Novelties in Mesoamerican Mistletoes (Loranthaceae and Viscaceae)

Author(s): Job Kuijt

Source: Annals of the Missouri Botanical Garden, Vol. 74, No. 3 (1987), pp. 511-532
Published by: Missouri Botanical Garden Press
Stable URL: http://www.jstor.org/stable/2399319
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 NOVELTIES IN MESOAMERICAN MISTLETOES
(LORANTHACEAE AND VISCACEAE)'
JOB KUIJT2

ABSTRACT
Five newspeciesofLoranthaceae (Cladocoleaprimaria,Psittacanthus
angustifolius,
P. minor,P.
pinicola,and Struthanthus
subtilis)
and sevennewspeciesofViscaceae(Dendrophthora davidsei,D.
talamancana,Phoradendronfasciculatum, P. molinae,P. nitens,
P. tardispicum,
and P. zelayanum)
are describedfromMesoamerica. The new combinationPsittacanthusrhynchanthus (Bentham) Kuijt
(includingthevar. wurdackii(Rizzini) Kuijt) is proposed forwhathas usuallybeen called P. calyculatus
(DC.) G. Don in Mesoamerica. The lattername is now restrictedto a different species endemic to
Mexico.

The following novelties result from recent triadsbelow and a pair above, followedby a pair
studies in connectionwiththe Flora de Nicara- of ebracteolatemonads and a terminalflower,
gua and Flora Mesoamericana. the 4 basal triadscrowdedin the leaf axil. Flow-
ers bisexual, pale yellow, 4-partite;petals di-
1. Cladocolea primaria Kuijt, sp. nov. TYPE: morphic, 2-2.5 mm long; anthers very small,
Panama. Panamai: Cerro Jefe,2 km along sessile on the shorterpetals and withveryshort
road to Altos de Pacora fromjunction with filamentson thelongerones; ovary 1.5 x 1 mm;
road to peak, low cloud forest,800 m, Syts- stylemore or less straight,the capitate stigma
ma & Knapp 4797 (holotype,MO; isotype, reachingthe petal tips. Fruit 6 x 4 mm, red,
LEA). Figures 1, 2. becomingdark purple,ellipticin outline; calyc-
ulus inconspicuous;embryodicotylous,slender,
Plantaeglabrae,pauceramosae;ramiquasi teretes,
saepe lenticellis
insignibus ornati,rectisat rigidique. the haustorialpole scarcelyexpanded.
Folia bina,aliquantumcoriacea,laminaelatelanceo-
lataevel ovatae,obtusaevel leviterapiculatae;costa Additionalspecimensexamined. PANAMA. PANAMA:
insignis,petioliad 15 mmlongi.Florespallideflavi, CerroJefe, Clusiaforest
nearradiotower,900m,D 'Arcy
bisexuales.Inflorescentia solitaria,
determinata,subtus & Hamilton14817(LEA, MO); in forest nearroadto
triadis3- vel 4-paribus, supraparisingulo,tuncpari CerroJefenearjunctionwithroadtoAltosde Pacora,
unomonadarum ebracteolatarum etdeniquefloreter- Mori & Kallunki72763 (LEA, MO); CerroJefe,6.6
minalisequentibus; triadaebasales ad axillasaggre- mi. above GoofyLake,disturbed cloud forest,
850-
gatae; inflorescentiae paribus nonnullisfoliorum 900 m,Sytsmaetal. 2839 (LEA, MO).
squamiformium, crassorum, fuscorum suffultae.Flo-
res 4-partiti;petaladimorpha,2-2.5 mm longa;an- Cladocoleaprimaria presentsconsiderabledif-
theraeperparvae, filamentis ficultiesin genericassignment.When I mono-
sessilesad petalabreviora,
brevissimis insertaead longiora;ovarium1.5 x 1mm; graphedCladocolea (Kuijt, 1975), I proposedthe
stylusrectus,stigmacapitatum.Fructus6 x 4 mm, notionthatStruthanthus
ruber,obscureviolascens,ellipsoideus. is polyphyletic,at least
many species being derived froma number of
Plants sparsely branched, twining,glabrous. independentsources withinCladocolea. Thus I
Stems tereteor slightly4-ridged,oftenwithcon- spoke of connectingbridges,thesein some cases
spicuous lenticelswhenolder,straightand rather characterizedby species pairs, one member of
rigid,with occasional epicorticalroots. Leaves which was placed in Cladocolea, the other in
paired, somewhat leathery,the blades broadly Struthanthus. Withsome veryminorexceptions,
lanceolate to ovate,-venationinconspicuous,the thisleftCladocolea as a genus withdeterminate
apex mostlyblunt or slightlyapiculate; midrib spikesofmonads, theflowers4-, 5-, or 6-partite,
conspicuousand runningintoapex; petiolesstout, and eitherbisexualor thespeciesdioecious. Stru-
to 15 mm long. Inflorescencessubtendedby sev- thanthusremaineda strictly dioeciousgenus(with
eral pairs of thick,brown scale leaves, solitary the solitaryexception of the highlyaberrantS.
in leaf axils, determinate,with 3 or 4 pairs of panamensis, which has bisexual flowers and

1 Acknowledgmentsaredue to Dr. KarelU. Kramer,Zurich,fortheLatindescriptions,

and to theNatural
Sciencesand Engineering
ResearchCouncilofCanada forcontinuingfinancial
support.
2 DepartmentofBiologicalSciences,University
ofLethbridge,
Lethbridge,
AlbertaTIK 3M4, Canada.
ANN. MISSOURI BOT. GARD. 74: 511-532. 1987.

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512 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VOL. 74

cm

FIGURE 1. Cladocolea primaria Kuijt (Sytsma & Knapp 4797). Pendent branch.

bracteolatemonads), its inflorescencemade up colea, however,itis thefirstspecieswhichis truly

of mostlytriads.The inflorescenceof Struthan-
thusis generallyindeterminate,but in a few
species it also bears two or fourmonads at the
tip, followedby a trulyterminalflower.
Cladocoleaprimariapresentsa predicament
fromwhich thereis no completelysatisfactory
escape, forplacementin eithergenusleads to the
need of significantmodificationof that generic
concept. As a species of Struthanthus it would
be the firstfour-partitespecies, and the second
one with bisexual flowers.If placed in Clado-
triadic. I feel that placement in Cladocolea is
more acceptable,althoughI cannot deny an ele-
ment of arbitrarinessin this regard.I continue
to hold the view that union of the two genera
would tendto obscurethe complex relationships
betweenthem,and thatthesedifficulties concern
veryfewspecies. In fact,if the evolutionaryre-
lationshipsare as I have proposed, such difficult
intermediatespecies would be expected.
Viewed in the above context,C. primaria is
closely related to C. lenticellata (Diels) Kuijt

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1987] KUIJT-MESOAMERICAN MISTLETOES 513

mm

"/,

mm E
E

FIGuRE2. Cladocoleaprimaria Kuijt (Sytsma& Knapp 4797).-a. Inflorescence,

theuppertriadsremoved.-
b. Flower dissection.-c. Mature fruit.-d. Embryo.

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514 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VOL. 74

(which it greatlyresemblessuperficially)and C. species. The closest relativeof D. davidsei,how-

roraimensis(Steyerm.)Kuijt, while in Struthan- ever,is D. paucifolia(Rusby) Kuijt,whichranges
thusit is especiallyS. leptostachyus(Kunth) G. fromBolivia to Venezuela. The major difference
Don and S. polystachyus(Ruiz & Pavon) Blume betweenD. davidseiand D. paucifolia is thatthe
that are related. Our species representsthe sec- vegetativebranchesof the latterinvariablyhave
ond Cladocolea reportedforPanama and seems a prominentpair of basal cataphyllsplaced high
to be limitedto the Cerro Jefearea. above the leaf axil (see Fig. 39 in Kuijt, 1986a).
This is not seen in any of the ten individuals
presentin the two Davidse collections,but basal
2. Dendrophthoradavidsei Kuijt, sp. nov. TYPE:
cataphyllsare presenton some (not all) vegeta-
Costa Rica. Lim6n: Cordillerade Talaman-
tivelateralsoftheLent specimen.The latterlooks
ca, Atlantic slope, unnamed cordillerabe-
extremelyslenderbut appears to belong to the
tweentheRio Terbi and theRio Sini, 2,400-
presentspecies. A second and apparentlycon-
2,750 m, elfin forestedge, Davidse et al.
sistentdifference is thatleaf size in D. paucifolia
28990 (holotype,MO). Figure 3.
dwindles upwardlyuntil the uppermostlateral
Plantaad circ.15cmalta,monoeca,erecta, olivacea, spikes are subtendedby leaf scales; even though
parvifoliata;
cataphyllabasalianullavelirregularia,
ap- perhaps a slightdiminution takes place in D.
pendicesbasalesin planomedio.Folia (ob)lanceolata, davidsei,none of the leaves on the main stem
succulenta, ad 9 x 2 mm,acuta. Floresfemineiad
partessuperioresinternodium fertilium,plerumque ever reach scale size. The relationshipof thetwo
quam masculisubtusinsertihaudplures.Spicaesoli- species is certainlyveryclose, but separationap-
pedunculocirc.7 mm
tariae,axillaresterminalisque, pears justified.
longoinclusoad 5 cm longae;internodia 2 vel
fertilia Terminal inflorescenceswere not seen in the
ad 20 pro bracteafertili.
3, raro4; floresuniseriati,
Fructus ad globosus,1.5mmdiam.,
albus,ellipsoideus type but are representedin the otherwiseiden-
petalispatentibus. ticalcollectionDavidse et al. 28755a. In thetype,
as shown in Figure 3a, the tips of the larger
Plants monoecious, to ca. 15 cm high,erect, branches have remained static, giving the
olive green,small-leaved;basal cataphyllsabsent impressionof buds with crowded small leaves.
throughoutin some individuals,irregularly pres- Such shoot tips undoubtedlywill expand into
ent in others;basal appendages orientedin me- new systemsofinflorescences, and perhapsmark
dian plane; youngparts oftenwith sparse, stiff, a seasonal change.In fact,in thethirdcollection,
white bristlelikehairs. Leaves (ob)lanceolate, Lent 3826, thereare some brancheswheresuch
succulent,to 9 x 2 mm, acute. Spikes solitary a new expansion seems to have takenplace, de-
and axillary as well as terminal,to 5 cm long marcatedfromtheolderportionby severalpairs
includingthe peduncle ca. 7 mm long; fertile of verysmall leaves.
internodes2 or 3, rarely4. Flowers uniseriate,
to 20 per fertilebract;femaleflowersin theupper 3. Dendrophthoratalamancana Kuijt, sp. nov.
part of the fertileinternodes,not generallyout- TYPE: Costa Rica. Limon: Cordillerade Ta-
numberingthe male flowersbelow. Fruitwhite, lamanca, Atlantic slope, Valle de Silencio,
ellipsoid to globose, 1.5 mm diam.; petals along the Rio Terbi, 0.5-1.5 airline km W
spreading. of the Costa Rican-Panamanian border,
Additional specimens examined. COSTA RICA. 2,300-2,400 m, Davidse et al. 28755b (ho-
LIMON: Cordillera de Talamanca,Atlanticslope,Valle lotype,MO). Figure 4.
de Silencio,alongtheRio Terbi,0.5-1.5airlinekmW
oftheCostaRican-Panamanian border, 2,300-2,400 Plantatenuis,monoeca,erecta,ramosissima, circ.
m, Davidseet al. 28755a (MO). HEREDIA: open road 30 cmalta,ad basemsquamata;caulesinflorescentias
side,1 kmN ofSan Rafaelde VaraBlanca,1,900m, gerentes haud ultra1 mm crassi.Cataphyllabasalia
Lent3826 (F). omninocarentia, appendicesbasalespositionemedia.
Spicae solitariae,axillares,ad 15 mm longae,inter-
Dendrophthoradavidsei is the sixth known nodiofertili uno.Floresuniseriati, floribus
usquead 6
Mesoamerican species of subgenus Dendroph- supraquamquebracteamfertilem, masculiirregular-
thora,countingD. talamancana described con- iterdispersiet femineis multopauciores;bacca alba,
2 mmdiam.,ovoidea,sepalisclausis.
currentlyin this paper. The othersare D. gua-
temalensis Standley, D. mexicana Kuijt, D. Plants monoecious, ca. 30 cm high,profusely
squamigera (Benth.) Kuntze, and D. terminalis branched,slender,erect,squamate to the base.
Kuijt. No seriousconfusionis possiblewiththose Inflorescence-bearing stems 1 mm or less in

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 1987] KUIJT-MESOAMERICANMISTLETOES 515

1~~~~~

FIGURE 3. Dendrophthoradavidsei Kuijt.-a. Habit (Davidse et al. 28990).-b. Same collection,old inflo-
rescence.-c. Terminal portionof compound inflorescence(Davidse et al. 28 755a).

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516 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VOL. 74

X)~~~~~~~~~~~~~ -

talamancanaKuijt (Davidseetal. 28755b).-a. Habit.-b. Fruitinginflorescence.

FIGURE4. Dendrophthora

thickness;basal cataphyllsabsentthroughout, the ularlydistributedand much outnumberedby the

basal appendagesin a median orientation.Spikes
solitary,axillaryonly,to 15 mm long,the single
fertileinternodeslightlylongerthan the pedun-
cle, the apex acute. Flowersuniseriate,to 6 flow-
ers above each fertilebract; male flowersirreg-
femaleflowers.Berrywhite,2 mm diam., ovoid;
sepals closed.
This distinctive plant represents the third
completelysquamate Mesoamerican species of
theothersbeingD. squamigera
Dendrophthora,

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1987] KUIJT-MESOAMERICAN MISTLETOES 517

(Benth.)Kuntze and D. terminalisKuijt. It differs time whetherthis clusterof species is obligately

from the formerin its extremelyfine, much- hyperparasitic; plantsgrowingnear the base of a
branchedstemsand shortfertileinternodes,and primaryhost may easily be mistakenforbeing
fromthe latterin its size, bluntscale leaves, and parasiticdirectlyon the host tree. Anotherfea-
lack ofterminalspikes.It is conceivable thatthe tureapparentlyheld in common bythesevarious
unusually slender Friedrichsthals.n. specimen species is thatseveralstemsoriginatefroma bas-
fromGuatemala, as cited in Kuijt (1961), will al cushion, as illustratedin Figure 5a (arrow). I
turnout to be this species. I know of no similar add a comparable illustrationof a small plant of
South Americanplants. PhoradendrondipterumEichlerfromNicaragua
(Fig. 6), whichhappens to be parasiticon a leaf,
4. PhoradendronfasciculatumKuijt, sp. nov. the host again being a Phoradendron.A clear
TYPE: Panama. Chiriqui: JaramilloArriba,
basal cushionis visible (arrow).A thirdexample
near Boquete, trailto Rio Palo Alto, 1,100 is PhoradendronaequatorisUrban fromEcuador
m, near paved road, hyperparasiticon Pho- (Kuijt, 1986a, fig. 2), which also has basal
is parasitic on a Phoradendron.
radendronundulatumEichl., in turnpara- sproutingand
sitic on Psidium guajava, Churchill& Kuijt That thisis notaxillarybranchingfromthenodes
5106 (holotype, MO; isotypes, BM, CR, ofa much shortenedbase is demonstratedby the
EAP, MEXU, NY, LEA, PMA). Figure 5. usual lack ofbasal cataphylls,in contrastto what
occurs elsewherein the plant.
Plantaeerectae,glabrae,basi rami4-6 e pulvino The explanation of the basal cushion almost
communiorientes;internodiavetustiorabicarinata, certainlylies in theoriginalhaustorialdisk ofthe
novellapraecipuestatusicco leviterquadrangularia,
ad 10 cm longa,ramilateralesparibuscataphyllarumseedling. It has recentlybeen shown in an un-
singulis.Ramisolumnovellifoliosi.Foliacarnosa,8 x related species of Viscaceae, Viscum minimum
5 mm,apicerotundata, moxdecidua.Monoeca;spicae Harvey, that the marginof the haustorial disk
paribuscataphyllorum steriliumvel nullis;internodia regularlyproduces aerial shoots (Kuijt, 1986b).
fertilia5 vel 6. Floresmasculi1-3 ad apicemareae The same is truefor Viscumalbum L.
floriferae (Tubeuf,
suprabracteamfertilem, femineiusque ad
12probracteaetiis suppositae, Fruc- 1923), and for Ixocactus hutchisoniiKuijt of
bi-veltriseriati.
tuslate ovoideus,3 mm diam.;petalisinconspicuis, Loranthaceae(Kuijt, 1987). In Phoradendron, this
clausis. featurewould appear to have some taxonomic
Plants monoecious, erect,glabrous, fascicled constancyin the group of species under discus-
sion.
fromthe base with 4-6 stems froma common
cushion. Stems usually lackingbasal cataphylls; Additional specimen examined. COLOMBIA.
internodes 2-keeled when older, somewhat ANTIOQUIA:
highwaybetweenUramita and Canfasgor-
quadrangularwhen young,especiallywhen dry, das,on Phoradendron
piperoides
(H.B.K.) Trel.,Bark-
ley& Gutierrez
535457(US).
to 10 cm long,stout;lateralbrancheswith 1 pair
ofbasal cataphyllsca. 4 mm above base, spread- 5. Phoradendronmolinae Kuijt, sp. nov. TYPE:
ingwhendry.Leaves fleshy,soon deciduous,8 x Nicaragua.Madriz: cutovercloud forestarea
5 mm, the apex rounded,sides parallel,the base on Volca'n Somoto, 10 km S of Somoto,
clasping.Spikes with 1 pair of sterilecataphylls 1,400 m, Williams & Molina 20270 (holo-
or without,5-6 fertileinternodessometimespro- type,US; isotype,F). Figure 7.
terminallyintoa second seriesofyoun-
liferating
gerfertileinternodes.Male flowers1-3 at thetip Internodia compressacarinataque, ad 6 cm longa;
of the flowerarea above each fertilebract, the ramilateralesparibuscataphyllorum basaliumvalde
inferissingulis.Folia ad 10 x 4.5 cm; laminatenuis,
female flowersto 12 per bract below them,bi- ovata,basi in petioluminsignem, cuneiformem, fere
or triseriate.Fruitbroadly ovoid, 3 mm diam., ad 1.5 cm longumabruptecontracta.Inflorescentia
thepetalsverysmall,inconspicuous,moreor less femineasolitaria,paricataphyllorum steriliumprae-
closed. sentevel carente;pedunculusstatufructifero circ.3
mmlongus;spicafructifera circ.3 cm longa,interno-
This distinctivespecies undoubtedlybelongs 2 vel 3, floribus
diisfertilibus probracteafertilitribus,
to the P. dipterumgroup of species, in which insertis
paulumsupramediuminternodii. Fructus ovo-
ideus,laevis,3 x 2 mm,petalaclausa.
hyperparasitismis the rule. Phoradendronfas-
ciculatumis no exception,in thatall plants seen Plants dioecious (onlythe femaleseen), stems
of the type collection were growingon P. un- withcompressed,keeledinternodesto 6 cm long,
dulatum Eichler. It is impossible to tell at this basal cataphylls one very low pair on lateral

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518 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VOL. 74

cm

FIGURE 5. Phoradendron fasciculatum & Kuijt5106).-a. Base of youngplant,showing

Kuijt (Churchill
sproutingfrombasal cushion(arrow).
-b. Habitof olderplant.- c. Olderinflorescence,
thebasal portionin
theterminal
fruit, portionproliferated
and in flower.

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 1987] KUIJT-MESOAMERICAN MISTLETOES 519

mm

FIGURE 6. Phoradendron
dipterum
Eichlerparasiticon Phoradendron
sp., Nicaragua(Stevens& Montiel
17931, LEA). The hyperparasiteis sproutingfroma basal cushion (arrow)attachedto a leafof the primaryhost.

branches. Leaves to 10 x 4.5 cm; blade thin, divide between Tres Rios and Cartago, on
more or less palmatelyveined, ovate, the base Euphorbiaceae, Kuijt 2465 (holotype,CR;
abruptlycontractedintoconspicuous,cuneiform isotype,UBC). Figure19 in Kuijt (1964) and
petiole to nearly 1.5 cm long. Female inflores- Figure 5 in Kuijt (1986a).
cence solitary,less than 2 cm long, oftenwitha
sterilepair ofcataphylls;peduncleca. 3 mm long Plantamagna,monoeca,carnosa, ramificatione
saepe
in fruit;fruitingspike ca. 3 cm long, with 2-3 furcata;innovationeslateralescataphyllisbasalibus
fertileinternodesand 3 flowersper fertilebract magnisbinis.Ramulinovellialiquantumcompressi,
just above the middle of the internode. Fruit demumteretes. Folia usque ad 15 x 7 cm vel ultra,
statuvivo crassaet nitentia,rigida,obovatavel fere
ovoid, smooth,3 x 2 mm; petals closed. elliptica,
basiangustata velabruptecontracta;petiolus
validissimus,planus,ad 6 mm latus.Floresmasculi
6. Phoradendron nitens Kuijt, sp. nov. TYPE: rarissimi,inferiores
seriatimdispositi;spicaad 4 cm
Costa Rica. Cartago:east side ofcontinental longa,cataphyllis
nullis,internodiafertilia
3 vel4, flo-

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520 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VOL. 74

inflorescence.-
FIGURE7. Phoradendronmolinae Kuijt(Williams & Molina 20270).-a. Habit.-b. Fruiting
c. Basal cataphylls.

Fructusflavus,3 x 1.5
resprobractea6-9, biseriati. a large pair of basal cataphyllsjust above the
petalaclausa.
mm,ellipsoideus;
axil; stems somewhatcompressed when young,
Plants fleshy, monoecious, the branching becomingterete.Leaves to 15 x 7 cm or larger,
mostlyforked,wherepercurrentperhaps witha thick and shiny when fresh,rigid, obovate to
pair ofintercalarycataphylls.Lateral shootswith nearlyelliptical;base taperingor abruptlycon-

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 1987] KUIJT-MESOAMERICAN
tracted;petioleverystoutand flat,to 6 mm wide.
Spike to 4 cm long, lacking cataphylls,fertile
internodes3-4, with flowers6-9 per bract,bi-
seriate;male flowersveryrare,lowest in series.
Fruityellow,3 x 1.5 mm,ellipsoid;petalsclosed.
Additionalspecimensexamined. NICARAGUA.
ZELAYA: Municipiode Siuna,Rio El Bambfx, arriba,
200-400m,Ortiz1665(HNMN, MO); Rosa Grande,
Rio Labfx(rio abajo), 300 m, Ortiz1031 (HNMN,
MO); CerroWaylawas(PefiasBlancas),5 kmal S de
Wany,costadooestedelcerro,on Crescentia,Grijalva
& Burgos1665 (HNMN, MO); Municipiode Rama,
caminodesde"SantaJulia"hasta"La Palmera,"60-
95 m,Robleto680 (HNMN, MO). (Alsosee thespec-
imenslistedunderPhoradendron obliquum(Presl)Ei-
chlerin Kuijt,1964).
This species, which in Mesoamerica has been
erroneouslyreferredto Phoradendronobliquum
(Presl) Eichler,is oftenextremelydifficult
to sep-
arate fromvigorous specimens of P. robustissi-
mumEichlerifonlyherbariummaterialis avail-
able. In the freshcondition, its usually larger
shinyleaves and yellowishberriescontrastsharply
with those of the latter species. Phoradendron
robustissimumis also strictlydioecious, but the
rarityof male flowersin P. nitenscan be very
misleading.Phoradendronobliquumis presently
placed in synonymyunder Dendrophthoraobli-
qua (Presl) Wiens (Kuijt, 1986a).
The speciesis now knownfromPanama, Costa
Rica (Kuijt, 1964), and Nicaragua, and has been
reportedrecentlyfromEcuador (Kuijt, 1986a, as
Phoradendron#5,fig.5). We may thus antici-
pate thatit will be foundin Colombia.
7. Phoradendron tardispicum Kuijt, sp. nov.
TYPE: Panama. Chiriqui:bridgeover Rio San
Felix,on Panama Hwy.,50 m,on Ficus along
riverdirectlyS of bridge,Churchill& Kuijt
5107 (holotype,MO; isotypes,BM, LEA,
MEXU, PMA). Figure 8.
Plantaedioeca (pistillatae solumvisae),vivae ob-
scurevirides, glabrae.Caulesdeindeteretes;internodia 8. PhoradendronzelayanumKuijt,sp. nov. TYPE:
recta,5-9 cmlonga.Cataphylla basaliaplerumque par
unumvelpariadua,valdeinconspicua. Cataphylla in-
tercalariapraesentia, paria singulaparibusfoliorum
interspersasedad caulesirregulariter ad in-
distributa,
ternodianonnullacarentia.Folia amplexicaulia, cor-
data,viva nitida,usquead 7.5 x 6 cm,margineun-
dulata,venationepinnata.Inflorescentiae satgraciles,
6-8 cm longae,fereomnesad internodia vetustiora
defoliata,
aliquotpronodo,spicaquaqueparibusduo-
bus cataphyllorum sterilium basaliuminternodiisque teretes; innovationes
fertilibus
circ.7; flores ad 10probracteafer- binis.Folia lateovatavel orbicularia,
biseriati,
tili.Fructusovoideus,luteo-viridis,3 mmlongus,pe- 8 cm,palmato-venosa;
talisclausis.
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MISTLETOES 521
Plantsdioecious (onlythefemaleseen),bright,
dark greenwhen fresh,glabrous, 1-2 m diam.,
somewhat pendulous. Upper part of internode
slightlyflattenedwhen young, soon becoming
terete;internodesstraight,5-9 cm long. Basal
cataphyllsmostly 1 or 2 pairs, very inconspic-
uous, if 1 pair presentnearlyaxillary,if 2 pairs
presentthe second pair to 20 mm above axil,
rarelyto 4 pairs. Intercalarycataphyllspresent,
one pairbetweensuccessivepairsoffoliageleaves
but irregularly distributedalong the branch,ab-
sent from some internodes,always inconspic-
uous. Leaves amplexicaul, cordate, shinywhen
fresh,to 7.5 x 6 cm, ratherthin; marginundu-
late; venation pinnate but obscure. Inflores-
cences ratherslender,6-8 cm long,nearlyall on
older,leaflessinternodes,severalpernode, rarely
one in axils of intercalarycataphylls;spike with
2 pairs of sterilebasal cataphyllsless than 5 mm
above base, followedby ca. 7 fertileinternodes.
Flowersbiseriate,to 10 per fertilebract,yellow-
ish green,each fertileinternodewith stalk and
flowerlesstip 2-4 mm long. Fruitovoid, yellow-
ish green,3 mm long,the petals closed.
This is a remarkablespecies forits irregularly
distributedintercalarycataphyllsand forlate de-
velopmentof inflorescences.It is difficultto see
what known species mightbe related.As faras
I am aware, only in Phoradendronparadoxum
Urban fromVenezuela do intercalarycataphylls
alternatein occurrence,but there this pattern
seemsto be regular(Trelease, 1916). That species
and the closely relatedP. fendlerianumEichler,
however,have triseriateflowersand long-peti-
oled leaves and do not seem closely related. In
the newlydescribedP. balsleviiKuijt fromEc-
uador (Kuijt, 1986a), a similarirregularity
exists,
but there0-3 cataphyllsmay be presenton an
"internode"; nor does this species seem to be
closelyrelatedto P. tardispicum.
Nicaragua. Zelaya: N of abandoned airstrip
near Alamikamba, along tributaryof Cafio
Alamikamba, 10 m, galleryforestamong sa-
vanna, on Symphonia globuliferaL.f., Ste-
vens 21717 (holotype, MO; isotypes,
HNMN, LEA). Figure 9.
Plantamonoeca,dichotoma, apiceabortivo.Caules
lateralescataphyllis
infrapositis
laminaad 8 x
petiolusvalidus,planus,supra
expansus.Inflorescentia
cataphyllissterilibus
nullis;in-
522 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VOL. 74

b 4
IE JP
FIGURE 8. Phoradendron
tardispicum & Kuijt5107).-a. Habit.-b. Inflorescence.
Kuijt(Churchill

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1987] KUIJT-MESOAMERICAN MISTLETOES 523

cm

FIGURE9. Phoradendron
zelayanumKuijt (Stevens21717). -a. Habit. -b. Inflorescence.

tria;flores13-15 pro bracteafertili, 9. Psittacanthus angustifoliusKuijt, sp. nov.

ternodiafertilia
spica4 cm longa;spicaehermaphroditae.
biseriati; TYPE: Nicaragua. Madriz: 0.5 km al S de San
Jbosede Cusmapa, I ,200 m, parasitandoen
Plants monoecious,forking, theapex aborting,
un Pinus, Moreno 24419 (holotype, MO;
theinflorescencesand youngshootsdull golden-
isotype,HNMN, LEA). Figure 10.
yellow. Stems terete,stout; internodesto 8 cm
long;thelateralshootswithone low pair ofcata- Caulesacutequadrangulares. Foliabina,anguste fal-
phylls.Leaves broadlyovate to orbicular;blade cata,tenua,venatione pinnata;laminaad 17 x 2.5 cm,
basiacuta,apicegraciliter attenuata,petiolusad 5 mm
to 8 x 8 cm, with 5 or 7 veryconspicuous pal- longus.Inflorescentia terminalis,e triadis4 vel 6 con-
mate veins runningfartowardsthe apex; petiole sistens. triaderum
Pedunculi circ.1cmlongi,inferiores
stout,ca. 8 mm long,flatand expandingdistally. bracteisfoliaceisad 2 cm longis;pedicellicupulater-
Inflorescences bisexual, lacking sterile cata- minalinotabili.Petalaaurantiaca, 7.5-8cmlonga;ala-
phylls,thepeduncle 3 mm long,thisfollowedby bastrumrectumvel aliquantumcurvatum. Ligulaeet
pili staminum nulli,sed apicespetalorum cristacen-
3 fertileinternodeseach with 13-15 biseriate trali,carnosaliguliformi ornati.Staminadimorpha, 5
flowersper fertilebract, the entire spike 4 cm cm longa,antherae6 mmlongae.Stigmaantheras su-
long. perans,capitatum.

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524 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VOL. 74

FIGURE10. Psittacanthus (Moreno24419) -a. Habit.-b. Tip ofpetal.

angustifolius

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1987] KUIJT-MESOAMERICAN MISTLETOES 525

Stems sharply quadrangular. Leaves paired, straight,not dilated at base. Petals ca. 3.7 cm
narrowlyfalcate,thin;the blade to 17 x 2.5 cm, long, red-orange;ligular area weakly differen-
venation pinnate,the base acute, the apex slen- tiated. Stamens dimorphic; the anthers 3 mm
derlyattenuate;petiole to 5 mm long. Inflores- long, dorsifixed,the 2 series scarcely overlap-
cences terminal,consistingof 4 or 6 triads;triad ping, the filamentsattached ca. 21 mm above
peduncles ca. 1 cm long, the lowest ones with the base, some 16 mm long. Ovary more or less
foliaceous bractsto 2 cm long; pedicels 1.5 cm cylindrical,4 x 2 mm; stylenearlyas longas the
long, withconspicuous terminalcupule, the ca- petals; stigma weakly differentiated; calyculus
lyculussmooth. Bud stout,more or less straight smooth. Fruits ovoid, 1.5 x 1 cm, with con-
or somewhatcurved; base and tip 5 and 4 mm spicuous calyculus,blackish.
wide,respectively, thelatterblunt.Petals orange,
Additionalspecimensexamined. NICARAGUA. MA-
7.5-8 cm long; innerpart of flowerhairless,the TAGALPA: PuertasViejas, 2 km al N sobre la Carretera
petalbase 5 mm wide,withoutligule;petalapices Panamericana,"San Vicente,"600 m, on Acacia, Mor-
4 mm wide, blunt,each witha fleshy,ligule-like eno 18288 (HNMN, LEA, MO); same, San Juanillo,8
median crest extendinginwards. Stamens di- km al SE de Ciudad Dario, 500 m, on Fabaceae, Gri-
morphic;filamentsattachedat ca. 2.5 cm above jalva 2618 (HNMN, LEA, MO); same, Entrada Paso
de Carretera,quebrada, 460-480 m, on Bignoniaceae,
petal base, 5 cm long; anthers6 mm long. Ovary Moreno 16698 (HNMN, LEA, MO). MANAGUA: cam-
5 x 6.5 mm. Stigmaplaced above anthers,very ino Dario-Presa Santa Barbara,8 km al NW de Ciudad
small, capitate. Fruitunknown. Dario, on Leguminosae,Grijalva2693 (HNMN, LEA,
MO).
Additional specimenexamined.Same dataas type,
Soza et al. 155 (HNMN, MO). Psittacanthus minor is closely related to P.
mayanusStandley& Steyerm.,whichappears to
This is an extremelydistinctivespecies,known
be limitedto the Yucata'n region,has quadran-
from what is essentially a single collection.
gular stems,and fruitswhich are about half as
Whetherthe species is restrictedto Pinus, as ap-
large as those of P. minor. Psittacanthus ma-
parentlyis Psittacanthuspinicola, can only be yanus
is much smallerin general.
shownby further fieldwork.The two species can-
not be confused,as P. pinicola is dyadic and has 11. Psittacanthuspinicola Kuijt, sp. nov. TYPE:
irregularlywhorled,blunt,more leatheryleaves. Belize. Belize: Manatee Pine Ridge, on pine,
The type locality suggeststhat the species may 1931-32, Gentle82 (holotype,GH; isotype,
well be presentin neighboringHonduras. MO). Figures 12, 13.

10. Psittacanthusminor Kuijt, sp. nov. TYPE: Caules plus minusve teretes;folia symmetrica,ter
verticillata,ad 11 x 2.5 cm, angusteellipticavel lan-
Nicaragua. Matagalpa: SW slopes of Cerro ceolata, apice rotundata,basi in petiolum circ. 5 mm
El Pilon and adjacent Laguna Tecomapa, longum angustata.Inflorescentiaelaterales,ad nodos,
roadside, low thornscrub and pastures on umbellulas e dyadis 2 vel 3 formantes.Petala circ. 4
rocky slopes, on ant acacia, Stevens 9466 cm longa, rubra,apice luteo-viridescentia,medio au-
(holotype, MO; isotypes, HNMN, LEA). rantiaca,basi ligula prominente;alabastrumsupra ali-
quantum curvatum,circum ovarium ad latitudinem
Figure 11. circ. 5 mm dilatatum,ad apicem tenuissimum,leviter
curvatum,circ. 1.5 mm latum angustatum;calyculus
Plantaeparva,caulibusteretibus, foliisbinis.Folia laevis. Stamina dimorpha; antheraedorsifixae,dorso
tenua;laminaad 5.5 x 3.5 cm,ovata,utraqueextre- paulo pilosae. Ovarium plus minusvecylindricum,4.5
mitateobtusavel fere;petiolus3-5 mmlongus.Inflo- mm longum; styluslongus, basi aliquantum torsus;
rescentiaterminalis,
paribustriadarum4-6. Petalacirc. stigmaellipsoideum,subtiliterpapillosum. Fructusel-
3.7 cm longa,rubro-aurantiaca; area ligulaepaulum lipsoideus,calyculoinconspicuo, 13 x 5 mm, saturate
distincta.
Alabastrum rectum,basihauddilatatum, ca- purpureus.
lyculuslaevis.Staminadimorpha; antherae 3 mmlon-
gae,seriesduaeviximbricantes; filamentacirc.16mm Stems more or less terete,becoming coarsely
longa;styluspetalisfereaequilongus;
stigmaaegredis-
fissuredand blackish when old; leaves symmet-
tinguendum.
rical, in (oftensomewhatirregular)whorlsof 3,
Stems terete,phyllotaxypaired. Leaves thin, to 11 x 2.5 cm, narrowlyellipticalto lanceolate;
the blade to 5.5 x 3.5 cm, ovate, the apex and apex rounded; base taperinginto petiole ca. 5
base obtuse or nearlyso; venation more or less mm long. Inflorescenceslateral, axillary,often
palmate;petiole3-5 mm long.Inflorescencester- also on older,leaflessstems,each beingan umbel
minal, consistingof 4-6 pairs of triads on pe- of2 or 3 dyads;inflorescencepeduncleto 13 mm
duncles ca. 12 mm; pedicels 10 mm long. Buds long;dyad pedunclesand floralpedicels 5-7 mm

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526 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VOL. 74

17~~~~~/
b c

SI
...4 ,:.
....

FIGURE11. Psittacanthus minorKuijt(Stevens9466).-a. Habit,withimmature

inflorescence.-b.
Mature
bud.-c. Base ofpetal.-d. Fruit(Moreno18288).

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 /
C

U~~~~~~~E

I~~~~

E~~~~~~~~~~~

N)~

FIGURE 12. Psittacanthus pinicolaKuijt.-a. Habitwithimmature (Pipoly4013).-b. Same

inflorescences
collection,
portionofinflorescencewithmaturebuds,a singlepetalshownseparately.
- c. Completeinflorescence
justafteranthesis(Gentle82).-d. Maturefruit (Stevens7600).
527

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 a

E~~~

E~~~~~~~~

E~~~~~~~~~~~

ic f

pinicola(Gentle82).-a. Leafyinnovation.-b. thestyleandanther

Matureflower,
FIGURE 13. Psittacanthus
shown separatelyto the right.-c. Relative positions of anthersand style.
528

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 1987] KUIJT-MESOAMERICAN MISTLETOES 529

long,the latterscarcelyexpanded at the tip. Pet- Syst.3: 415. 1834. TYPE: Mexico.Cuemavaca:
als ca. 4 cm long,red withyellow-greentip, or- Berlandier1150 (G-DC).
ange in the middle, prominentlyligulateat the An attractivePsittacanthusin whichthe buds
base. Buds somewhat curved above, inflatedat are distinctively curved and beaked occurs
the ovary to a width of nearly 5 mm, tapering throughoutMesoamerica,at low elevationsfrom
to a veryslender,slightlycurvedtip ca. 1.5 mm southernMexico to Venezuela. In the past, this
wide. Stamensdimorphic,sometimesapparently species has been called P. calyculatus(DC.) G.
nearly trimorphic(Fig. 13c); filamentsof the Don or,earlier,P. chrismardi Urban. Afterstudy-
longertypeattached 1.5 cm above the base, 1.5 ing the types of both Loranthus rhynchanthus
cm long; anthers 3-4 mm long, dorsifixed, Benthamand L. calyculatusDC., I conclude that
sparselypubescenton back. Ovary more or less theseare two distinctspecies. Consequently,the
cylindrical,green, 4.5 mm long, 2 mm diam. name P. rhynchanthus must be applied to the
below, expanding slightly above; calyculus wide-rangingspecies mentionedabove. True P.
smooth. Style4.6 cm long more or less straight, calyculatus seems limited to Mexico, the type
but thebase somewhattwisted;stigmaellipsoid, originatingfromthe area of Cuernavaca, further
finelypapillate. Fruit ellipsoid, with inconspic- collections having been seen fromPuebla and
uous calyculus, 13 x 5 mm, "deep purple." Morelia.
Additionalspecimensexamined. BELIZE. BELIZE: Notwithstandingtheirgeneral similarity,the
westernhighway,Mile 30, beside track,on pine, White- two speciesmaybe consistently separatedmostly
foord2442(MO); same,ThePlace,onpine,Whitefoordon the basis of floralfeatures.The mature,un-
2562 (MO). DISTR. UNKNowN: P. Cayo, in roadside opened bud of Psittacanthuscalyculatusis very
park on ridgearea overlooking1,000 ft.falls,in area
ofPinuscaribea,MountainPine Ridgearea,Huston nearlystraightand has a ratherblunttip; thatof
615 (MO). NICARAGUA. ZELAYA: Rio Tronceraatjunc- P. rhynchanthus shows a distinctivecurvaturein
tion with carreterabetweenWaspam and Puerto Ca- the distal portion,the apex being sharplyacute,
bezas, elevation less than 200 m, poorly developed and more beaklike. Psittacanthusrhynchanthus
galleryforestinsavanna,onPinuscaribea,Pipoly4013 has -smoothpollen
sacs behind which are borne
(HNMN, LEA, MO); nearTala Has and PuenteMango
(over Rio Kisalaya), 40-60 m, pine savanna, on Pinus long,conspicuous,reddishstamenhairs;thepol-
caribea,Stevens7600(LEA, MO); Comarcadel Cabo, len sacs ofP. calyculatusare distinctlylobed, and
Kornuk Creek above Puente Pozo Azul, old bridge, stamenhairsare lacking.Furthermore, thestylar
Robbins5831 (LEA, MO). NUEVA SEGOVIA: El Jicaro, base in P. rhynchanthus bearslow protuberances,
7 km sobre la carreteraa Murra entradaal Quebracho,
600-620 m, on pine, Moreno 8305 (HNMN, LEA, and each adjacent petal base shows a ligulecon-
MO). sistingof a low, V-shaped ridge;the stylarbase
in P. calyculatusis smooth, and ligules are ab-
Other dyadic species northof Panama are P. sent.Leaves of P. calyculatustend to be smaller
sonorae (Watson) Kuijt, P. palmeri (Watson) (to 8 x 4 cm), mostlyless than twice as
long as
Barlow' & Wiens, P. nudus (Molina) Kuijt & wide, and approximately symmetrical,while
Feuer, and P. ramiflorus(DC.) G. Don. Psitta- those of P. rhynchanthus are usually larger(to
canthuspinicola is similarto the last species but 12 x 4 cm), more than twice as long as
wide,
seems more closelyrelatedto P. dichrous(Mar- and strikingly falcate.
tius)Martius(see Eichler,1868, especiallyP1. 5). In Venezuela, at least some individuals of the
Not onlyinflorescencestructureand generalap- latterspecieshave extremely narrowleaves; these
pearance indicate this affinity, but even-thepe- plantsbelongto Psittacanthusrhynchanthus var.
culiarcurvatureof thestylarbase is seen in both. wurdackii(Rizzini)
Kuijt, comb. nov. (P. calyc-
ulatus(DC.) G. Don var. wurdackiiRizzini,Rod-
12. Psittacanthus rhynchanthus (Bentham) riguesia41: 15.
1976). I have notyetencountered
Kuijt, comb. nov. Loranthus rhynchanthus the species fromthe Caribbean lowlands of Co-
Bentham,Bot. Voy. Sulphur102-103. 1845. lombia, but it would be
surprisingifit were not
TYPE: "Dr. Sinclair," Tiger Island (Hondu-
presentthere.
ras, Bay of Fonseca) (K). Figure 14d-f.
Psittacanthus
chrismarii Urban,Bot. Jahrb.24: 331. 13. Struthanthussubtilis Kuijt, sp. nov. TYPE:
1897.TYPE (here designated):Costa Rica. Forets
de Nicoya, Tonduz 13706 (hololectotype,US; iso- Panama. Cocle: nearcontinentaldividealong
lectotypes,CR, GH). lumberingroad, 2.2 km beyond sawmill in
Psittacanthus
calyculatus
auct.,non(DC.) G. Don,Gen. forestabove El Cope, 900 m, Hammel 998

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530 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VOL. 74

a d

E E

LEA).-a, d. Maturebuds.-b, e. Ptals.-c,f. Sty basee

C~~~~ f
E E~~~~~~~~~~~~~~
E E~~~~~~

FIGURE 14. Psittacanthus (DC.) G. Don (a-c; Mexico, Puebla, slopes of Popocatepeti, Ugentet
calyculatus
aL.1359,-WIS) and P. rhynchanthus
(Bentham)Kuijt (d-f; Nicaragua,Managua, Cuatro Esquinas, Moreno4400,
LEA). -a, d. Mature buds. -b, e. Petals. c f. Stylarbases.

(holotype,MO; isotypps,LEA, PMA). Fig- laminaangustelanceolataad lateovata,4-12cmlonga,

ure 15. 2-6 cm lata,apice caudiformis, petioluscirc.3 x 1
mm. Inflorescentiaesingulae,axillares,bracteistria-
Plantae inconspicuae, subtiles; caulis gracilis,plus darum2 vel 4 constitutentes;pedunculuscommunis
minusveteres.Folia tenua;venatiomanifesta,pinnata; ad 8 mmlongus,triadarum ad 5 mm,uterque0.5 mm

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 cm

E
E

E~~~~~~~~~

E~~~~
e
~~~~~~1 ~ ~ ~ 3

FiGuE 15. Struthanthus subtilis

Kuijt.-a. Leafyshoot(Croat49004) and leaves(Croat44584).-b. Inflo-
rescence(Folsom& Lantz 1894).- c. Samecollection, - d. Samecollection,
petalsand style,maleflower. petals
- e. Maturefruits
and style,femaleflower. (Hammel998).

531

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532 ANNALS OF THE MISSOURI BOTANICAL GARDEN
velminuscrassus;flores anthesipedicellis0.5
laterales
mmlongis;bracteaebracterolaequeminutae,caducae;
petala2-2.5 mmlonga;stigmacristisindistinctispa-
subglobosus,5 mm
pillosis6. Fructusaurantiacus,
pedicellifructuum
diam.,calyculoinconspicuo; later-
aliumad 3 mmelongati.
Plantsinconspicuous,ratherdelicate,branched.
Stems slender,more or less terete;stem roots
occasional,thin.Leaves thin,blade narrowlylan-
ceolateto broadlyovate, alwayswithcaudate tip,
2-6 x 4-12 cm, the evident venation pinnate;
petiole ca. 3 x 1 mm. Inflorescencespale green,
solitaryin leaf axils, subtended by 4 minute,
probablycaducous bracts,consistingofa raceme
of 2 or 4 triads;inflorescencepeduncle to 8 mm
long; triad peduncles to 5 mm, both 0.5 mm or
less thick. Lateral flowerson pedicels 0.5 mm
long at anthesis; bracts and bracteoles minute,
caducous. Petals 2-2.5 mm long. Stamens di-
morphic; upper portion of sterile stamens pa-
pillate; anthers0.4 mm long; style2 mm long,
thestigmawith6 indistinctpapillatecrests.Friits
orange,nearlyspherical,5 mm diam.; calyculus
inconspicuous;pedicelsoflateralfruitselongated
to 3 mm.
Additionalspecimens examined. PANAMA.COCLE:
nearcontinentaldividealonglumberingroad,1.5 mi.
N ofEl Cope,Croat44584-(CR,LEA, MO); El Cope-
El Potrosa,AtlanticslopeofAltoCalvario,700-850
m,Folsom & Lantz 1894 (LEA, MO, PMA); alongroad
fromLa Pintedato El CopebywayofPiedrasGordas,
sawmillaboveEl Cope,cloudforest, 100m,Hammel
2640 (LEA, MO); alongroadbetweenLlano Grande
andCoclesito(N ofPintada),4 mi.N ofLlanoGrande,
600 m, Antonio3575 (LEA, MO); trailfromCanio
Blancodel Norteto continentaldivideN ofEl Cope,
on Hedyosmum,400 m, Davidse & Hamilton 23654
(BM,LEA,MEXU,MO, PMA);El Cope,Atlantic side,
1,200m, Antonio1153 (LEA, MO); betweenconti-
nentaldivideabove El Cope and El Potrososawmill
andtheRio Blancoto thenorth,330 m,Sytsmaetal.
2409 (EAP, LEA, MO); El Cope,alonggravelroadto
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[VOL. 74
therightbeforesawmill,800 m,Antonio2207 (LEA,
MO, NY, PMA); AltoCalvariocloudforest, 5.3 km
aboveEl Cope,continental
divide,abovesawmill,930
m,Antonio3044 (LEA,MO, PMA); aboveEl Potroso
sawmillat continental
divide,1,200-1,300m,Sytsma
1820 (LEA, MO). vEiRAGuAs:alongSantaF-Calove-
boraroadbeyondEscuelaAgricolaAltoPiedra,along
majorstream,3 mi. fromforkin roadat school,
first
700 m, Croat 49004 (LEA, MO).
This species was previously listed as S. aff.
dichotrianthusEichler (Kuijt, 1978). Struthan-
thusdichotrianthus(andS.phyllyraeoides(Kunth)
Blume) are indeed relatedto S. subtilis,as is S.
quercicola(Cham. & Schlecht.)Blume ofwestern
Panama to Mexico, but the extremeslenderness
and small racemesof our plant,theleaf size, and
especiallythe consistentlycaudate to acuminate
leaf apex, leave littledoubt thatthisis a distinct
species. Struthanthussubtilisappears to be en-
demic to the Cocle-Veraguas region.
LITERATURE
CITED
EICHLER,A. W. 1868. Loranthaceae. In C. F. P. Mar-
tius, Flora Brasiliensis5(2): 1-135.
KuIJT,JOB. 1961. A revisionof Dendrophthora (Lo-
ranthaceae).Wentia 6: 1-145.
. 1964. A revisionofthe LoranthaceaeofCos-
ta Rica. Bot. Jahrb.83: 250-326.
1975. The genus Cladocolea(Loranthaceae).
J. Arnold Arbor. 56: 265-335.
1978. Commentaryon themistletoesofPan-
ama. Ann. Missouri Bot. Gard. 65: 736-763.
1986a. Viscaceae. In Flora of Ecuador, 24:
13-112.
1986b. Observations on establishmentand
earlyshoot emergenceof Viscumminimum(Vis-
caceae). Acta Bot. Neerl. 35: 449-456.
1987. Miscellaneous mistletoenotes, 10-19.
Brittonia(in press).
TRELEASE,W. 1916. TheGenusPhoradendron. Univ.
Illinois Press, Urbana.
TUBEUF,K. 1923. Monographie der Mistel. Olden-
bourg,Berlin.