TAXONOMY OF BARKERIA (ORCHIDACEAE)

L. B. THIEN AND R. L. DRESSLER

Thien, L. B. (Dept. of Botany, University of Wisconsin, Madison) & R. L.

Dressler (Smithsonian Tropical Research Institute, Balboa, Canal Zone).
Taxonomy of Barkeria (Orchidaceae). Brittonia 22: 289-302. 1970.--The genus
Barkeria (Orchidaceae) consists of four species and eight subspecies (the treat-
ment includes a key) of epiphytic plants endemic to Mexico and Central America.
Barkeria is separated from the genus Epidendrum on the structure of the rostellum
and column in the flower, plus shape of the pseudobulbs. The authors suggest that
Barkeria is most closely related to Caularthron based on the general shape of the
flowers, widely spreading fleshy column wings, and structure of the rostellum.

INTRODUCTION
From the latterpart of the nineteenth century to the present, there has been disagree-
ment regarding the taxonomic relationship of Barkeria to Epidendrum. The decision
to maintain Barkeria as a distinct genus within the Epidendrum alliance has depended
upon whether Epidendrum is considered to comprise an artificial or natural assemblage
of plants (Dressier, 1961). I n this paper Barkeria is separated from Epidendrum
primarily on the structure of the rostellum and column in the flower, and shape of
the pseudobulbs (Table I; Dressier, 1961 ).
The genus Barkeria was first described by Knowles & Westcott in 1838, based upon
Barkeria elegans, in honor of George Barker of Springfield, England, who had im-
ported the plant from Mexico. The showy and lasting qualities of the flowers created
great interest and demand for the plants among growers and collectors. The remaining
members of the genus were described in the middle and latter parts of the nineteenth
century.
The genus Dimerandra was included in Epidendrum sect. Barkeria b y Ames, Hub-
bard, & Schweinfurth (1936), but Dimerandra is very different in habit and in-
florescence, in the form of the lip and column, and in the rostellum. We suggest that
Barkeria is most closely allied to Caularthron. This relationship is suggested by the
general shape of the flowers, widely spreading fleshy column wings, and the structure
of the rostellum (Table I ) .
TAXONOMY
Barkeria: Knowles & Westcott, Floral Cabinet 2: 7. 1838.
Epiphytic plants occurring on rocks, trees, or shrubs, small to 12 cm or more tall,
erect, sometimes caespitose. Stems forming slender pseudobulbs, bearing leaf sheaths.
Leaves 2-7, linear-lanceolate to broadly ovate, concentrated on the upper part of the
stem or arranged along the stem. Inflorescence a terminal raceme, sometimes branched,
with closely clasping scarious bracts. Pedicel 1.5-4 cm long. Flowers I-6 cm in diam-
eter, perianth parts reflexed or projected forward. Sepals and petals similar, or the
petals wider than the sepals. Lip free or adnate to the column, up to half the length
of the column, simple, callus absent or consisting of three thickened keels extending
beyond the apex of the column, sometimes verrucose and extending along the lateral
veins of the lip. Column 2-16 m m long, with widely spreading membranaceous or
fleshy wings, appresscd against the lip or divergent. Rostellum tongue-shaped, without
a viscidium. Anther terminal, 2-celled, with four equal, waxy pollinia, in one series,
sometimes united in loose pairs by the caudicles.
BRITTONIA 2 2 : 289-302. October-December, 1970.

289
290 BRITTONIA [VOL. 22

K e y to the Species of B a r k e r i a
1. Column 2---4 mm long; callus of the lip with keels, and lateral veins papillose-verrucose.
2. Ovary without a conspicuous visible nectary; flowers cream, sometimes with reddish
veins, la. Barkeria chinensis ssp. chinensis.
2. Ovary with a conspicuous visible nectary; flowers reddish or pale lilac.
3. Nectary goiter-like, 2 mm lo.ng; flowers reddish; only in Mexico from Juchit~in
(Oaxaca) to the Rio Balsas on the Pacific slope, lc. Barkeria chinensis ssp. naevosa.
3. Nectary sac-like, 3--4 mm long; flowers lilac; only in Mexico in the states of Colima,
Jalisco, Nayarit, and Sinaloa. lb. Barkeria chinensis ssp. palmeri.
1. Column more than 6 mm long; callus of the lip absent or consisting of three thickened
keels, the lateral veins of the lip not thickened nor verrucose.
4. Column entirely free, diverging from the lip. 6. Barkeria melanocaulon.
4. Column basally adnate to lip and appressed against it.
5. Column adnate to lip for about one-half the length of the column; column wings
fleshy. 3. Barkeria skinneri.
5. Column adnate to lip up to one-third of the length of the column; column wings
membranaceous or slightly fleshy.
6. Callus absent, or if present, not extending beyond the column.
7. Sepals and petals subequal, clawed; callus absent; the column 1--1.4 cm long.
4. Barkeria barkeriola.
7. Sepals longer than the petals; callus consisting of 3--5 thickened keels not
extending beyond the column; column 1.5--2 cm long. 5. Barkeria elegans.
6. Callus present, consisting of three thickened keels at least three-fourths the length
of the lip.
8. Base of tip enfolding column; upper one-third of lip dark lilac; Costa Rica.
2a. Barkeria lindleyana ssp. lindleyana.
8. Base of lip not enfolding column; upper one-third of lip spotted; Mexico and
Guatemala.
9. Column 1.5--1.8 cm long; lamina of lip oval to elliptic (apex may be
mucronate); flowers pale lilac with dark purple spots on lip; Chiapas,
Mexico, into Guatemala. 2b. Barkeria lindleyana ssp. spectabilis.
9. Column 1--1.4 cm long; lamina of lip not oval to elliptic; Mexico (Oaxaca,
Puebla, Guerrero, Michoac~n, and Veracruz).
10. Flowers red-magenta; Mexico, in the state of Michoac~in.
2d. Barkeria lindleyana ssp. cydotetla.
10. Flowers pale lilac; Mexico, i.n the states of Puebla, Guerrero, Oaxaca, and
Veracruz. 2c. Barkeria lindleyana ssp. vanneriana.

la. BARKERIA CHINENSIS ( L i n d l e y ) T h i e n ssp. CHINENSIS

Barkeria chinensis (Lindley) Thien i~n Dressier, Taxon 15: 241. 1966.
Broughtonia chinensis Lindley in Hooker's London Jour. 1: 492. 1842. TYPE: "Hong Kong,"
[actually Guatemala, fide Reichenb. f.], Hinds s.n. (HOLOTYPE: Herb. Lindley).
Epidendrum nonchinense Reichenb. f. in Walpers, Ann. Bot. Syst. 6: 324. 1862.
Laeliopsis chinensis (Lindley) Lindley ex Reichenb. f. in Saunders, Refugium Botanicum 2:
t. 139. 1882. (pro. syn.)
Barkeria nonchinensis (Reichenb. f.) Schlechter, Orchideen 206. 1914.
Epidendrum chinense (Lindley) Ames, Schedulae Orchidi~nae 7: 4. 1924.
Epidendrum strophinx Reichenb. f. in Linnaea 41: 78. 1876. T~PE: GUATEMALA (without
precise locality), H ort. Schiller.

P l a n t erect. Stem fusiform, 1.5-11 cm long, b e a r i n g 2-7 leaves, usually leafless

w h e n in flower. L e a v e s a l t e r n a t e , lanceolate to linear-lanceolate, a c u m i n a t e , 3 - 9 cm
long, 5 - 1 5 m m broad. I n f l o r e s c e n c e terminal, f r o m 2.5-35 cm tall, 2- to 20-flowered.
F l o w e r s a p p r o x i m a t e l y 1-2 cm across, cream to pinkish. Pedicels 1-2 cm long. Sepals
linear-lanceolate, a c u m i n a t e , 7-15 m m long, 1-3 m m wide, petals g r e a t l y resembling
sepals, linear-lanceotate to lanceolate, a c u m i n a t e , 7-15 m m long, 2 - 4 m m wide. Lip
s o m e t i m e s a d n a t e to c o l u m n for a b o u t 0.5 m m , deltoid-ovate, apex m u c r o n a t e , 1-1.5
c m long, 5-12 m m wide, callus consisting of three verrucose keels r u n n i n g n e a r l y to
a p e x of lip, the lateral veins of the lip also verrucose. C o l u m n with m e m b r a n a c e o u s
wings, 2 - 4 m m long, ca. 2 m m wide, anther cap s o m e t i m e s bright red. T h e c l i n a n d r i u m
1970] THIEN & DRESSLER: BARKERIA 291

TABLE I
A COZ~PARISONOr Barkeria WITg RELATED CENERA.

Epidendrum Barkeria Dimerandra

Range North America (North Mexico, Central America Mexico to northern

Carolina), Central South America
America, throughout
tropical South America
Approximate size 800 species 5 species 1 species
of group
Stem reed-like, sometimes fusiform pseudobulb elongate pseudobulbs
forming a pseudobulb
Callus wide variety of types, but absent or when present transverse lamellae
.not like Barkeria nor consisting of three raised
Dimerandra keels (verrucose)
Column completely adnate to lip appressed against lip free from the lip
(except 1 species),
adnate to lip up to
one-half the length of the
column, or free
Column wings absent widely spreading fleshy absent (two horns
or membranaceous projecting from the
apex)
Rostellum deeply slit (shallow in tongue-shaped without small tongue-like (in a
oerstedella) with a a viscidium wide notch) without a
distinct viscidium ~ziscidium

s m o o t h or v a r i o u s l y l o b e d ( s o m e t i m e s w i t h d e n t a t e m a r g i n s ) . O v a r y o c c a s i o n a l l y w i t h
a n e x t e r n a l l y v i s i b l e sac-like n e c t a r y .
Specimens examined:
MEXICO: MORELOS: Barranca de San Antonio near Cuernavaca, Ostlund (NageI) 2102 (MO,
US), on trees along river, B. Cruz 2130 ( F ) ; vicinity of Cuernavaca, Rose & Painter 1906 (US),
Nagel 4850 (MO) ; Barranca de Rio Joya, near Cuernavaca, 1500 m, Nagel 3172 (MO) ; Bejucos,
on a copal, Hinton 7390 (NY). STATE OF MEXICO: Tejupilco, on a Cirian, 1400 m, Hinton 5583
(K) ; Ixtapan, on a Cirian, 1400 m, Hinton 3649 (F, K, MO, NY, US). MICHOAC~N: Cuahuilote,
o.n a Cirian, Hinton 5566 (BM, US). GUERRERO: between Iguala and Telolo~pan, 1000 m,
Sawyer 845 (F) ; Barranca de Joveritos, Ostlund 3285 (MO) ; Vallecitos, o.n a tree, Hinton 11724
(BM, MO, NY, US); S of Chilpancingo, Clark 7199 (MO). V~RACR~YZ: Cd. Veracruz, Purpus
16273 (MICH). OAXACA: Road between Nopala and Mixtepec 800-1000 m, E. Nelson 2444 (US).
C~7~PAS: Stiltepec, Matuda 371 ( M I C H ) ; ~scuintla, Matuda 361 (MICH, MO, NY, US).
GUATEMALA: without precise locality, Salvi~ & Godman 47 (K); Gual~n, Dean 212 (MICH,
NY); Anibal, J. Smith 452 (US); Alta Verapaz, near Pancajache, Standley 91926 (F); Fiscal,
1100 m, Standley 80352 ( F ) ; Palo Gando, F. Morton 425 (F); without locality (no collector)
Herbarium numbers, 827, 55, 49, 5988 (W). 1 COSTA RICA: Rio Tirribi near San Jos~,
A1]aro 34000 (US), 23 Dec 1924, Al]aro s.n. (US) ; S~n Sebastian, Standley 32693 (US), 1160 m, 23
Feb. 1926, Standtey s.n. (US); vicinity of La Verbena, 1200 m, Standley 322251 (F, US); Las
Pavas, 1070 m, Standley 36067 (MICH, US) ; Heredia, 1000--1100 m, P. Allen 557 (F), 1924-1925,
Brenes s.n. (NY) ; Camino Hatilo, Alajuelito, 1150 m, Solis 96 (F) ; Rio Maria Aguilar near San
Jos~, Alfaro 36024 (MICH), 1100 m, Solis 69 (F) ; no locality, Herbarium numbers 13833, 35426
(W). NICARAGUA: Low hills near Granada, Maxon, Harvey, & Valentine s.n. (US); vicinity
of Managua near Lago de Nejapa, Maxon, Harvey, & Valentine 7565 (US). EL SALVADOR:
Tonacatepeque, Standtey 19429 (NY, US), S. Calderdn 216 (US); San Antonio del Monte,
250 m, Standley 22147 (US). PANAMA: Upper Chiriqui Valley, White & White 100 (F).

1Most specimens from the Vienna Herbarium (W) lack collection data, but are listed because
of the numerous colored drawings by Reichenbach.
292 BR~TTONIA [VOL. 22

FIC. 1. Distribution maps.

 1970] THIEN & DRESSLER: BARKERIA 293

HONDURAS: Comuatepeque, Edwards 364 (K); Las Mesas, 800 m, Rodriguez 2047 (F);
Zamorano, on tree, Standley 2167 (F), 1730 (F), L. Williams & Molina 11546 (MICH, MO, UC),
800 m, R. WiUiams s.n. (F); Rio de Capa Rosa, Rodriguez 3665 (F), 2147 (F), Standley
2167 (F).
The plants usually occur in great abundance in hedge rows and trees and apparently
become extablished very readily in any new openings in the habitat. Barkeria chinensis
ssp. chinensis occurs along the arid western coast of Central America from the state of
Michoac~n, M~xico, to the Upper Chiriqui Valley in Panama (Fig. 1), flowering from
December through February.
This widespread subspecies varies greatly in the size of the plants and in the number,
size, and shape of the leaves.
Barkeria chinensis ssp. chinensis is very closely related to ssp. naevosa and ssp.
palrneri. Barkeria chinensis ssp. palmeri usually has a greater number of flowers per
inflorescence, slightly larger flowers than ssp. chinensis, and a long sac-like nectary
visible near the summit of the ovary. Barkeria chinensis ssp. naevosa has a conspicuous
goiter-like nectary near the summit of the ovary (Fig. 2).
lb. BAI~KERIACHINENSIS ssp. p a l m e r i (Rolfe) Thien, comb. nov.
Epidendrum palmeri Rolfe in Kew Bull. 1893: 6. 1893. TYPE: MEXICO: COLIMA: Palmer
1201, Jan 1891. (ILOLOTYPE: NY; ISOTYPE: K).
Barkeria palmeri (Rolfe) Schlechter, Beih. Bot. Centralbl. 36: 470. 1918.
Epidendrum cubense Lindley, Bot. Reg. Misc. 17. 1843. TYPe: CUBA. (I~OLOTYPE:K).
Broughtonia cubensis Lindley ex Cogn. in Urban, Symbolae Antillanae 6: 542. 1910.
Laeliopsis cubensis Lindley ex Cogn. in Urban, Symbolae Antillanae 6: 543. 1910.
Plant slender, erect. Stem fusiform, 2-15 cm long, bearing 2-4 leaves. Leaves al-
ternate, linear-lanceolate, acuminate, 3-8 cm long, 7-13 mm wide. Inflorescence ter-
minal, 7-30 cm long, sometimes branched, bearing 2-30 flowers densely packed upon
the inflorescence. Flowers 2-2.5 cm across, pale lilac with a tinge of yellow in the
center of the lip. Pedicels 1.3-2 cm long. Sepals and petals resembling each other,
swept forward; sepals linear to linear-lanceolate, acuminate, 1-1.7 cm long, 2-4 mm
wide. Petals lanceolate, acute, 1-1.6 cm long, 3-4 mm wide. Lip sometimes adnate
to column for ca. 0.5 ram, lip obovate, apex rounded, often with a mucro, I-1.5 cm
long, and ca. 1 cm wide at the broadest part, margin of lip denticulate to various
degrees; callus consisting of three verrucose keels extending almost to the apex of lip,
the lateral veins of the lip also with verrucose thickenings. Column 2.5-4 mm long,
ca. 2 mm wide at the broadest part with widely spreading membranaceous wings.
Ovary slightly curved and with an externally visible sac-like nectary 3-4 mm long.
Specimens examined:
MEXICO: COLI~A: Rio Salado, Ikhanyma 3182 (MEXU). Herbarium numbers 42787, 53,
42793, 42792, 42789, 42791, all collected by Roezl (W). NAYARIT: Volcan Ceboruco near
Ahuacatl~in, an trees and shrubs, 1300 m, Nagel 5074 (F, MO) ; 20 mi S of Compostela, epiphytic
on Hura and other trees in canyon, Dressler 1062 (MO). JALISCO: Mountain trail from Puerto
Vallarta to San SebastiAn, near town of "Las Palmas," 600 m, Mexia 1317 (UC).
This subspecies occurs only in Mexico in the states of Colima, Jalisco, Nayarit, and
perhaps Sinaloa (Fig. 1), flowering from November through January.
We have tentatively listed Epidendrum cubense Lindley as a synonym for this
subspecies, as discussed by Dressler (1966). The fragmentary type suggests this
disposition, but future collections may show that E. cubense is truly Cuban and not
a Barkeria.

ic. BARKERIACHINENSIS ssp. n a e v o s a (Lindley) Thien, comb. nov.

Epidendrum naevosum Lindley, Folia Orchidacea 4. 1853. TYPE: MEXICO: Galeotti.
(HOLOTYPE: 5330 in Herb. Lindley; ISOTYPE: Herb. Reichenbach.)
294 BRITXONIA [VOL. 22

2
FIc. 2. A-E. Flowers of Barkeria. A. Barkeria elegans (flower X1; column Xl.5). B.
Barkeria barkeriola (flower X l ; column X1.5). C. Lip and spur of Barkeria chinensis ssp. palmeri
(lip X 2.5; spur X 1.5). D. Lip and spur of Barkeria chinensis ssp. naevosa (lip X 2.5; spur X 2).
E. Lip and column of Barkeria chinensis ssp. chinensis (lip > 1.5; column )<2.5).
1970] TH:EN & DRESSLER: BARKERIA 295

Barkeria naevosa (Lindley) Sehlechter in Repert. Sp. Nov. Beih. 19: 46. 1923, in textu.
Epidendrum chinense vat. naevosum (Lindley) L. O. Williams, Ceiba (Orchidaceae of Mexico)
2: 159. 1951.

Plant slender, erect. Stem terete, thickened, 4-15 cm long. Inflorescence 6-45 cm
tall, 3- to 12-flowered. Sepals and petals reddish, lip creamy gray with reddish veins.
Pedicels 1-2.3 cm long. Flowers 2.5-3.5 cm across. Sepals and petals similar. Sepals
lanceolate, acuminate, 1.3-2 cm long, 3-6 m m wide. Petals lanceolate, acuminate,
1.3-2 cm long, 3-6 mm wide. Lip sometimes slightly adnate to column; lip deltoid
to ovate in shape, 1.3-1.9 cm long, 9-12 m m wide (at broadest part), apex somewhat
rounded to sharply acute-acuminate; keel consisting of three verrucose lines running
almost to apex of lip, the lateral veins with verrucose thickenings. Column 3-4 mm
long, 2.5-3.5 mm wide, with widely spreading membranaceous wings. Ovary with a
swollen conspicuous goiter-like nectary near summit, 2 mm long, 1 mm wide.
Specimens examined:
MEXICO: GUE~Rr.RO: Km. 450 between Acapulco and Tierra Colorada, on trees, Sawyer
76 (F); along Rio de la Colorada near Carrizal, on cacahuananche tree, 600 m, Sawyer 830 (F).
OAXACA: (no collector) Herbarium numbers 46, 5989, 48, (W).
Mr. Glenn Pollard (personal communication) states that this subspecies is indigen-
ous only on the Pacific slope in Mexico from Juchitan (Oaxaca) approximately to
the Rio Balsas (Fig. 1 ), which forms the boundary between the states of MichoacAn
and Guerrero. Records show that it grows in dry and sparsely wooded areas on trees
near flowing creeks.
This subspecies flowers from J a n u a r y through March.
2a. B A ~ K ~ A LINDLEYANABateman ex Lindley ssp. LINDLEYANA
Barkeria lindleyana Bateman ex Lindley in Bot. Reg. 28: 2. 1842. TYPE: COSTA RICA,
Skinner.
Epidendrum lindleyana (Bateman ex Li.ndley) Reiehenb. f. in Walpers, Ann. Bot. Syst. 6:
375. 1862.
Barkeria lindleyana var. centerae Reichenb. f. in Gard. Chron. 1873: 1597. 1873. TYPE:
COSTA RICA, Endres. s.n.
Epidendrum lindleyana var. centerae (Reichenb. f.) Veitch, Manual Orchidaeeous Plants
6: 105. 1890.
Plant slender, erect. Stem terete, slightly thickened, 12-24 cm long, bearing 3-5
leaves. Leaves alternate, slightly thickened, lanceolate-elliptic, acute, 8-15 cm long,
2-3.2 cm broad, keeled below. Inflorescence terminal to 50 cm tall, I0- to 15-flowered.
Flowers as much as 5 cm across, dark lilac and white. Pedicels 3-3.5 cm long. Sepals
lanceolate, acute about 2.5 cm long, 8-10 mm wide. Petals ovate, sharply acute, 2.6-3
cm long, 1.3-1.5 cm wide. Lip oval to elliptic to oblong in shape. Lip base slightly
enfolding the base of the column. Margin of the lip undulate, lip dark or pale lilac
with a white to yellow spot in the center and with the apex (upper third of lip) a
very dark lilac. Callus consisting of three thickened keels running almost to the apex
of the lip, white to yellow with several small purple dots. Lip 2.4-3.3 cm long, 1.7-2
cm wide, apex of lip sharply apiculate to acute. Column 1.2-1.4 cm long, about 5 mm
wide, with slightly fleshy wings, back of the column pale lilac with the wings white,
yellow, and green with numerous small dark purple dots.
Specimens examined:
COSTA RICA: San Jos~, Pittier 6909 (US), 11oo m, Brade 16671 (US), Sep 1924, Allaro s.n.
(US) ; Orosi, 9 Aug 1924, Al/aro s.n. (US), Endres 40 (W) ; Caseajal, 1600 m, Al/aro s.n. (US).
The " l i n d l e y a n a complex" is composed of ssp. l i n d l e y a n a , ssp. spectabilis, ssp.
cyclotella, and ssp. v a n n e r i a n a . This group has been exceedingly difficult to delimit,
296 BRITTONIA [VOL, 2

~84184184184184

3
 1970] THIEN & DRESSLER: BARKERIA 297

I I W

BARKEItlALINDtEYAE
S S P L'NDLEY.~NA
SS~ CYCLOTE,LLA
ANNERIANA
SSI)~
4
FIG. 4. Distribution map.

and its study is hampered by a lack of representative specimens. The members of the
complex occurring in Mexico north of the Isthmus of Tehuantepec occupy definite
altitudinal zones.
Barkeria lindleyana ssp. lindleyana (Fig. 3) is found only in Costa Rica (Fig. 4),
and flowers from July through September.

2b. BARKERIALINDLEYANAssp. spectabilis (Bateman ex Lindley) Thien, comb. nov.

Barkerla spectabilis Bateman ex Lindley in Bot. Reg. 28: Misc. 43. 1842. TYPE: GUATE-
MALA: Skinner. (HOLOTYPE: in Herb. Lindley.)
Epidendrum spectabile (Bateman ex Lindley) Reichenb. f. in Walpers, Ann. Bot. Syst. 6: 375.
1862; nan Focke, 1853.

Plant stubby to slender, erect. Stem terete, slightly thickened, 5-12 cm long,
bearing 2-4 leaves. Leaves alternate, lanceolate, acute, 2-11 cm long, 1-3 cm wide.
Inflorescence terminal from 7-22 cm tall, 2- to 10-flowered. Flowers 4-7 cm across,
pale lilac and white. Pedicels 2-4 cm long. Sepals lanceolate to elliptic acute, 2-3.8
cm long, 0.5-3.8 cm long, 0.7-1.8 cm wide. Lip sometimes adnate to column for 1-3
mm, lip oval, ovate to elliptic in shape, apex obtuse to acute sometimes with a very
slight mucro, 2-3.5 cm long, 1.4-2.3 cm wide, keel (callus) consisting of three lines
running three-fourths the length of the lip, lip pale lilac with numerous red spots.
Column with membranaceous wings, 1.4-1.7 cm long, 4-7 mm wide.

<.-
Fro. 3. A-E. Flowers of Barkeria. A. Barkeria lindleyana ssp. lindleyana (flower X 1; column
X1.5). B. Lip and column of Barkeria lindleyana ssp. vanneriana (lip Xl.5; column X1). C.
Barkeria lindleyana ssp. spectabilis (flower X l ; column X2). D. Lip and column of Barkeria
lindleyana ssp. cyclotella (lip X1; column X1). E. Barkeria melanocaulon (flower X3; column
• 1.5).
298 BRITTONIA [VOL. 22

Specimens examined:
MEXICO: CmAPAS: Pan American Highway between Las CasEs and Comit~n, Carlson 1693
(CN, M, UC); SW of Comit~in, dark oak forest on trees, 1600 m, Nagel (Ostlund) 5690 (CN,
US); La Grandesa, 2016 m, Matuda 5603 (MO, UC, US). GUATEMALA: without precise lo-
cality, Bernoulli 767 (NY); DEPT. SOLOL.~: Chocoyes, Steyermark 46386 (CN). DEPT. Qt~IC~:
Santa Cruz Quich6, Lehmann 1505 (K, US). DEPT. SACATEI"~-QUEZ:Embaulada, 1800 m, Heyde 6"
Lux 4600 (US).

This subspecies commonly grows on oak trees and is found in the state of Chiapas,
Mexico, and south into Guatemala (Fig. 4). Subspecies s p e c t a b i l i s differs from ssp.
l i n d l e y a n a in having a larger lip that is lighter in color with numerous red spots
(Fig. 3). T h e plants tend to be s t u b b y in a p p e a r a n c e and bear two or three flowers.
The plants bloom in March, May, and December.

2c. BARKERIA LINDLEYANA ssp. v a n n e r i a n a (Reichenb. f.) Thien, comb. nov.

Barkeria vanneriana Reichenb. f. in Gard. Chron. n.s. 24: 678. 1885. TYPE: Hort. Vanner
ex Sander. (ltOLOTYPE: in Herb. Reichenbach).
Epidendrum vannerianum Reichenb. f. in Gard. Chron. n.s. 24: 678. 1885, in textu.

P l a n t slender, erect. Stem terete, slightly thickened, 6-15 cm long, bearing 3-7
leaves. Leaves alternate, lanceolate to b r o a d l y ovate or elliptic, acuminate to acute,
3-11 cm long, 1.5-4.5 cm broad. Roots massive 0.5 cm in diameter. Inflorescence
terminal from 15-60 cm tall, 3- to 15-flowered. Flowers 3-4.5 cm across, pale lilac
and white. Sepals lanceolate, acute to acuminate, 2-2.5 cm long, 9-14 m m wide. Lip
a d n a t e to column for 2-3 mm, lip slightly p a n d u r a t e to obovate or sub-rectangular,
apex rounded to square with a slight mucro, 1.7-2.3 cm long, 1-1.7 cm wide, callus
consisting of three keels running almost to apex of lip (middle keel running beyond
the other two). Column with widely spreading membranaceous wings, 1-1,4 cm long,
4 - 6 mm wide at the broadest part.
Specimens examined:

MEXICO: PI~EBLA: near Texcala, very dry country on mesquite trees, 1800 m, Nagel 2566
(CN, MO, US); limestone ledges near Tehuac~n, 5000 ft, Pringle 6769 (BM, CN, K, MEXU,
MO, UC, US, W); vicinity of San Luis Tultitlanapa, C. Purpus 3555 (CN, MO, NY, US);
Tehuadm, Sawyer 201 (CN), Rose, Rose & Painter 9937 (US). OAXACA: near Santa Maria
de Tule, dry mountains not far from the town of Oaxaca, 1600 m, Velde (Ostlund) 5430 (MO,
US); SW of Chilpancingo, Mt. Prieto, mostly on rocks, 1800 m, Nagel 3223 (CN); Cerro
de Tule, near Cd. Oaxaca, 1900 m, Conzatti 7072 (MO) ; El Pari6n, Conzatti 1553 (CN), on lava
at 5000 ft, Galeotti 5068 (K); Santa Catarina, Rusby 39 (NY, US); San Felipe, Conzatti 2007
(CN); (No collector given) Herbarium numbers 42795, 42798, 42807, 42708, 42806, 42812,
42811 (W). GUERRERO: Idolo Vinatita, high on cliffs above Rio del Oro, 1250 m, Hinton
9732 (BM). VERACRIIZ: Orizaba, Purpus 1363 (UC).

Subspecies v a n n e r i a n a is found in Mexico in the states of Puebla, Guerrero, Oaxaca,

and Veracruz (Fig. 4) growing on rocks, cacti, and mesquite. I t blooms from J u l y
through October.
Subspecies v a n n e r i a n a differs from ssp. l i n d l e y a n a in having a 3-keeled callus and
a lip that varies in shape from pandurate to sub-rectangular (Fig. 3).

2d. BARKERIA LINDLEYANA ssp. e y e l o t e l l a (Reichenb. f.) Thien, comb. nov.

Barkeria cyclotella Reichenb. f. in Gard. Chron. n.s. 13: 72. 1880. TYPE: Hort. Bull 6" Day.
(I:fOLOTYPE: in Herb. Reichenbach.)
Epidendrum cyclotellum Reichenb. f. in Gard. Chron. n.s. 13: 72. 1880, in textu.
Epidendrurn lindleyana var. cyclotellum (Reichenb. f.) Veitch, Manual Orchidaceous Plants 6:
105. 1890.
1970] T t I I E N & DRESSLER: BARKERIA 299

Plant slender, erect. Stem terete, slightly thickened, 8-15 cm long, bearing 3-6
leaves. Leaves alternate, ovate-elliptic, acute to acuminate, 5-10 cm long, 1.5-3.5
cm broad. Roots massive, 5 mm in diameter. Inflorescence terminal from 10-35 cm
tall, 3- to 9-flowered. Flowers 3-4.5 cm across, deep magenta-red. Pedicels 2-3 cm
long. Sepals lanceolate, acuminate, sometimes slightly clawed, 1.5-2 cm long, 4-6
mm wide. Petals ovate to elliptic, acute 1.7-2.4 cm long, 6-10 mm wide. Lip adnate
to column for 2-3 mm, lip obovate, apex rounded with a small mucro, 1.5-2 cm long,
1.2-1.5 cm wide, callus consisting of three slightly thickened keels running almost
to the apex of the lip. Column with widely spreading membranaceous wings, 1-1.2
cm long, 4 - 6 mm wide at the broadest part, with numerous dark purple dots on back
of the column.
Specimens examined:

MEXICO: MICHOAC~A'r Lago de Cuitzeo, near Chucandiro, Mt. Coronillas, dry places on
cacti, trees, and rocks. 1800 m, Vera (Ostlund) 7110 (MEXU, MO, US).
This subspecies is found in Mexico within the states of MichoacAn and Oaxaca
(Fig. 4), flowering in November and December. It differs from ssp. lindleyana in
having dark red-magenta flowers and an obovate lip with a 3-keeled callus (Fig. 3).

3. BARKERIA SKINNERI (Bateman ex Lindley) Richard & Galeotti.

Barkeria skinneri (Bateman ex Lindley) Richard & Galeotti in Comptes Rend. Acad. Sci.
Paris 18: 506. 1844, nomen; Lindley ex Paxton, Mag. Bot. 15: 1. 1849, in textu.
Epidendrum skinneri Bateman ex Li.ndley in Bot. Reg. 22: t. 1881. 1836. TYPE: GUATE-
MALA, Skinner.
Barkeria sklnneri var. major Paxton, Mag. Bot. 15: 1. 1849.
Dothiolophis purpurea Rafinesque, Flora Telluriana 4: 39. 1838. (based on Epidendrum
skinneri Lindley f., Bot. Reg. 1881)
Epidendrum Fuchsil Regel, Index Seminum Hort. Turic. 4: 1850; in Schweiz. Zeitschr.
Gartenb. 9: 202. 1851. TYPE: GUATEMALA: Hort. Fuchs.
Epidendrum skinneri Bateman ex Lindley [var.] superbum Warner, Select Orchideous Plants
1: 38. 1862-1864. TYPE: GUATEMALA: Skinner ex Hort. Veitch.

Plant slender, erect or ascending, loosely caespitose. Stem slightly thickened,

fusiform, bearing 3 or 4 leaves, (sometimes leafless when in flower). Leaves alternate,
linear-lanceolate to lanceolate, acuminate to acute, 3-18 cm long, 1-2.8 cm broad.
Inflorescence terminal, raceme rarely compound, from 9-45 cm long, 4- to many-
flowered. Flowers 1.5-2.5 cm across, rose-purple or red-magenta. Pedicels 2-4 cm
long. Sepals elliptic to lanceolate, acute-acuminate, 1.5-2.5 cm long, 4-5 mm wide.
Petals elliptic to ovate, acute to sharply acuminate, 1.5-2.4 cm long, 7-12 mm wide.
Lip adnate (along margins) to column for about one-half the length of the column,
lip ovate in shape often resembling the petals in size, apex acute to sharply acuminate,
1.5-2 cm long, 7-13 mm wide, callus consisting of three yellow, thickened keels
running to the apex of the lip. Column club-shaped with more or less fleshy wings,
6-7 mm long. Ovary sometimes with an externally visible nectary on the lower side.
Specimens examined:
GUATEMALA: without precise locality, 1800 m, Sala 1471 (CN), Aguilar 143 (CN);
Amatitl~in, 1102 m, Morales 1163 (CN), SaIvin & Godman 161 (K); mountains along the road
between Jalapa and San Pedro Pinula, 1400-1800 m, Standley 77064 (CN); vicinity of Jalapa,
1360 m, Sandley 76776 (CN); Huehuetenango, Shannon 462 (US); Los Llanitos, near San Jos6
Acatempa, 1200 m, Standley 60590 (CN); Santa Rosa, Casillas, Hcyde & Lux 4276 (NY, US),
in t862, Salvin & Godman 101 (K) ; bought in market in Cd. Guatemala, Standley 57847 (CN) ;
Herbarium numbers (no collectors) 42815, 42831, 42824, 42823, 42825, 42826, 42828, 42897, 42830,
3864, 42829, (W) ; Nov., 1865, G. Bernoulli (W).
3 00 BRITTONIA [VOL. 22

5 Fic. 5. Barkeria skinneri. A. Flower X1. B. Lip (X1.5) and column (X2).

Barkeria skinneri (Fig. 5) occurs in Guatemala (Fig. 1) and flowers in November

and December.

4. BARKE.RIABARKERIOLAReichenb. f.
Barkeria barkeriola Reichenb. f. in Gard. Chron. n.~. 22: 616. 1884. TYPE: Hort. Sander.
Epidendrum barkeriola Reichenb. f. in Gard. Chron. n.s. 22: 616. 1884. (pro syn.)

Plant slender, erect. Stem terete, slightly thickened, 4-16 cm long, bearing 4 or 5
leaves. Leaves alternate, lanceoIate, acuminate to acute, 1.5-10 cm long, 7-20 mm
broad, reddish underneath with a reddish margin above. Inflorescence terminal, from
4-27 cm tall, 2- to 10-flowered. Flowers 2.5-3.5 cm across, pale lilac and white.
Pedicels 1.5-2.5 cm long. Sepals slightly longer than petals, not reflexed, clawed,
lanceolate to oblanceolate, acute, 1.5-3 cm long, 3-9 mm wide. Petals ovate to
oblanceolate, acute, 1.5-2.5 cm long, 5-9 m m wide. Lip adnate to the column
for about 3 mm, pandurate, apex mucronate to rounded, 2-3.5 cm long, ca. 5
mm wide near the base, callus absent except for two slightly incurved lines near
the base. Column with membranaceous wings, 1-1.4 cm long, 4-7 mm wide at the
broadest part, white-yellow with numerous purple spots on the back. Ovary some-
times with a sac-like nectary on the lower side.
Specimens examined:
MEXICO: COLIMA: foot of Volcan Colima, Ostlund 4028 (F, NY, US), and 4028.4 (MO).
SINAZOA: "La Rata," 4 leagues from "La Noria," Mexia 365 (UC) ; San Ignacio, Montes & Salazar
675 (US) ; Los Tepemesquites, Ortega 5718 (US).

This species is found in Mexico in the states of Colima, Sinaloa, Jalisco, and Nayarit
(Fig. 1 ). It flowers in October and November.
Barkeria barkeriola is closely related to B. elegam but differs in having sepals and
petals that are subequal, smaller flowers, and lacks the widely spreading mem-
branaceous column wings (Fig. 2).
1970] THIEN & DRESSLER: BARKERIA 301

5. B A R K E R I A ELEGANS K n o w l e s & W e s t c o t t .
Barkeria elegans Knowles & Westcott, Floral Cabinet 2: 7. 1838. TYPE: MEXICO: Hort.
Barker.
Epidendrum elegans (Knowles & Westcott) Reichenb. f. in Walpers, Ann. Bot. Syst. 6 : 374.
1862.

Plants slender, erect, caespitose. Stems terete, slightly thickened, 4-16 cm long,
bearing 2-6 leaves (sometimes leafless when in flower). Leaves thin, alternate, linear
to lanceolate, acuminate to acute, 3-12 cm long, 6-20 mm wide. Inflorescence a
terminal raceme from 5-26 cm long, 1- to 4-flowered. Flowers 4-6 cm across. Pedicels
2.5-3 cm long. Sepals and petals mallow-pink. Sepals sometimes clawed, elliptic-
lanceolate, acute, 2.5-4 cm long, 8-12 mm wide. Petals obovate to elliptic-ovate
acute to obtuse, 2.5-3.5 cm long, 1-1.5 cm wide at the broadest part. Lip white with
a large purple blotch on apex, adnate to column for 3-4 mm, appressed against lip,
pandurate to obovate in shape, apex emarginate to apiculate, 2-3 cm long, 1.5-2.5 cm
wide, callus consisting of three to five thin keels. Column with widely spreading
membranaceous wings, 1.5-2 cm long, 8-13 mm wide at the broadest part.
Specimens examined:
MEXICO: MICWOAC~N: Coalcom~in, 1000 m, Hinton 12500 (F, MO, NY, US); Naranjillo,
Hinton 16053 (US); E1 Temalscal, 750-850 m, Moore, Herndndez, & Porras 5674 (UC); Sierra
Madre, Cerro Verde, 300-1000 m, Langlassd 610 (K, MEXU, US). GUERRERO: Vallecitos,
Hinton 11468 (BM, MO, US) ; San Juan de las Flores, Galeana, Hinton 11214 (K, US) ; Barranca
de Joveritos near Km. 350 on road to Acapulco, 1500 m, Ostlund 3282 (F, MO, US); Campo
Morado, 1240 m, Hinton 11155 ( E M ) ; between Petatl~n and Chilapa, E. Nelson 2153 (US);
Paraje Joveritos, Ostlund 5202 (US); "La Cebada" near Plan del Molino, N of Atoyac, in dry
mixed forest, Sawyer 2048 (MEXU) ; La Junta, Santo Tom,is, 750 m, Matuda 29415 (MEXU) ;
entre Malpai y Santa Barbara, 1300 m, Matuda 30134 ( M E X U ) . MEXICO: Calera,
Temascaltepec, 770 m, Hinton 2557 (EM, K) ; Pungarancho, 950 m, Hinton 5128 (US) ; Pericones,
1500 m, Dec., 1932, no collector (EM). OAXACA: without precise locality, Galeotti 5318 (W);
Herbarium numbers (no collector) 42818, 42821, 42816, 42820, (W).

The species (Fig. 1) occurs in Mexico from the states of Mexico and Michoadm
to Oaxaca (Fig. 2), and flowers from October through January.
6. BARKERIA MELANOCAULON R i c h a r d & G a l e o t t i
Barkeria melanocaulon Richard & Galeotti in Comptes Rend. Acad. Sci. Paris 18: 512. 1844,
nomen; in Ann. Sci. Nat. Bot. III. 3: 22. 1845. TYPE: MEXICO: Galeotti 5069.
(~OLO~YPE: P; ISOTYPE: W )
Epidendrum melanocaulon (Richard & Galeotti) Reichenb. f. in Walpers, Ann. Bot. Syst. 6:
375. 1862.
Epidendrum whartonianum Schweinfurth, in Amer. Orchid Soc. Bull. 1 7 : 316-318. 1948.
TYPE: MEXICO: OAXACA: Nizanda, Ames 64690. (HOLOTYPE: AMES)

Plant slender, erect. Stem terete, thickened, to 17 cm long, bearing 2-5 leaves.
Leaves alternate, elliptic to lanceolate, acute to acuminate, 4-12 cm long, 1-2.4 cm
broad. Inflorescence a terminal raceme to 8 cm tall, from 4- to many-flowered
(18-20). Flowers 2-3 cm across, rose-lilac. Pedicels to 2.5 cm long. Sepals and
petals reflexed. Sepals lanceolate-elliptic, acute to acuminate, 12-15 mm long, 4-8
mm wide. Lip free or slightly adnate to column at base, not appressed, lip heart-
shaped to more or less suborbicular, apex rounded to emarginate with a slight mucro,
the margins undulate, the base more or less cordate, 10-16 mm long, 8-13 mm wide,
with three keels running almost to apex of lip, the anterior portion usually slightly
higher than the rest of the keel and with a few dark purple dots and streaks (white
incurved teeth near base). Column 7-10 mm long, green and purple, 6-8 mm wide,
the wings of column usually fleshy.
302 BRITTONIA [VOL 22

Specimens examined:
MEXICO: OAXACA: Sierra San Pedro, Talea, Jurgensen 830 (W); Talea, Galeotti 5836 (W);
Herbarium numbers (no collector) 42796, 42797, and 42800 (W).
This species (Fig. 3, E) is apparently found only in the state of Oaxaca, Mexico,
(Fig. I), where the two definite localities are known (Nizanda and Talea). At
Nizanda the plants characteristically grow on the north side of cacti and rocks. The
plants flower in December and January.

ACKNOWLEDGMENTS
We thank the curators and staff of the following herbaria for the opportunity of
examining material in their care: AMES, BM, CN, EM, F, K, MEXU, MICH,
MO, NY, UC, US, and W.
L I T E R A T U R E CITED
Ames, O., F. T. H u b b a r d , & C. S c h w e i n f u r t h . 1936. The Genus Epidendrum in the United
States and Middle America. Botanical Museum, Harvard Univ., Cambridge, Massachusetts:
xi q- 233 pp.
Dressier, R. L. 1961. A reconsideration of Encyclia (Orchidaceae). Brittonia 13: 253-266.
1966. Nomenelatural notes on the Orehidaceae III. Taxon 15: 241-243.
Knowles, C. B. & F. VC'esteott. 1883. Barkeria elegans. The Floral Cabinet. 2: 7, 8.
Ostlund, E. Sketches and notes on Mexican orchids (unpublished), on file at Smithsonian
Institution, Washington, D.C.