3 mm wide, the lateral angles or lobes acute, the apex
obtuse, acuminate into an apiculum, with a pair of rounded calli, the disc with a central, flat callus;
column stout, 1 mm long, pollinia 6.
Plat^ 19. Brachionidium escobarii Luer
ICX)NES PLEUROTHALUDINARUM
SYSTEMATICS OF BRACHIONIDIUM
tLuer&Hirtz
ICONES PLEUROTHALUDINARUM
SYSTEMATICS OF BRACHIONIDIUM
Plate 21. BrachionidiumfolsomiTyKS&\Qt
ICONES PLEUROTHALUDINARUM
systemaucs of brachionidium
Plate 22. BrachionidiumfomcatumlMtxkVim
ICONES PLEUROTHALUDINARUM
SYSTEMATICS OF BRACHIONIDIUM
Plate 23. Brachionidiumfurfuraceum'L\J&T
ICONES PLEUROTHALUDINARUM
SYSTEMATICS OF BRACHIONIDIUM
Plate 24. Brachionidiim galeatumhvsT
ICONES PLEUROTHALLIDINARUM
SYSTEMATICS OF BRACHIONIDIUM
Plate 25. Brachionidium haberi Lnei
PLEUROTHALUDINARUM
SYSTEMATICS OF BRACHIONIDIUM
Plate 26. Brachionidium hirtzti Luer
5 PLEUROTHALUDINARUM
SYSTEMATICS OF BRACHIONIDIUM
Plate 27. Brachionidium irrq)eriale Luer & Escobar
SYSTEMATICS OF BRACHIONIDIUM
SYSTEMATICS OF BRACHIONIDIUM
Plate 29. Brachionidiumjulianii Camevaii & Ramirez
ICONES PLEUROTHALLIDINARUM
between ramicauls, enclosed by 3 loose, loosely fitting, n
Ramicauls subercet, 3-5 mm long, aiclosed by 2 T '
eUiptical, subacute, apiculate, 7-15 mm long with a petiole 0.5-2 mm long, 4-11 mm wide, 7- to 9-
!. 3-veined;
caudate synsepal 10-11 mm long including the tail 5_
apex obtuse, contracted into the slender tail, minutely bifid at the ti|
^ique, acute, acuminate. 9 mm long including the tail 3 mm long, 3 n
ty obcordate, 2 mm long. 3.25 mm wide, the lateral angles obtuse to roui.^™, u«.
shortly apiculate, lightly coicave above the middle, the margins microscopically e
low,flat.triangular.shortly pubescent caUus with asm ”--
2 pairs of which are connate at the ^x.
PANAMA: Prov. of Code: Alto del Calvario. north
of Aserradero H Cope, alt. 800 m, 5 Jan. 1980, R. L.
Dressier 5858 (Holotype: US; Isotype MOV same
^a. alt. 900 m, 19 Feb. 1985, C. Luer. J. Luer, R.
Dressier & K. Dressier 10631 (MO).
ECUADOR: Prov. rf Napo: epiphytic on the eastern
slope of Volcan Reventador, alt. 1850 m 8 Feb
1986. C. Luer. J. Luer. A. Hirtz. W Flores 17
Embree 11710 QAO).
This species was discovered growing
on a fallen log in central Panama in 1980
by Janet Kuhn with her daughter Helen
and Robert Dressier. Plants in flower were
found still growing there five years later.
Since the log is gradually deteriorating,
been moving individual
Brachionidium kuhniarum is recog¬
nized by the short, erect, leafy rhizome____
^ mLmdfSr »<* P^PJ®
one pair onLh side united"™w'*
em flank of VokS. RevS/ "S' <>"«»<« of Ecuador on die east-
SYSTEMATICS OF BRACHIONIDIUM
Plate 30. Brachionidium kuhniarumDKSsl&:
ICONES PLEUROTHALUDINARUM
Brachionidium lehmannii Luer,!
Ety.; Named in honor of F. C. I
ogisu who discovered this sp<
Plmta parva ihizomate erecto vel suberecto, pedunculo foliis ellipticis quinque-nervatis n
giore, florc parvo luteo, sepalo mediano synsepalo petalisque ciliatis longicaudatis, labello p
tnmsverse ovato apiculato maigine senato, disco concavo cum caUo basah pubescenti t’ransverso.
^iwccu rainicauls, enclosed by 2 imbricating, long-
the base. Ramicauls suberect, 2-4 mm long, enclosed
, elliptical, acute, apiculate. 8-15 mm long
ng me peoicet ana part of the ovary;
f 3 mm long; sepals minutely ciliate.
IB ovate, oblique, densely minutely'dliaieT W long, Tmm A^rideJ^v^^S
d into a slender tail 8 mm long; lip transversely ovate, 1.25 mm long, 2 mm
EOTADOR: Piov. of Pichincha?: Pil„.a -
1470^*'”'*”" (Holotype: K). C. Luer
by Consul Lehmann in the nineteenth
century. Unusual among Lehmann’s
collecnons, no altitude, date, or collection
number is given. The specimen was
^ong those purchased by Kew in 1906
from Lehmann's heirs in Colombia.
The most remarkable feature of the
IitUe, erect plant is the length of the
peduncles, more than twice as long as the
pl^t IS laU. The small flowers were noted
to be lemon yellow. The sepals and petals
ai^ minutely ciliate with filiform tails
^t twice as long as the blades. The tiny
Iip IS transverse, serrate, and concave with
a transverse callus across the base.
SYSTEMATICS OF BRACHIONIDIUM
Plate 31. BrachiomdiumlehmanniilMSX
SYSTEMATICS OF BRACHIONIDIUM
Plate 32. Brachionidiim longicaudatum /
ICONES PLEUROTHALUDINARUM
brevibus, pedunculo foliis suborbiculat
- 'suoerect, 5-7 mm long, enclosed by 2
Leaf suberect, conaceoos.descnbed as clear, bright green. 18-- ' ' '
long, the Wade broadly elliptical to suborbicdar. 6-8
ablate, dte rounded base contracted into the petio.... . a s,
ty a peduncle 10 nun Img. from near the apex of the ramicaul; floral
6 mm long; pedicel thick, 0.5 mm long -- ^ o
e, yeUow toward the api
acpaia oant reu-purpie, yell
rai^ate. long-caudate. 26
■ expanded, 4-veined, the filiform apices «rf the
WUU...6 luauicnt 7-8 mm long; ovary 3 i
--„ensely short-ciliate, the middle sepal ova
? including the slender tail 15 mm long, 7 n ’ "
imina, 26 mm long, 8.5 n
., —..imate, long-caudate, 22 ram Ion;
-r-arsely ovoid, 3.5 mm long, 6 mm wi(
apex «ath an obtuse apiculum, the margins with low. loni
iitral, flattened, minutely pubescent callus; column stout, 2
ECUADOR: Prov. of Loja: terrestrial among vegeta¬
tion 15 Km east rf Loja, alt. 2400 m, 27 Dec. 1947
R. Esmnosa & A. Espinosa 2292 (Holotype: NY).’
-Chinchioe: 14 Km Met i -i.
unauia-vnincnipe: 14 Km east of Loia al
21 May 1971,B. MacBryde 330 (AMES).
Although this species occurs in an area
that has been thoroughly collected by
many botanists for many years, this dis-
ttncuve species has apparendy been recol-
I^ted only once since it was first found by
Reinaldo and Arsenio Espinosa in 1947.
Brachionidium loxense is related to
large, robusl B. dalsmemii thal is found in
a nearby mountain-pass. Both species are
Stout and erect with short peduncles; rela-
tively large, long-tailed flowers; and
sirnilar, transversely ovate lips. Brachio-
niamm loxense is distinguished by the
smalle r habit: small, round leaves: and
sand petals.
SYSTEMATICS OF BRACHIONIDIUM
i
Plate 33. Brachionidiumloxensehuei
ICONES PLEUROTHALUDINARUM
SYSTEMATICS OF BRACHIONIDIUM
Plate 34.
ICONES PLEUROTHALUDINARUM
VENEZUELA: Edo. de TrujiUo: bet_
and La CuchiUa, alt. 2800 m, 3 Nov. 1951. /.
7507 (Holotype: Heib. J. Renz); between Santa Cruz
de Mora and El Molino. alt. 2800 m, 12 Nov. 1949. /
Renz 6110 (Herb. J. Renz); between Tovar and
Guanaque. alt. 2400 m, 18 July 1951. /. Renz 7252
Brachionidiunt meridense is a small
weak, creeping species found in the East¬
ern Cordillera of Colombia and western
Venezuela where it was first discovered by
Dr. Jany Renz. The flowers of the
Colombian collection are sUghtly smaller
than those of the type from Venezuela, and
the tail of the synsepal of the latter is
minutely bifid.
The small, more or less upright nar¬
rowly elliptical leaves are spaced along a
slender rhizome that creeps in loose,
mossy smaU. ihe translucent, pale brown Bowers with s.„.ua.,
sepals and petals are nearly as large as the leaves. The Up is broadly
SYSTEMATICS OF BRACHIONIDIUM
Plate 35. Brachionidiim meridense Gaay
SYSTEMATICS OF BRACHIONIDIUM
Plate 36. BrachionidiumminusculumhmT&DKSshi
ICONES PUEUROTHALUDINARUM
I
SYSTEMATICS OF BRACHIONIDIUM
85
Plate 37.
SYSTEMATICS OF BRACHIONIDIUM
iLuesr&Hirtz
ICONES PLEUROTHALLIDINARUM
r
SYSTEMAUCS OF BRACHIONIDIUM
89
?. Bracfttonidium parvifolium (Lindl.) LumH.
MLLIDINARUM
SYSTEMATICS OF BRACHIONIDIUM
ICONES PLEUROTHALUDINARUM
The aptly named B. parvum is widely distributed at relatively low altitudes of
both the Greater and Lesser Antilles, Venezuela, and Ecuador. Four different col¬
lections were cited by Cogniaux in the original description. The first is a collection
by Harris of B. syme-morrisii from Mt. Moses, Jamaica; the second is a sterile
collection by Duss {Duss 3730) from Guadeloupe (not Martinique); and the third
and fourth collections are by Broadway from Grenada. Congiaux’s description
applies to the last two collections, and not to either the Jamaican collection, nor to
the sterile collection from Guadeloupe.
The Grenadan collection of W. E. Broadway 268 at Kew has been designated as
the lectotype of B. parvum instead of the Guadeloupean collection of Dms 3730 at
New York indicated by Garay, because Cogniaux’s description applies to the
Broadway collection, and because the Duss coUection is without flowers. Because
Duss 4149 from Guadeloupe, the type of B. dussii, has been lost at B, a recent
collection from Guadeloupe has been designated as a neotype. These small-flow¬
ered plants from Guadeloupe are indistinguishable from B. parvum.
Garay (1956) reduced B. longicaudatum to the synonymy of B. parvum, appar-
entiy basing his conclusion on vegetative similarities of B. longicaudatum from the
mountains of Amazonian Venezuela and the sterile collection from Guadeloupe by
Duss. Brachionidium longicaudatum is distinguished by the small but stout, erect
habit; tails of the sepals and petals longer than the blades; and an acutely deflexed
lip, the disc of which is mostly covered by a broad, pubescent callus. Garay (1956)
published drawings of B. longicaudatum under the name of F. parvum.
Brachionidium parvum is distinguished by the very small, erect habit; either
purple or yellow sepals and petals less than one centimeter long including the
acuminate, tail-like apex; and a minute, transversely trapeziform, obtuse lip with a
pubescent callus. It is very similar to a small form of B. sherringii, and was consid¬
ered a variety of that species by Stehld. Forms intermediate between B. parvum and
B. sherringii occur.
The recent discovery of essentially identical plants in western Ecuador would
seem to indicate that this species is more widely distributed in South America than
” referring to the sh^ of the lip.
n long between ramicaals, enclosed by'2-3 ii
near the base. R^icanls suberect. 5-6 mm long, enclosed by 2 mucronate sheaths. Leaf erect tc
Mte, apiculate, 12-15 mm long including the petiole 1-2 mm long. 5-7
•art the ovary; pedicel 0.5 mm Icmg, with a
SYSTEMATICS OF BRACHIONIDIUM
Plate 41. Brachionidium peltarion hucT
ICONES PLEUROTHALLIDINARUM
PANAMA: Piov. Code: Cerro Gaital, mesa north of
El Valle de Anton, near western peak, alt. ca. 900 m,
14 Mar. 1981, R. L. Dressier & E. Hagsater 6005
(Holotype: MO),C. Luerillustr. 14556.
This little species is known only from a
single collection by Dressier in the moun¬
tains of central Panama. It is characterized
by the small, suberect habit; a small,
yellow-green flower, tails of eciliate sepals
and petals only 1 mm long; and an acutely
deflexed, shield-shaped, obtuse lip that is
essentially without a callus. The deflexed,
basal portion of the lip is microscopically
pubescent, reminiscent of the more coarse
pubescence on the deflexed portion of the
lip of B. longicaudatum.
COLOMBIA: Dept, of Norte de Santandcn Munic. of
Toledo, Alto de Mefue, epiphytic in cloud forest, alt
2600 m. 25 May 1982. C. Luer, R. Escobar & D
PoitiUo 7968 fS£L>.
This long-repent species is found
growing in damp, mossy humus of cloud
forests in the Eastern Cordillera of
Colombia and adjacent western Venezuela
where it was first discovered by Dr. Jany
Renz.
The erect, elliptical leaves are pro¬
duced at intervals of 1.5-2 centimeters
along the rhizome. The bright purple.
SYSTEMATICS OF BRACfflONIDIUM
SYSTEMAUCS OF BRACfflONIDIUM
Plate 43. Brachionidiumpiunt^^^-^^
ICONES PLEUROTHALUDINARUM
Brachionidium polypodium Luer, sp. nov.
iW to a petiole 1-2 mm long. InfloresceL____._
t, filifonn peduncle 14-16 mm long, with a bract below the middle
1; floral bract infundibular, mucronate, 3 mm long,
og, with a filament 3 mm long; ovary 1.5 mm long;
long including the Hlifotm i«l 6 nin loog/s romae to”
including the fo*ed, liUfom. toil 5
1, shordy cUiate, ovate, oblique,
1 long, 2.75 n
SYSTEMATICS OF BRACHIONIDIUM
44. BrachionidiumpolypodiumLaer
SYSTEMATICS OF BRACHIONIDIUM
101
45. BrachionidiumportUlae Luer
SYSTEMATICS OF BRACHIONIDIUM
103
Plate 46. BrachionitUumpteroglossumUm
systemaucs of brachionidium
105
Plate 47. Brachionidium puraceenseLusr
MXIDINARUM
f
SYSTEMATICS OF BRACHIONIDIUM
107
ICONES PLEUROTHALUDINARUM
teo pro planta grande, sepalis
; peltato obtuso maigine den-
m long, enclosed 1^ 2 re
15-30 re
the petiole. ]
ig, with a bract below the middle; floral bract long-mucronate, 3.5-4 mm
part of the ovary; pedicel 1-1.5 mm long, with a protmding filament 3.5
sepal elliptical, 10 mm
n wide, 3-veined, the apex
into a sknS uflTsTc^
xite, acuminate, the blade 9
27 mm Iraig; lip transversely pelute, 2
slender tail 3-3.5 c
ig, 6.5 n
3-3.5 cm long, the tip bifid for about 7-8 mm; petals minutely ciliate, ovate,
tinate, the blade 9 mm long, 4.5 mm wide, 3-veined, contracted into a slender tail 25-
im long, 4 mm wide, with subacute lateral angles, the apex
r
SYSTEMATICS OF BRACfflONIDIUM
Plate 49. BracHomdiumrenziil^
no
ICONES PLEUROTHALUDINARUM
Brachionidiuni restrepioides (Hoehne) Pabst, Orch. Bras. 166,1975.
Bas.: Yolmida restrepioides Hoehne, Arq. Mus. Nac. Rio de Janeiro 22:72,1919.
Ety.: Named for a fancied resemblance to the genus Restrepia H.B.K.
Syn.: Brachionidium dungsu?!b&t, Bradea 1:268,1972.
BRAZIL: Estado dc Sao Paulo: Esta^So Biologica do
Alto de Seira Sfio Paulo, Dec. 1920, F. C. Hoehne
2152 (Holotype:? ; Isotype: MO). C. Luer illustr.
14372;F ' •
W. J. Keay K-67 (K). Estado do Rio de Janeiro, near
Nova Friburgo, alt. 1300 m. 7 May 1972, F. Dunes
SJi. (HB, type of B. dungsii).
This species is apparently the only
representative of the genus in southern
Brazil where it was first discovered by I>r.
Hoehne. He described it in a new genus,
Yolanda, because it had eight pollinia. At
that time, the number of pollinia in a
species of Brachionidium had been report¬
ed only twice, each species bearing six.
Therefore, it was assumed that a Brachio- ____
nidmm could have only six pollinia. However, the type-species of the g=..us,
^ -J**" A “corapanying Ulusiration
r!Z “ «°'ypa reconstituted in
concentrated ammonium hydroxide.
™<repio,Vfex is inaccurate. TTie
™ apj^ce of the hp is piacticaUy exactly as drawn for the type-descripuon of
B. dungsii, a purple-colored form discovered in 1972.
r
SYSTEMATICS OF BRACHIONIDIUM
111
Plate 50. BrachionicUum i
, restrepioides (Hoelme) Pabst
systemaucs of braoiionidium
Plate 51. Brachionidiumrugosum\
iLuer&Hirtz
114
ICONES PLEUROTHALUDINARUM
mm wide, 3-veined, the tail filiform, 13 cm long; lateral sepals connate into an ovate, acute, acuminate,
synsepal. concave with the middle third convex, the blade 11 mm long, 6.5 mm wide expanded, 4-
veined, the tail filiform, 14 mm long, the tips free for 2 mm; petals deep wine red, reflexed, minutely
lip purple, subquadrate-trilobed, acutely deflexed near the middle, 1.5 mm long, 2.5 mm wide, the lateral
angles subacute, erect, the middle lobe broadly obtuse, concave centrally below the callus, the disc with a
rounded, curved, pubescent callus, protruding above the cavity of the blade; column stout, 1 nun long,
pollinia 6.
"PERU: Bci Cochobamba,” 1907, Dr. Filomeno sm. (Hdotype destroyed at B).
This small species, collected from an unknown locality by an obscure collector,
IS known only from the type-description. Tlie holotype was destroyed at Berlin, and
no sub^quent collection is known to have been made. The above description was
made from Schlechter’s original publication.
Apparently B. serratum is a small, repent species with ovate leaves about 15
miUimeters long. The flower is very small and purple without sepaline tails, the
nvnte sepals being only eight millimeters long. The equally long petals are de¬
scribed as “scharf gesagten”
(sh^ly saw-toothed). The lip, described as thickly
rhomboid and obtuse, ^parentty is similar to forms commonly
SYSTEMATICS OF BRACfflONIDIUM
115
Plate 52.
PLEUROTHALUDINARUM
SYSTEMATICS OF BRACfflONffilUM
Plate 53. BrachiomdiumshernnguKoUe
118
ICONES PLEUROTHALUDINARUM
This species, frequent and widely distributed near the tops of the mountains of
the islands in the Lesser Antilles, was first collected on Grenada in 1890 by R. V.
Sherring for the West India Exploration Committee. The accompanying iUustration
was made from a dried flower from the type-collection reconstituted in concentrated
Brachionidium sherringii is variable in size vegetatively, ranging from small to
relatively large. The habit is commonly long-repent, but more or less erect forms
are not infrequent, presumably depending upon the amount of exposure and light.
The long-tailed flowers vary in size from relatively small to medium in size for the
genus. The color varies from yellow-green to dark red-purple. The sepals are not
ciliale, but the petals are microscopically ciliate. The transverse lip with a central
callus is basically similar to the lips of many other species: transversely subovate,
broadly obtuse, minutely apiculate, with longitudinally callous margins, and with an
ovoid, shortly pubescent, central callus. The tips of the lateral angles of the lip are
minutely uncinate in the type-collection, as well as in others, but these angles are
simply obtuse on the lips of some other collections.
Small forms grade into B. parvum. Although the sepals of the larger B. sherrin¬
gii are eciliate, and the sepals of the small P. parvum are minutely ciliate, excep¬
tions occur. The two taxa could be considered merely vegetative and floral forms
of one variable species.
ECUADOR: Without locality, “Quito,” W. Jameson
s.n. (Holotype: AMES). Prov. of Morona-Santiago:
The type-specimen of Brachionidium
simplex was collected in the nineteenth
century by Professor Jameson who lived in
Quito, Ecuador. No collection data are
known, and how the specimen arrived in
the personal herbarium of Garay is also
unknown. The original description was
drawn from a reconstituted, dried flower.
SYSTEMATICS OF BRACHIONIDIUM
119
1 Brachionidiim Simplex Gat2iy
HALUDINARUM
r
SYSTEMATICS OF BRACHIONIDIUM 121
Plate 55.
122
ICONES PLEUROTHALLIDINARUM
BrachioDidiuin syme-morrisii Luer, sp. nov.
Ely.: Named in honor of Geojge Syme and Sir £>aniel Morris, who first collected the species.
JAMAICA: Mt. Moses, alt. 3000-5000 ft., Nov.
1881. G. Syme & D. Morris 2361 (Holotype:K,
Isotype: W), C. LueriUustr. 14459; Mt. Moses, alt.
1000 m, W. Harris 7802 (Herb. Hot. Dept.); Mt.
Moses, 1908. received from W. Harris s.n. (N^; St.
Andrew. Mt. Horeb. Port Royal Mts. G. R. Proctor
This species was first collected by
George Syme and Sir Daniel Morris on
Mt. Moses in Jamaica in 1881, and again
by Syme on 20 Sept. 1882 is appended to
the type-specimen at Kew. Reichenbach’s
sketch present in his herbarium at W
(Vienna) has the intended name Brachio-
nidium morrisii. This species is endemic
jg (jjg Qjjjy species of the
-.lud Kiiuwii to occur on tne island.
In his original description of B. parvum
Cogniaux cites four separate collections as ---^^
^ collection of this species by Harris from Mt. Moses in
n Jamaican species is smaller in
(mm r “an^iy long-repenl B. sherringii. One of the two specimens cited
OrrhidK '^Jj^ Orchidaceae Antillanae and Fawcett and Rendle in the
Evav author si i*!® ^yme and Morris’ collection as B. sherringii.
syme-n^rrisU is disL-
and a distinctive cl ^ habit; minutely ciliate sepals and petals;
, c e ID with ftrprt 1 ^ broad, concave
SYSTEMATICS OF BRACfflONIDIUM
123
ICONES PLEUROTHALUDINARUM
SYSTEMATICS OF BRACHIONIDIUM
125
ICONES PLEUROTHALUDINARUM
Biiil
SYSTEMATICS OF BRACfflONIDIUM
127
Plate 58.
128
ICX)NES PLEUROTHALUDINARUM
in 0. Rangel & J. Aguirre 12223 (COL. NY). Dept, of Santander: paramo east of Bucaramanga
rd BerUn 5t 2950 m. 7 May 1984, C. Luer, J. Luer. R. Escobar & E. Valencia 10170 (MO). Dept,
a Manizalcs and Fresno, alt. 3200 m, 6 Apr. 1956, /. Renz 8619 (Herb. J. Renz).
ECUADCMt: Prov. of Azuay: moist, forested slojws bordering Ri'o Collay south of El Pan, alL 2650-3290
m. 6 July 1943. J. A. Steyermark 53360 (F), specimen sterile.
BOLIVIA: without locality. 1846. Bridges s.n. (Holotype: K). Dept, of Cochabamba: Prov. of Chapare,
tetiestrial in forest northeast of Codiabamba, alt. 1900 m, 29 Jan. 1984, C. Luer, J. Luer & R. Vdsquez
9476 (SEL); same area, 18 Jan. 1988, C. Luer, J. Luer, R. Vdsquez, T. *S D. Mulder & A. Vogel 12867
(MO).
The epithet tuberculatum is misleading. There is indeed a thick callus in the
center of the lip, and to Lindley it seemed tuberculate in a rehydrated or poorly
preserved flower. The locality and vegetative appearance of the plant and the
sketch of the flower leave no doubt that it is a well-known, frequent and somewhat
variable species found in all three cordilleras of Colombia, adjacent Venezuela,
Ecuador and Bolivia. Specimens of Schlim’s original collection are present in
several Eurqiean herbaria.
This species was first collected in Bolivia by Bridges a century and a half ago,
and it had not been collected there again until 1984. Lindley described it as B.
concolor simultaneously with B. tuberculatum, but the later is chosen for the name
because it is widely known in a wide distribution.
A specimen from western Venezuela was redescribed by Garay as B.floribun-
dum, and he applied the epithet tuberculatum to B. tetrapetalum (Garay, 1956),
which is also relatively common but restricted to the high paramo and subparamo
Brachionidium tuberculatum is characterized by a stout rhizome that usually
grows more or less erect terrestrially in the leafy humus of wet cloud forests. The
rhizome is enclosed by a series of mucronate sheaths that are more or less tuber¬
culate or fimbriate on their lower margins. Old plants seem to cease growing in
height and leaves become lost or degenerate so that herbarium specimens vary in
appearance. Specimens of young, actively growing plants retain more and fresher
leaves.
The solitary, purple flower with acuminate sepals and petals is held just above a
leaf. The petais are microscqjically but densely ciliate. The broadly cordate lip is
furnished with an elevated but flat, minutely pubescent callus channeled on the
front surface into a glenion. Freshly pickled flowers from several, distant localities
have been examined. Plate 58. is an illustration of a collection from Colombia, and
Plate 59. is an illustration of a collection from Bolivia.
SYSTEMATICS OF BRACHIONIDIUM
129
Plate 59. Brachionidium tuberudatumhindl
e sheaths. Leaf erect or suberect, coriaceous, elliptical, acute, 15-23 mm
long including a petiole 2-4 nun long, 6-9 mm wide, 5- to 7-veined. Inflorescence a solitary, non-resu-
pinate flower home by a slender, erect peduncle 10-20 mm long, with a mucronate bract near the middle,
from near the apex of the ramicaul; floral bract inflated, acute, mucronate, 3-4 mm long, enclosing the
r, pedicel stout, 0.5 mm long, with a fdament 4 mm long; ovary 2 nun long; sepals
red-purple stripes along the veins, shortly ciliate, the middle sepal ovate, acute,
m long, 6 mm wide, 3-veined, the lateral sepals completely connate into an ovate,
n long, 8.5 mm wide, 4-veined; petals ovate, obhque, acute,
2, 3-veined; lip yellow-orange, transversely obovoid, 2 mm
long, 3.5 nun wide, the lateral angles acute, the apex obtuse, shortly apiculate, the disc thickened above
Ae CTtire margins, with a rounded, elevated but flat, pubescent callus above the base, the base thin,
BOLIVIA: Dept of Cochabamba: Chapare, northeast
of Cochabamba toward Villa Tunari, terrestrial in
deep humus in wet forest, alt 2800 m, 14 Nov. 1992,
C. Luer, J. Luer, R. Vdsquez, D. Ric & W. Teague
i64id(Holotype:MO).
This distinctive, showy species is
known from a single discovery by Jane
Luer. Several plants were growing at the
mossy base of a large, ancient tree on the
north slope of central Bolivia where it is
presumably endemic. See Frontispiece.
At that time of its discovery, this orchid-
rich forest was in the process of being
destroyed for fire-wood by local people.
Vegetatively this species appears
similar to the widely distributed B. tuber¬
culatum. The colors of the flowers are
unique in the genus: white with prominently red-striped sepals and petals and a
utterscolch-colored lip. The sepals are slightly acuminate, not caudate; the petals
are acute and shortly caudate. The 1^ is transverse with acute, lateral angles.
systemaucs of brachionidium
131
132
ICONES PLEUROTHALLIDINARUM
Brachioiiidiuiii valerioi Ames & Schweinf., Sched. Orchid. 10: 53,1930.
Ety.: Named in honor of Juvenal Valerio, co-discoverer (rf this species.
Plant smaU but forming large mats, epiphytic, the rhizome slender, long-repent, occasionally
branching, up to 20 cm long, 4-12 mm long between ramicauls, with 3 imbricating, mucronate, tubular
closed by 2 similar sheaths. Leaf erect to suberect, coriaceous, elliptical, acute, apiculate, 15-20 mm long
including the pctide 1-2 mm long. 5-7 nun wide, 7-veined, the base cuneate into the petiole. Inflores¬
cence a single, non-resupinate flower borne by a slender peduncle 15-18 mm long, with a bract near the
middle, from near the apex of the ramicaul; floral bract infundibular, acuminate, mucronate, 3 mm long,
enclosing the pedicel and ovary; pedicel thick, 0.5 mm long, with a protruding filament 3 mm long;
ovary stout. 1.5 nun long; sepals translucait pale green, the middle sepal elhptical, the blade 5 nun long,
3 mm wide, 3-veined, the apex acute, long-acuminate into a filiform tail ca. 15 cm long, the lateral sepals
connate into a ovate, deeply ccuicave synsepal, 5 mm long, 5 mm wide expanded, 4-veined, the apex
acute, acuminate into a Uil ca. 15 mm long, bifid above the middle; petals translucent pale green,
minutely ciliate, ovate, obUque, the blade 5 mm long, 3 mm wide, 3-veined, the apex acute, acuminate
into a filifonn tail ca. 15 mm long; lip transversely subtrapeziform, 2 mm long, 3 mm wide, the lateral
angles acute, the apex broadly rounded, prominently apiculate, the margins coarsely denticulate-erose,
with a submarginal crested caDus, with minute crystalline substance within the lip, the disc with a cen¬
tral, flat, ovoid, minutely pubescent callus shallowly deft anteriorly; column stout, 0.5 mm long, 1 mm
broad; poUinia 6.
COSTA RICA: Prov. of Heredia: Cerros de Zurqui
northeast of San Isidro, alt. 2000-2400 m, 8 Mar.
1926. P. C. Standley & J. Valerio 50759 (Holotype:
AMES; Isotype: US).C. Luerillustr. 14465; Cerro
Zurqui, alt. 2000 m. 13 Mar. 1979, C. Todzia 563
(CR>, Yerba Buena, northeast of San Isidro, alt. 2000
m, 22-28 Feb. 1926. P. C. Standley & J. Valerio
50136 (AMES, US); Cerro de las Caricias north of
San Isidro, 11 Mar. 1926, P. C. Standley & J. Valerio
52366.52479 (AMES, US).
This species is apparently confined
locally in the province of Heredia in cen¬
tral C(Kta Rica where it has been found on
few occasions. It is a weak, long-repent
species with a slender rhizome and little,
elliptical leaves. Large, loose mats may
accumulate. The sepals and petals of the
small flowers are long-caudate. The very
distinctive lip is coarsely fringed with an
additional fringed callus lying behind the
margin of the Up.
The illustration pubUshed by Garay (1956) for B. valerioi is B.folsomii which
was undescnbed at that time. The illustration was copied from a drawing made by
Endres in Costa Rica in the nineteenth century. Garay did not include an illustra¬
tion of fi. vo/erioL
SYSTEMATICS OF BRACHIONIDIUM
133
1. BrachionUtium valerioi Ames & Schwemf.
134
ICONES PLEUROTHALLIDINARUM
Bracbionidium vasquezii Luer, Phytologia 55: 177,1984.
Ely.: Named in honor of Roberto Vasqaez Chavez of Cochabamba, Bolivia, illustrator and co-author
of the Orchids of Bolivia, who discovered this species.
BOLIVIA: Dept, of Cochabamba: Prov. of Chapare,
Balconcillo, terrestrial in moss along the trail, alt.
3250 m. 7 Dec. 1980, R. Vdsquez 477 (Holotype:
SEL; Isotype: Herb. Vasquezianum); same area, 17
Jan. 1984, C. Luer, J. Luer & R. Vdsquez 9356
(SEL); 18 Nov. 1984, C. Luer. J. Luer & R. Vdsquez
10493 (MO). Dept, of U Paz: Prov. of Nor Yungas,
terrestrial northwest of Chuspipata, alt. 3100 m, 26
Aug. 1983. J. C. Solomon 10663 (LPB, MO).
Brachionidium vasquezii was first
discovered by Roberto Vdsquez in 1980.
It grows terrestrially at high altitudes over
30(X) meters above sea level on the north¬
ern slopes of the Andes of Bolivia. Erect
rhizomes stand in wet, mossy habitats with
the showy, purple-striped flower held high
by an exceedingly long peduncle, which is
unknown in any other member of the
genus.
The sepals and petals are broadly ovate, obtuse and acuminate into slender tails
considerably shorter than the blades. Only the sepals are minutely cUiate. The lip
is transversely ovate and concave with a flattened margin. The relatively small
callus is elevated with an anterior glenimi.
systemaucs of brachionidium
135
PLEUROTHALLIDINARUM
SYSTEMATICS OF BRACHIONIDIUM
137
ICONES PLEUROTHALLIDINARUM
REFERENCES
COGNIAUX, A., 1909. Orchidaceae Antillanae in Urban, Symbolae Antillanae 6:450-452.
Dressler, R. L., 1981. The orchids: natural history and classification. Harvard University Press,
Cambridge.
-^ 1982. Dos Brachionidium nuevas en Panama. Orquideologra 15:152-164.
Fournet, J., 1978. Flore iUustre des phanerogames de Guadeloupe et de Martinique. INRA Paris.
Garay, L. A., 1956. Studies in American orchids, 11. the genus Brachionidium Lindl. Canad. J. Hot. 34:
721-743.
Garay, L A., 1974. Orchidaceae in Howard, Flora the Lesser Antilles, 111-113.
Linduby, j., 1859. Fdia Orchidacea Brachionidium. J. Matthews, Covent Garden, London 1.
Luer, C. a., 1986. IccMies Pleurothallidinarum I. Systematics of the Pleurothallidinae. Monogr. Syst.
Bot Missouri Bot. Gard. 15:20-21.
Sastre, C., 1991. Retrouvailles avec Brachionidium dussii Cogniaux sur La Soufriere de Guadeloupe.
L’Orchidonhile 00-
SYSTEMATICS OF BRACHIONIDIUM
139
INDEX OF SCIENTIFIC NAMES
,5.32,74,124
1, 22, Plate 6.
i 1,6,24, Hate 7., 78
re 1.8,26, Hate 8.. 66
um 7,28, Plate 9.
or 10.126,128
5,30, Plate 10.
6,32, Hate 11.. 76
1,6,34, Plate 12.
;13.
10,90,92
arense 9,14,42, Plate 16., 46,104
IS 5,16.44, Plate 17.. 56
lenim 1,9,46, Plate 18.
8.48, Plate 19.
), 50. Plate 20.
2,10.126.128
18.38,52, Hate 21.. 60,108,
1.6,54, Plate 22.. 88
i. Hate 23.
,5.58. Plate 24., 112
haberi 1.6,60, Plate 25.
hirtzii 6,9.62, Hate 26.
imperiale 1.6.64, Plate 27.. 88
j 1.8,26,66. Plate 28.
8.68, Plate 29.. 108
2,7,70, Plate 30.
1,8,72. Plate 31.
7,74. Hate 32.. 92.94
1,6,76, Hate 33.
6.24.78. Plate 34.
I, Hate 35.
8,82. Hate 36., 106
9.82, Plate 37.
parvifdium 1,2,4.6.10,14,44.54,64.86,
Plate 39.
parvum 7.28,74.90. Hate 40.. 92.118,122
pdtarion 1.7,92, Hate 41.
phalangifenm 9,62.94. Plate 42.
piuntzae 1.6. %, Hate 43.
polypodium 1,9.98, Hate 44., 136
poitUlae 7,100, Hale 45.
pteroglossum 7.102, Hate 46.. 120
puraceense 1.9.104. Plate 47.
pusinum 8.20.82.106. Hate 48.
renzii 1,8,108, Plate 49.
restrepioides 4.7,84,110, Hate 50.
ragosum 6,58,112. Plate 51.
1.6.24,78.112, Hate 52.
c 53.. 122
1,8.92,122. Plate 56.
S,7.102.120,124. Hate 57.. 128
2,3,5,126, Plates 58., 59.. 128,
kcgellii 2
parvifolia 1,4,86
SteUs 3
YoUnda 2,4.110
restrepioidcs 4,110
140
ICONES PLEUROTHALUDINARUM
ADDENDA TO DRESSLERELLA, PLATYSTELE AND PORROGLOSSUM
V spedes of Dresslerella Luer, Platystele Schltr. and Porro-
. The plates are numbered as continuations of their
Dresslerella archilae Luer & Behar, sp. nov.
Platystele bovilinguis Luer, sp. nov.
1 archilae Luer & Behar, sp. nov.
ed in honor of Fredy Archila of Coban, Guatemala, who discovered this species,
es ^ns Dresslerellae Luer jdanta parva, folio ovato longipubescenti, ovario sepalisque
sepalo dorsali ovato acuto liberi, synsepalo saccato, petalis glabris oblique
This litUe species is distinguished by
the ovale, long-pubescent leaves; a small,
purplish, long-pubescent flower with the
dorsal sepal free from a deeply saccate
synsepal; and by short peduncles. The
petals are obliquely obovate. The lip is
oblong, vemicose above the middle, and
with a pair of smaU, uncinate lobes below
the middle. The four equal-sized pollinia
^ from tte usual unequal paiis seen in
Other members Of the genus.
SYSTEMATICS OF DRESSLERELLA
141
9. Dresslerella archilae Luer & Behar
MUDINARUM
SYSTEMATICS OF PLATYSTELE
143
Plate 90. PlatystelebovilinguisLm
cal,obtuse, 1.2-1.5cm wide,^^e^eSow^mo^Ae^^-
vat^ sHghUy cleave, 6 mm lo^! W wi^/
SYSTEMATICS OF PORROGLOSSUM
145
Plate 36. Porroglossumjosei huci
146
ICONES PLEUROTHALUDINARUM
A RE-EVALUATION OF THE PLEUROTHALLID
SUBGENERA SATRYIA AND SILENIA (ORCHIDACEAE)
In Systematics of Myoxanthus (Luer 1992) two subgenera of Myoxanthus Poepp.
& Endl. were proposed for seven species that varied significanUy from the perimit-
ers that defined the genus. The species of these taxa have greater affinities with
PleurothalUs R.Bt. subgenus Acianthera (Scheidw.) Luer (Luer 1986) than they
have with Myoxanthus. Their transfer to secUons of sugbenus Acianthera restores
the integrity of Myoxanthus.
Pleurothallis subgen. Acianthera sect Satyria Luer (staL nov.)
Pleurothallis balaemceps Luer & Dressier, Orquideologia 16:47.1986.
Syn.: Myoxanthus balaeniceps (Luer & Dressier) Luer, Monogr. Syst Bot. 44: 94,1992.
Pleurothallis lapp^ormis Heller & LO.Williams, Fieldiana Bot. 31: 42 1964
Syn : Myoxanthus tappifomus (HeUer & L.O. Williams) Luer, Monogr. Syst. Bot 44: 96, 1992.
P/enrette/ttspan Luer, SdbyanaS: 360,1976.
Syn.: Myoxanthus pan (Luer) Luer, Monogr. Syst Bot 44:98,1992.
Pleurothallis stemei Luer, Phytdogia44:171,1979.
Syn.: Myoxanthus stonei (Luer) Luer, Monogr. Syst Bot 44: 98.1992.
,BOTplandia3:75.1855.
/.Gem. PL 2:667,1891.
.1. „• -. T-’ --Veg. Beih. 19: 10
Syn.: pastensis Luct, Sdbyana 3:156,1976.
“ ) Luer, Monogr. Syst Bot 44: 100, 1992.
literature aiED
of Pleurothallis. Monogr. Syst. Bot. 20:12.
uer, CA., 1992. Systematics of Myoxanthus. Monogr. Syst Bot 44: 5-6,94-108.
-A 1753 8840