The hexandrous species of Topobea (Melastomataceae)

13 Pages·2000·5.2 MB·English

by  Frank Almeda

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume52, No. 9,pp. 97-109,4 figs. October 18, 2000
The Hexandrous Species of Topobea (Melastomataceae)
by
FranktAAlmeda ^ pHnn
DepartmentofBotany, CaliforniaAcademyofSciences
Golden GatePark, San Francisco, California 94JJ8 ^Ooog^
Akeyandupdatedgeographicandtaxonomicnotesarepresentedforthefiveknownspe-^
ciescomprisingthe hexandrousctadeofTopobea. Topobeaarboricola, a Panamanian en-
demic described and illustrated here, is distinguished by its six stamens per flower,
2-locularovary,elliptictoovateleaveswith acaudate-acuminateto long-attenuateapex,
andunappendagedantherconnectives.Theneedforrecordingfielddataontaxonomically
useful floral and fruitcharactersisemphasized.
Resumen
Sedescribeunaespecienueva, Topobeaarboricola dePanama.Seproveendescripciones,
notassobredistribucion yfenologia,ydiscusiones referentes a lasafinidadesdelascinco
especiesde Topobeaquetienenseisestambres.Tambiensepresentaunaclaveymapasde
ladistribucionparatodasellas.Elcolorytamanodelospetalosyestambres,formadelos
poros de las anteras, y el numero de loculos en el ovario son a menudo caracteres
taxonomicos muy utilesen Topobea.
Topobea, with approximately 70 species, is aberry-fruited genus ofmostly shrubby epiphytes
and hemiepiphytes restricted to wet forests of tropical America. Over 55 species occur in the
biodiversity hotspot extending from Costa Rica south to Colombia.
Traditionally, Topobea has been defined by its 6-merous isomerous flowers with 12 stamens,
6-locularovary, andelongateantherthecaewithbroadconfluentapical pores. Although thischarac-
terization applies to a majority ofspecies in Topobea, discovery ofnew species in southern Central
Americainthepasttwodecadeshasnecessitatedabroadenedcircumscriptionofthegenustoaccom-
modatetheaccumulationofnewandunusual speciesandareconsiderationofgeneric limitsbetween
Topobea and its sistergenus Blakea (Almeda 1990).
I here provide a review ofthe most specialized evolutionary line within the genus that T infor-
mallyrefertoasthehexandrous species. These species,which are largelycenteredin Panama, share
several synapomorphiccharactersthatinclude: (1)6-merousflowerswithsixstamens,eachofwhich
isattachedtothehypanthial torusoppositeacalyx lobe; (2) subsessileorshort-pedunculateflowers;
and(3) 2-locularor4-locularcompletelyinferiorovaries. Until recently, mostofthe speciestreated
herewere known fi-om fewcollections. A studyofadditional material generatedbymyrecent field
workandthatofcolleaguesnowmakesitpossibletoemendandaugmentmorphologicalanddistribu-
tional information fordescribedspecies, propose anothernew species in this alliance, andprovide a
preliminary assessmentofinterspecificrelationships within this well-defined clade.
Understanding intraspecific variation in woody epiphytes has always been a challenge. Many
epiphytic species ofMelastomataceae are composed ofwidely dispersed, low density populations.
This coupledwith spottysamplingbycollectors andthe fragmentary preservafion or loss ofcritical
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Volume52, No. 9
floral characterson driedspecimenshascontributedtothepaucityofadequate studymaterial. Petal
shapeandcolor,stamennumber,morphologyofantherpores,andovaryloculenumberareessential
charactersforthedelimitationofspeciesin Topohea.Unfortunately,someofthesecharactersaredif-
ficult to evaluate once specimens have been pressed and dried. Ideally, notes on these characters
shouldberecordedinthefield,ifthepreservationofflowersandfruitsinaliquidmediumisnotpossi-
ble.
KeytotheHexandrousSpeciesof Topobea
la. Leavessessileandcordate-clasping;calyxlobesoblong-ovate;petalswhite,8-13mmlong;antherssubulate,5-7mm
long T. cordata
lb.Leavesdistinctlypetioled,varyingfromelliptic-ovatetoobovateorsubrotund;calyxlobeslance-triangular;petals
pink,4.5-7mmlong;anthersbluntlyoblong,2-2.5mmlong.
2a.Ovary4-locular,antherporesdorsallyinclinedatthetruncateapex;antherconnectiveelevateddorso-basallyintoa
bluntspurlikeappendage T. crassifolia
2b.Ovary2-locular,antherporessomewhatventrallyinclinedatthetruncateapex;antherconnectiveunappendaged
dorso-basally(ifconnectiveisappendagedthenuppermostcaulinenodesbesetwithspreadingtuftsofhairs).
3a. Leavesthickandleatherywhendry,theapexobtusetorounded,theabaxialsurfaceglandularpunctate.
4a.Hemiepiphyticshrubwithadventitiousrootsonthecaulineintemodes; uppermostcaulinenodesbesetwith
caducoustuftsofspreadinghairs;leavesovate-elliptictosubrotund;CerroJefe,Panama . . . T. hexandra
4b.Terrestrialshrubwithoutadventitiousrootsonthecaulineintemodes;uppermostcaulinenodeslacking
tuftsofspreadinghairs;leavesobovatetooblanceolate;CerroTute, Panama T. caliginosa
3b. Leavesnotthickandleatherywhendry,theapexcaudate-acuminate,theabaxialsurfacenotglandular
punctate T. arboricola
Species Description
1. Topobea arboricola Almeda, sp. nov.
Fig. 1
—
Type. PANAMA.BocasdelToro/Chiriquiborder:windsweptcloudforestonslopesandval-
leysofthe Cerro Coloradoregion, elev. 1450 m, 27Jan. 1989,Almedaetal. 6456(holotype: CAS!;
isotypes: MO!, PMA!).
Frutex epiphyticus. Lamina 3.8-7.1 x 1.6-3.9 cm elliptica vel ovata apice caudato-acuminata
vel attenuata basi acuta vel rotundata, 3-5-nervata papyracea et integra, nervis secundariis
nervulisque invisis; petioli 0.9-2.8 cm longi. Flores 6-meri sessiles vel subsessiles in quoque nodo
superiori singuli; bracteae omnino liberae; bracteae exteriores 2.5^ x 1.5-4 mm, ellipticae vel
ovato-ellipticaeapiceacutovelobtuso;bracteaeinteriores2-3 x 2-3 mm,ovataevel suborbiculares
mm mm mm
apice rotundato. Calycis tubus 0.5 longus, lobis 2 longis. Petala 5 x 3.5 obovata vel
mm
obovato-elliptica apice obtuso. Antherae 6, ca. 2 x 0.5 oblongae inter se non cohaerentes;
connectivum nee prolongatum nee appendiculatum. Ovarium 2-loculare et omnino inferum apice
glabro (cono et collo non evoluto).
Epiphyticshrubto 1.5 mtall.Uppermostbranchletsglabrous,somewhatcompressedandbluntly
two-edged;olderbranchesbecoming±roundedwithnodularleafscars. Matureleavesofapairequal
to somewhat unequal in size, glabrous throughout; petioles 0.9-2.8 cm long; mature blades
papyraceous, 3.8-7.1 cm long and 1.6-3.9 cm wide, elliptic to ovate, apex caudate-acuminate to at-
tenuate, base acute varyingto rounded, margin entire, 3-5-nerved, the outermost intramarginal pair
oftendepressedandinconspicuous,thetransversesecondaryveinstypicallynotelevatedorconspicu-
ous. Flowers erect, solitary in leafaxils ofuppermost branches, sessile, subsessile, or with short
(1-2 mm) ill-defined peduncles formed by the compressed bases ofthe outer floral bracts. Floral
ALMEDA: TOPOBEA 99
Figure 1. TopobeaarboricolaAlmeda. A. habit, x ca. Vr, B. representativeleaf(abaxial surface), ^ 1; C. youngfruiting
hypanthium with decussate floral bracts, x 6; D. stamens, ventral view (left) andpartial profile view (right), x 7; E. petal
(adaxialsurface), x 7;F.berry(topview)showingcalyxlobes,ovarysummit,andtoruswithstamina!filamentscars, x4;G.
seeds, x 15.(A,Bfromtheholotype;C-EfromMcPherson 12813,MO;F,GfromMcPherson 7739,CAS.)
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bractsfreeandentire,glabrateorsparinglybesetabaxially(mostlytowardtheapex)withalanateor
stellulate-lepidote,copperybrown indument; outerbracts2.5-4 x 1.5-4 mm,concave, elliptictoel-
liptic-ovate, apex acutetoobtuse; innerbracts2-3 x 2-3 mm,depressed-ovatetosemicircular, apex
rounded. Calyxtube0.5 mmlong;calyx(infruit)erect,2 mm longand2 mmwidebasally,deltoidto
deltoid-ovate, entirebutcallose-thickened alongthe interlobe sinuses, sometimes sparingly beset at
the abaxial apex with an indument like that of the floral bracts. Petals 6, sparingly beset with
disc-shapedglandswhendry,5 x 3.5 mm,translucentpink,obovatetoelliptic-obovate,apexobtuse,
mm mm mm
entire. Stamens 6, filaments complanate and glabrous, 2.5-3 long; anthers 2 long, 0.5
wide,paleyellow,oblongwithtwobroadporesatthetruncateapex;connectivebarelythickenedand
unappendaged. Ovary completely inferior, 2-locular, apex glabrous, stylar scarevident but not ele-
vatedintoaprominentconeorstylarcollar.Stylesomewhatdeclinateandsigmoidapically,glabrous,
5 mm long; stigmapunctiform. Berry4-6 x 4.5-6 mm,whiteatmaturity. Seedscuneatetonarrowly
mm
deltoid, 1 long—, tan with a smooth, glossytesta.
Phenology. Flowering in February, May, July and August; fruiting specimens have been
collected in July, Dec—ember, andJanuary.
Distribution. Known only from wet cloud forests of western Panama from Valle de la
m
Sierpe (Chiriqui)—east to Cerro Tute (Veraguas) at 1000-1500 (Fig. 2).
Paratypes. PANAMA. Bocas del Toro/Chiriqui border: above Fortuna dam, along divide
trail, ca. 8°45'N, 82°)5'W, 4 Dec. 1985, McPherson 7739(CAS, MO). Chiriqui: Fortuna dam area,
alongQuebradaBonitatoEofroad,8°45'N,82°13'W,8Feb. 1984,Churchilletal 4759(MO);Valle
de la Sierpe, en direccion SE a lo largodeQuebrada Bonita, 17 May 1987, Correaetal. 5092 (MO,
PMA); Fortuna dam area, N of reservoir, ridge along continental divide and southward from
QuebradadeArena,Aug. 1984(nodaygiven),D'/lrcvcS Todzia /5959(CAS,MO).Veraguas:trailto
ReservaBiologicaSerraniadeTuteabout0.7 kmbeyondtheEscuelaAgricolaRioPiedrajustoutside
ofSantaFe, 18Feb. 1996,Almedaetal. 7622(CAS,MO,PMA,US);DistritodeSantaFe,Serraniade
Tute,8°33'N,81°07'W,5Jul. 1996,Arandaetal. 2676(CAS,SCZ);DistritodeSantaFe,Serraniade
Tute,8°33'N,81°07'W,5 Jul. 1996,Arandaetal. 2657(CAS, SCZ);vicinityofCerroArizona-Tute,
above Santa FeandAltosPiedra, alongtrail to summit, 8°30'N, 81°10'W, 28 Jul. 1988, McPherson
12813 (MO). —
Discussion. Topobeaarhoricolaappearstobeatrueepiphyticshrubthatneverhasrootcon-
tact with the ground. It is distinguished by its 2-locular ovary, comparatively long petioles
(0.9-2.8 cm), elliptictoovateleaveswith acaudate-acuminateto long-attenuateapex, andshortob-
long antherswith unappendaged connectives.
OfthethreehexandrousspeciesofTopobeawith2-locuIarovaries, T. arhoricola ismostsimilar
to T. caliginosa.Thelatterdiffersinbeingaterrestrial shrubandinhavingobovateleaveswithanob-
tusetoroundedapex, shorterpetioles (0.5-1 cm), longerfloral peduncles (3-5 mm), andabaxial fo-
liar surfaces that are inconspicuously glandular-punctate.
All collectionsofT. arhoricolacomefromascatteringoflocalitiesalongtheCordilleraCentral
ofPanama. Foranepiphytewith adispersedpopulationstructure,thisspeciesisremarkablyuniform
invegetativeandreproductivecharacters.AnexceptiontothishomogeneityisexhibitedbytheCerro
Tute population wh—ich has uniformlyovate leaves (vs. elliptic leaves).
Etymology. Theepithetforthis species,arhoricola, isderivedfromtheLatin word, arbor,
meaning dwelling in a tree, in reference to its epiphytic habit in the forest canopy.
ALMEDA: TOPOBEA
101
t
—
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2. Topobea caliginosa Almeda, Proc. Calif. Acad. Sci. 46(14):317. 1990.
Blakeamicrantha Almeda, Rhodora 82:614. 1980.
— NW
Xype. PANAMA.Veraguas:CerroTuteca. 1 km ofSantaFeonridgetopincloudforest
above 1000 m, 19 June 1975, Mori 6765 (holotype: CAS!; isotype: MO!).
Rigidlybranchedterrestrialshrubto 1 mtall.Uppercaulineintemodesquadrangulartodistinctly
carinate. Young vegetative buds andjuvenile foliage commonly besetwith a sparse coppery brown
furfuraceousindument,otherwiseglabrousthroughout.Leavescoriaceousandglabrouswhendrybut
inconspicuously punctate abaxially, 1-4 x 0.4-1.6 cm, obovate to oblanceolate, apex obtuse to
rounded, base acute to attenuate, the margin entire, often revolute when dry, 3-nerved; petioles
0.5-1 cm long. Flowers solitary in the axils ofdistal branches, sessile, subsessile orwith outermost
mm
floral bractscompressedintoan ill-definedpeduncle3-5 long. Floralbractsroundedattheapex,
sparselyfloccosetoglabrousatmaturity,marginentire;outerbracts3-5 x 2-3 mm,fusedatthebase
for 1-2 mm, elliptic-lanceolate; innerbracts 3^ x 2-3 mm, free, elliptic-ovate. Calyx lobesbluntly
deltoid, 1.5 mmlongand2 mmwidebasally.Petals6,glabrousbutsparselyverrucoseabaxiallywhen
dry,4.5-6 x 2.5-3 mm,pink,ovatetoelliptic-ovate, apexobtusetorounded, entire. Stamens6,fila-
ments 2-3.5 X 0.5 mm, erect, glabrous; anthers free, 2 x 0.5-1 mm, linear-oblong and erect, each
with two confluent pores at the truncate apex; connective not thickened or appendaged dorsally.
Ovarycompletely inferior, 2-locular, glabrous atthe apex which is elevated into a shortcone. Style
glabrous, 5-6 mm long; stigmapunctiform. Berry4-5 x 3-5 mm. Seedsmostly 1 mm long,cuneate
to narrowlypyrifo—rm.
Phenology. FloweringspecimenswerecollectedinFebruary,April,andJune;fruitingspec-
imens collected in Ap—ril, June, and July.
Distribution. Western Panama where it is known only from Cerro Tute (Veraguas) at
m
1400-1453 (Fig. 3). —
Representative Specimens Examined. PANAMA. Veraguas: windswept summit of
Cerro Tute, 18 Feb. 1996, Almeda etal. 7627(CAS, MO, PMA); distrito de Santa Fe, Serrania de
Tute, 8°33'N, STOT'W, 5 Jul. 1996, Aranda et al. 2724 (CAS, SCZ); summit ofCerro Tute above
Escuela Agricola Alto de Piedra, just W of Santa Fe, 8°32'N, 81°07'W, 5 June 1982, Knapp &
Dressier5394 (CA—S, MO).
Discussion. This species was originallydescribed as an epiphytic shrub based on informa-
tionprovidedbythecollectorofthetypeandonlyknowncollection atthetime(Almeda 1980). Dur-
ingrecentfieldworkI havefound T. caliginosagrowingonlyasaterrestrial shrubabovetreelineon
the shrubby summit ofCerro Tute.
Three other congeners in this hexandrous group of species also occur on Cerro Tute T.
arboricola, T. cordata, andwhat maybe an undescribed taxon most closelyrelated to T. hexandra.
Thetwoformertaxaappeartobetrueepiphyticshrubsthatgrowontreeswithinthecloudforestzone.
The third entity, which grows in the upper zone just below tree line, is a vinelike secondary
hemiepiphyte that gemiinates terrestrially, ascends nearby trees by adventitious roots, and laterbe-
comes epiphytic by losing root contact with the ground. This sympatry and parapatry suggest that
strongisolatingmechanismsareoperatingtomaintainphenotypicdistinctionsamongfourcloselyre-
latedspecies.Thatthisalloccursonatropical mountainof1453 melevationmakesitall themorere-
markable.
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3. Topobeacordata Gleason, Phytologia 3:354. 1950.
TYPE. -PANAMA. Code: Cerro Pajita, north ofEl Valle de Anton, elev. 1000-1200 m,Allen
4178 (holotype: NY!; isotype: MO!)
Epiphytic shrub to 1 m tall. Uppermost intemodes ± compressed and quadrisulcate, the angles
distinctly carinate sometimes varying to narrowly winged; older branches ± rounded to
rounded-quadrate.Leavesofapairsessile,equaltoslightlyunequalinsize,theyoungeronesofapair
typically fused basally at the node into a narrow collar, glabrous throughout; mature blades
coriaceous, 5.1-12.9 cm long and 3-5.8 cm wide, cordate-clasping to subcordate or ovate, apex
acute,baseroundedtocordate,margin entire, 5-7-nerved,thesecondaryveinstypicallynotevident.
Flowers erect, coconut-scented (fide McPherson 8738), solitary or paired in each upper leafaxil;
peduncles 3^ mm long,conspicuouslycompressed. Floral bracts entire, glabrateorsparinglybeset
abaxially (mostly toward the apex) with a coppery brown lanate indument; outer bracts
0.8-1.1 X 0.6-0.9 cm, fused at the base for 2 mm, elliptic, apex acute to obtuse; inner bracts
mm
0.8-0.9 X 0.4-0.6 cm,freetothebase,oblong-ovate,apexrounded.Calyxtube 1-1.5 long;calyx
lobes(atanthesis)4.5 mmlongand3 mmwide,oblong-ovate,±entire,essentiallyglabrousbutsome-
times beset with a coppery brown lanate indument toward the apex on both surfaces. Petals 6,
0.8-1.3 X 0.4-0.7 cm, translucent-white, obovate, apex rounded, entire. Stamens 6, filaments
complanateandglabrous,6-7 mmlong;anthers5-7 mmlong, 1-1.5 mmwide,yellow,subulatewith
two broad confluent ventrally inclined apical pores; connective not conspicuously thickened and
unappendaged. Ovarycompletelyinferior,4-locular, apex glabrousandelevatedintoabluntbasally
swollen cone 2 mm long. Style somewhat declinate and sigmoid apically, glabrous, 8-9 mm long;
mm
stigmatruncate.Berry 1 x i cm,redatmaturity.Seedsnarrowlypyriformtocuneate, 1-1.5 long,
beige, the testa smo—oth and glossy.
Phenology. Flowering from Februarythrough May, Septemberandprobablyduring inter-
vening months; the o—nly fruiting specimen was collected in September.
Distribution. Wetcloud forestsofwesternPanama from theFortunaregion (Chiriqui)east
to Cerro Pajita (Code) at 1050-1250 m (Fig. 3)—.
Representative Specimens Examined. PANAMA. Chiriqui: Fortuna dam region, along
Quebrada Arena near continental divide, 8°45'N, 82°15'W, 9 Mar. 1986, McPherson 8738 (CAS,
MO);FortunacaucedeQuebradaArena,9Apr. 1987, Valdespinoetat. 639(CAS,PMA).Veraguas:
Cerro Tute, trail pa—st agricultural school near Santa Fe, 17 Sep. 1979, Antonio 1875 (CAS, MO).
Discussion. Thetype specimen ofthis species lacks petals and stamenswhich accounts for
the incomplete descriptions provided in theprotologue (Gleason 1950) and subsequenttreatmentof
thisspeciesintheFloraofPanama(Gleason 1958).Thecollectionoffloweringmaterialwithattached
stamens facilitated a critical assessment ofits generic placement and relationships to other species
with six antesepalous stamens (Almeda 1990).
Topobea cordata is characterized by a distinctive suite of characters including sessile
cordate-claspingleaves,oblong-ovatecalyx lobes,comparativelylarge(8-13 mm)translucentwhite
petals, and subulate anthers with ventrally inclined apical pores. Topobea cordata has a 4-locular
ovary like T crassifolia, but the latter has petiolate leaves, lance-triangular calyx lobes, smaller
(5-7 mm) pink petals, and short (2-2.5 mm) apically truncate anthers.
One enigmatic collection (McPherson 9864, CAS) from the Fortunaregion ofwestern Panama
resembles T cordatainitsoverallvegetativemorphologyandwhitepetals. Incalyxshapeandstami-
nal details, however, it is a good match for T crassifolia, the species to which T cordata is most
closelyrelated. Inthesizeofits floral bracts,hypanthia,andpetalsthiscollection alsoapproaches T
crassifolia. Thiskind ofcharactersorting is suggestive ofhybridization or introgression between T
cordataand T. crassifolia. Typical T. cordatahasalsobeen collectedat Fortuna. I havenotseen au-
ALMEDA: TOPOBEA 105
thenticmaterial ofT. crassifoliafromFortunabutitmaywelloccurtheresinceitsrangeextendsboth
east and west ofthatregion.
4. Topobea crassifolia (Almeda) Almeda, Proc. Calif. Acad. Sci. 46(14):318. 1990.
Blakeacrassifolia Almeda, Rhodora 82:612. 1980.
—
Type. PANAMA.Code:LaMesaaboveElValleinforestonbothsidesofjunctionwithroad
to Cerro Pilon, ca. 800 m, 21 Jul. 1974, Croat25430(holotype: CAS!; isotypes: MO!, US!).
m
Epiphytic shrubto 1 tall. Uppermostbranchesquadrangulartorounded, glabrousthroughout.
Leaves thick, succulent or semisucculent, coriaceous or chartaceous when dry,
1.5-6.5(-12) X 1-4.3 cm, ovateto elliptic-ovate orelliptic-lanceolate, apex acute to attenuate vary-
ingtocuspidateormucronate,baseroundedtocordate,themarginentire,3-5-nerved,oftenwithonly
the median nerve elevated and conspicuous abaxially, essentially glabrous throughout; petioles
0.1-0.4 cmlong.Flowerssolitary,pairedorinsubfasciculateclustersofthreetofiveinaxilsofdistal
mm
branches, sessile, subsessile or with ill-defined peduncles 1-2 long. Floral bracts glabrous or
sometimes beset with an inconspicuous lanate or fioccose indument distally, margin entire; outer
bracts 5-8 x 2.5^ mm, fusedatthebaseforabout 1 mm,elliptic-lanceolate,apexacutetoobtuseor
rarely rounded; inner bracts 4-7 x 2.5^ mm, free but closely subtending the hypanthium, ellip-
tic-ovate,apexacutetoroundedorbroadlytruncate.Calyxlobeslance-triangular,3^.5 mmlongand
mm
2-2.5 widebetweensinuses.Petals6,glabrousandcoarselyverrucose,sometimesfringedwitha
scatteringofminutemattedhairs, 5-7 x 1.5-2.5 mm,reportedlywhiteorpink, oblong-lanceolateto
narrowly oblanceolate, apex acute, base somewhat clawed, entire. Stamens 6, filaments
3.5-4.5 X 0.5 mm, erect, glabrous; anthers free, 2-2.5 x 0.5 mm, linear-oblong and erect distally,
each with two confluent dorsally inclined apical pores; connective thickened dorso-basally into a
spurlikeappendage.Ovarycompletelyinferior,4-locuIarandglabrousatthetruncateapex. Stylegla-
brous,6 mmlong;stigmapunctiform.Berry6x5 mm.Seedsmostly 1 mmlongorless,rangingfrom
ovoid orclavate to—lunate orpyriform.
Phenology. FloweringinFebruaryandMaythroughAugust; fruitingcollectionshavebeen
made in May and Au—gust.
Distribution. Cloud forests in central CostaRica from Braulio CarilloNational Park (San
Jose) andvicinitydisjunctto Panama from Cerro Colorado (Bocas del Toro)andEl Vallede Anton
(Code) to El Llano-Carti region (Comarca de S—an Bias) at 350-1750 m (Fig. 4).
RepresentativeSpecimensExamined. COSTA RICA. Alajuela/Herediaborder: Colonia
Virgen del Socorro alongroad from Costa Rica#9 to the Colonia, 2 Aug. 1976,J. & K. Utley5629
(CAS,DUKE,F). SanJose: ParqueNacionalBraulioCarillo.LaMontura,25-30Jul. 1982, Todziaet
al. 1964(NY). PANAMA. Bocasdel Toro/Chiriquiborder: CerroColorado,roadalongtop, 13 Aug.
1977,Folsometal. 4681(CAS).Code:LaMesa,4 kmNofElValledeAnton,6May 1981,Sytsmaet
al.4364(CAS,MO); alongtrailtoLaMesaabout4.5 milesbeyondElValledeAnton,21 May 1970,
Wilbur & Luteyn 11697 (CAS, DUKE, F, MO, US). Comarca de San Bias: El Llano-Carti road.
Nusagandi, 9 km from Interamerican Hwy, 9° 9'N, 78°55'W, 26 Aug. 984,deNevers & deLeon
1 1 1
3767(CAS,MO);ElLlano-Cartiroad, km 19.1,9°19'N,78°55'W, 1 Jul. 1985,deNevers5943(CAS,
MO); entrada a Nergan Igar, km 15 de la carretera Llano-Carti, 9°20'N, 78°58'W, 2 Jul. 1994,
Galdames etal. 12—43 (CAS, SCZ).
Discussion. Amongthehexandrousspecies, T. crassifoliaisthemostwidelydistributedand
hasthebroadestelevational amplitude. Likemanywoodyepiphytes,thefragmentedbutsmall popu-
lation structure ofthis species has evidentlypromotedmorphological divergence.
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