Вы находитесь на странице: 1из 133

Organization for Flora Neotropica

Bignoniaceae: Part I (Crescentieae and Tourrettieae)

Author(s): Alwyn H. Gentry
Source: Flora Neotropica, Vol. 25, No. 1, Bignoniaceae: Part I (Crescentieae and Tourrettieae)
(Sep. 19, 1980), pp. 1-130
Published by: New York Botanical Garden Press on behalf of Organization for Flora
Stable URL: http://www.jstor.org/stable/4393736
Accessed: 21/12/2008 18:02

Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at
http://www.jstor.org/page/info/about/policies/terms.jsp. JSTOR's Terms and Conditions of Use provides, in part, that unless
you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you
may use content in the JSTOR archive only for your personal, non-commercial use.

Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at

Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed
page of such transmission.

JSTOR is a not-for-profit organization founded in 1995 to build trusted digital archives for scholarship. We work with the
scholarly community to preserve their work and the materials they rely upon, and to build a common research platform that
promotes the discovery and use of these resources. For more information about JSTOR, please contact support@jstor.org.

New York Botanical Garden Press and Organization for Flora Neotropica are collaborating with JSTOR to
digitize, preserve and extend access to Flora Neotropica.



'~OPI ,CCANCI ?.\ ?..0... 0TIOPIC OF




The New York Botanical Garden

Bronx, New York 10458

Issued 19 September 1980

Copyright ? 1980

The New York Botanical Garden

Library of Congress Cataloging in Publication Data

Gentry, Alwyn H

(Flora neotropica ; monograph no. 25 ISSN 0071-5794)

Bibliography: p.
Includes index.
1. Bignoniaceae. 2. Botany--America. 3. Botany
--Tropics. I. Title. II. Series.
QK495.B62G46 583' .54 80-10846
ISBN 0-89327-222-1

Published by
The New York Botanical Garden
Bronx, New York 10458

International Standard Serial Number 0071-5794

All material subject to this copyright may be photocopied for the non-commercial purpose of
scientific or educational advancement.
(Crescentieae and Tourrettieae)


Table of Contents
Introduction .. ................................................ ................... 2
HistoricalSummary................................................................ 2
Linnaeanperiod(before1789).................................................... 3
Post-Linnaeanperiod(1789-1837)................................................ 3
Candolleanperiod(1838-1845)........................................... ........ 4
Latenineteenthcentury(1845-1900)............................................ 6
Modernperiod(after1900) .............................................. 8
Morphology..................................... ................................. 10
Habit......................................................................... 10
Bark ......................................................................... 11
Wood andStemAnatomy....................................................... 11
Twigs ........................................................................ 12
Leaves........................................................................ 13
Indument ................................................................ .... 15
Inflorescence ....................................................... 15
Calyx................................................................ 17
Corolla. .................... ...................................... 17
Stamens ...................................................................... 18
Pistil ......................................................................... 19
Disk.......................................................................... 20
Fruit ......................................................................... 20
Seeds......................................................................... 22
Seedlings...................................................................... 22
PollenMorphology................................................................. 23
Chemotaxonomy ................................................................... 26
Cytology ..................................................................... ... 27
PhylogenyandEvolution............................................................ 31
PollinationEcology ................................................................ 32
Floralmorphologyandpollenvectors ............................................. 33
Phenology .................................................................... 37
Intra-community constancyandnichespecificity.................................... 38
DispersalEcology.................................................................. 38
Distributionand Phytogeography ..................................................... 40
Bignonieae.................................................................. . 40
Tecomeae ..................................................................... 41
Crescentieae................................................................... 43
Schlegelieae................................................................... 43
TourrettieaeandEccremocarpeae................................................. 44
Uses.................. ........................................ ................... 44
SystematicTreatment.................................... ......................... 47
Crescentieae ................................................................... 49
Amphitecna............................................................... 50
Crescentia................................................................. 82
Parmentiera............................................................... 97
Tourrettieae.... ........................................................... 110
Tourrettia ......................................................... .... 110
Acknowledgm ents.................................................................. 114
LiteratureCited... ................................ ................... 114
NumericalListof Taxa.............................................................. 118
Listof Exsiccatae........................................................... . 119
Indexof VernacularNames..................................... .................. 125
Indexof ScientificNames ....................................................... 127

'AssociateCurator,MissouriBotanicalGarden,P.O. Box299, St. Louis,Missouri63166.

2 FloraNeotropica


This is the first of a plannedthreepart monographof the 600 speciesof

neotropicalBignoniaceae.Five of the family'seight tribes-Bignonieae, Cres-
centieae, Eccremocarpeae,Schlegelieae,and Tourrettieae-are restrictedto
the neotropicsand a sixth-Tecomeae-occurs in both Old World and New
Worldswith a preponderanceof speciesin the Americas.This first part of the
Flora Neotropica monograph of the family treats tribes Crescentieaeand
Tourrettieaeand includesintroductorymaterialrelatingto the whole family.
The projectedsecond part of the serieswill treatTecomeae,Schlegelieae,and
Eccremocarpeae; the thirdwill coverthe largesttribeBignonieae.


The taxonomichistory of the Bignoniaceaecan be convenientlybroken

into five periods,each characterizedby a differentapproachto the classifica-
tion of the family. Linnaeusand his immediatesuccessorsregardedthe species
of this family as comprisingonly two genera, Bignonia and Crescentia.The
post-Linnaeanperiod from 1789 through 1837 saw the descriptionof many
new speciesand their divisioninto many additionalgeneraand severaltribes
with differencesin calyx and habit the primedeterminants.The most impor-
tant students of the Bignoniaceaeduring this period were Jussieu, Kunth,
Bojer,and G. Don. The Candolleanperiodfrom 1838to 1845is bracketedby
two importantworks of de Candolleand encompassesless importantstudies
by a numberof contemporaryauthors. Duringthis period, charactersof the
fruit becamethe chief taxonomicdeterminantswhile most of the problemati-
cal borderlinegenerarelatedto Bignoniaceaeweresortedinto tribesand fami-
lies moreor less in accordancewithmodernideasas to theiraffinities.The late
nineteenthcenturyfrom 1845to 1900saw a fragmentationof the studyof this
family betweenthe various Europeancountrieswith a great deal of generic
splittingin generalvogue. The most importantstudentsof the family during
this time were Miers and Seemannin England,Bureauand Baillon at Paris,
and K. Schumannin Germany.The modernperiod in the taxonomy of this
family from roughly 1900to the presenthas been markedby the introduction
of a numberof new charactersinto the classificationof this family with new
and often monotypicgenerabased on each new character;the ascendancyof
Sandwithas the world's pre-eminentstudentof the Bignoniaceaeresultedin
the merging of many superfluousspecies with the consequencethat many
generacame to includeonly a very few species. Indeedpriorto my own work
therewere fewer speciesper genus in Bignoniaceaethan in any other largeor
medium-sizedplant family(Gentry,1973a)and a mind-boggling54 monotypic
genera(Gentry& Tomb, 1979).I have concentratedon similaritiesratherthan
differencesin attemptingto redefinegeneraas largerand more variable,but
evolutionarilynatural, units and van Steenis (1978) has adopted a similar
approachin his treatmentof the familyfor FloraMalesiana.Perhapsthe latter
HistoricalSummary 3

half of the twentiethcenturywill see the emergenceof a comprehensivepicture

of phylogeneticrelationshipswithinthis family.

Linnaeanperiod(before 1789)

Unrepresentedin Europe,the familywas first broughtto the attentionof

Europeanbotanistsby the returningexplorersfrom Asia, Africa, and Ameri-
ca. The nameBignoniadates from Tournefort(1719)who gave a rathergood
characterizationof the family underthis genericname, includingan illustra-
tion of each diagnosticpartof the plant. Bignoniawas thus characterizedby a
monopetalous,tubular,somewhatbilabiateflower and a siliquiformfruit di-
vided into two locules by an intermediateseptum and containingmany flat,
wingedseeds. Linnaeus(1753, 1754)acceptedthe genusBignoniaand included
in it 13 species.He consideredCrescentiato be a distinctgenus with the single
species C. cujete which he placed near Bignonia in the DidynamiaAngio-
spermia.These 14 species, then, comprisedwhat we now know as the family
In Linnaeus'later publicationshe added more species to both Bignonia
and Crescentiabut did not changehis basic treatmentof them. It is perhaps
worthyof note that LinnaeusincludedCatalpa,whichhas only two fertilesta-
mens, in Bignoniaand underthe four-staminateDidynamiaAngiospermiain
accordwith its naturalaffinitiesbut in directoppositionto his sexualsystem.
This is anotherexampleof his clear understandingof naturalrelationshipsin
contrastto some of his successorswho followedthe sexualsystemto the letter
in removingCatalpafromthe DidynamiaAngiospermia.


The firstimportantchangein classificationof the Bignoniaceaecamewith

Jussieu(1789)who consideredthis group(alongwith the modernPedaliaceae,
Martyniaceae,and Cheloneof the presentday Scrophulariaceae) to constitute
one of his naturalorders,the Bignones.Crescentiawas placedfar awayin the
Solanes. He subdividedthe Bignonesinto three groups on the basis of habit
and fruit, partitioningLinnaeus'speciesinto a numberof new genera.Incar-
villea was placed togetherwith Cheloneand Sesamumon the basis of herba-
ceous habit and capsular2-valvedfruit; Tourretia(sic) was placed with Mar-
tynia, Craniolaria,and Pedalium in a group characterizedby herbaceous
stemsand a woody-coriaceousfruit dehiscingat the apex;the rest of the mod-
ern Bignoniaceaeas then known were retainedin a group characterizedby a
woody habit and a 2-valvedcapsularfruit with the generaMillingtonia,Jaca-
randa,Catalpa,and Tecomabeingrecognizedin additionto Bignonia.
Descriptionsof new species of tropicalplants accumulatedrapidlytoward
the end of the eighteenthcenturywith Vahl (1798), Jacquin(1760, 1763, 1797,
1800-1809),and Aublet (1775)makingthe most importantsuch contributions
4 FloraNeotropica

to the Bignoniaceae.By 1802, Willdenowincluded54 species in Bignoniaas

well as recognizingmost of the generaof Jussieu.
In 1810Brownreducedthe Bignoniaceaeto Jussieu'sbasicsection, segregat-
ing the thirdsectionof Jussieuas the Pedaliaceae.Kunth(1818), on the other
hand, kept this group with the Bignoniaceaebut treatedit as a separatesec-
tion, the Sesameae,characterizedby winglessseeds and simple leaves as op-
posed to the Bignoniaceaeveraewhichhad seedswith membranaceouswings.
Underthis classificationTourrettiawas transferredback to the Bignoniaceae
whereit and Eccremocarpuswereplacedalongsidethe extraneousnew genera
PlatycarpumBonpl. (now Rubiaceae)and CobaeaCav. (now Polemoniaceae).
Kunth(1819,in HBK.)was also apparentlythe first to note the relationshipof
Crescentiaand Tanaeciumwith the Bignoniaceaeplacing them as "Genera
Bignoniaceis affinia"in theNova Generaet SpeciesPlantarum.
Bojer(1837)and G. Don (1838)proposednearlyidenticalnew systemsof
classification.for the Bignoniaceae.Under these systemsof classificationthe
present-dayTecomeaeand Bignonieaewereplacedin the tribeBignonieaeand
Crescentiaand Tanaeciumin the Crescentieae.Bojer'sfinal tribe, Coleeae,in-
cludedColeaand Arthrophyllum(Phyllarthron),whilethat of Don, the Tour-
retieae, was composed of Tourretia,Eccremocarpus,and Calampelis, all
generauntreatedby Bojer. These systems,especiallythat of Bojer, were also
importantin introducingcalyx form as the most importantgenericdetermi-


WhereasSteudelin 1831had listedonly 131speciesof Bignoniaceae,only

seven years later de Candolle(1838) in his importantRevue Sommairede la
FamilleBignoniacdeslisted 357 speciesas more or less well known, partition-
ing them into two tribes,the Bignonieaewith 336 speciesand the Crescentieae
with21 species.The Bignonieaeof de Candolleweresubdividedinto threesub-
tribeson the basis of theircapsules,introducingan importantnew taxonomic
character. Eubignonieeswere characterizedby marginicidaldehiscence (a
namecoined by de Candollespecificallyfor this group)and valvesparallelto
the septum.Catalpeeshad loculicidaldehiscenceand the septumperpendicular
to the valves. The third group, Gelsemiees,was characterizedby fruits more
like those of the Apocynaceaethan like the other Bignoniaceae.This was per-
haps not surprisingconsideringthat the two generareferredto this subtribe,
Gelsemiumand Platycarpumare now referredrespectivelyto the Loganiaceae
and Rubiaceae!The Crescentieaewerelikewisedividedinto two subtribes,the
Crescentieesand the Tanaeciees,the first with fruit oval or globose and uni-
locular,the secondwith fruit elongate,2-locular,and the leaves alwaysoppo-
site or verticillate.
In additionto these majorgroups, de Candolleincludednine monotypic
"Generisnon satis nota seu anomala"in the family,threeof whichhe himself
described.Of these nine, only Rhigozumis now referredto the Bignoniaceae,
the othersbeing Apocynaceae(Peltospermum= Aspidospermum),Scrophu-
lariaceae (Wightia, Esterhazya),Oleaceae (Schrebera),Verbenaceae(Psilo-
HistoricalSummary 5

gyne = Vitex),Acanthaceae(Bravaisia),Martyniaceae(Holoregmia = Mar-

tynia),and Celastraceae(!) (Trigonocarpus= Kokoona).
Despitehis "generisanomala," de Candolle'sclassificationof the family
was clear and concise, his divisions based on fruiting characterseminently
more naturalthan the calyx-baseddivisionsof Bojer. Of his eight generaof
Crescentieae,only Boutonia (= Periblema)is now rejectedto the Acantha-
ceae;in the Bignonieaeall of the 13generaof Eubignonieaeand all but two of
the 14 genera of Catalpeae(Neowedia(= Ruellia;Acanthaceae)and Paul-
ownia (Scrophulariaceae but retainedin Bignoniaceaeby many authors))are
currentlyplacedin other families. However,some of the generaplacedby de
Candolle in the Eubignonieae have since been shifted to his Catalpeae
(Argylia,Astianthus, Stereospermum,Delostoma)and two of the genera of
the latter(TourrettiaandEccremocarpus)arenow placedin separatetribes.
Otherclassificationsof the familywhichappearedat about this time were
much fartherfrom the moderninterpretation.Rafinesque(1838)published19
new and often ill-conceivedgenera.Endlicher(1836-1840)put the Crescentieae
underthe Gesneriaceaeand the Sesameaebackinto the Bignoniaceae.He also
added some "GeneraAnomala and Dubia" includingMetternichia(Solan-
aceae), Oxera(Verbenaceae),and Ferdinandusa(Rubiaceae).Meisner(1840,
1841)includeda numberof new generawhichhe attributedto Martiusand de
Candolle,presumablytakingthem fromthe manuscriptof the latter'sProdro-
mus. He also includedthe Sesameaein the Bignoniaceaealong with four other
suborders(i.e. tribes), the Eccremocarpeae,Incarvilleae,Tourretieae,and
Bignonieae,the lattersubdividedinto the tribes(i.e. subtribes)Argylieae(dis-
sepiment contrary to the valves, seeds unwinged),Tecomeae (dissepiment
contraryto the valves, seeds winged),and Eubignonieae(dissepimentparallel
to the valves).
The first (andto datelast)comprehensivemonographof the Bignoniaceae
was de Candolle's(1845)treatmentin the Prodromus(publishedposthumous-
ly with additionalcommentsby Alphonse de Candolle). It was basicallythe
same as his earlierwork with the same two major divisions. However, the
"generaanomala" (except Bravaisia)were removedalong with the subtribe
Gelsemieae(Platycarpumwas placed in Catalpeae)and the Catalpeaewere
split into two additional subtribes, the Incarvilleaeand Eccremocarpeae.
Despitethe removalof 12of the 13extraneousgeneramaintainedin his earlier
treatment,de Candollenow included50 generain the family with a total of
525 species, 492 in Bignonieaeand 33 in Crescentieae.With the exceptionof
Bravaisiaand Platycarpum,this outlined the family exactly as we know it
The infrafamilialclassificationof de Candolle, however,was not nearly
so closely in accordancewith modernconcepts. He dividedeach of his major
subtribesof Bignonieaeinto two series ("divisions") the Monostictideswith
seeds uniseriateon each side of the septum and the Pleistictideswith seeds
multiseriateon each side of the septum,a characterratherdifficultto ascertain
in the frequentabsenceof fruitingmaterialencounteredin this group. Further-
morethereare a numberof ambiguousgenerawith seedsmore or less 2-seriate
on each side of the septum and some clearly relatedgenera are widely sep-
aratedby this artificialcharacter.Even worse was de Candolle'sover-useof
6 FloraNeotropica

calyx featuresas genericdeterminantswhichresultedin unnaturalcollections

of speciesunder, for example, Tabebuia(bilabiatecalyx), Spathodea(spatha-
ceous calyx), and Tecoma(sub 5-toothed calyx). Other genera were equally
poorly defined. Bignonia was maintainedas a veritablehodge-podgeof 173
mostly unrelatedspecies. The new genus Pachypteraincludedfour "species"
identicalto Paragoniapyramidataalong with a single species each of Mac-
fadyena and the modernPachyptera.Adenocalymmaincludeda numberof
completelyheterogeneouselementsand Tanaeciumincludedthe present-day
Schlegelia.But despiteits obviousdeficiencies,the monographof de Candolle
providedthe chief referencepoint and a frameworkfor latertaxonomyof this

Subsequentto the Prodromus,study of this familywas evidentlycarried

on moreor less independentlyin the variousEuropeancountrieswith contrast-
ing ideas and considerableduplication in descriptionsof new genera and
species.Thiscreateda good dealof taxonomicconfusion.
The Englishworkersin the familywereperhapsthe most numerous.See-
mannand Mierspublishedimportantstudiesof the familyin the 1860's. Ben-
tham& Hooker's(1876)treatmentwas followedby that of Hemsley(especially
1882).This school was characterizedat first by excessivesplittingwith Cres-
centiaceaeconsidereda separatefamilyby Lindley(1847)and Seemann(1860)
whileMiers(especially1863)publisheda numberof new generaand extensive
lists of new species,mostlyas nominanuda.
This trendwas abruptlyreversedby Bentham& Hookerwho, followedby
Hemsleyand manyearlyAmericanauthors,reducedCrescentiaceaeto Bigno-
niaceae,includingin the compositefamilyonly 52 generaand 450 speciesand
following very closely the treatmentof de Candolle. The treatmentof Ben-
tham & Hooker, however, correctedmany of the most serious flaws in the
Prodromus.The misplacedgeneraof the Tecomeae(Delostoma,Astianthus,
Cybistax)were removedto their correctposition in the Tecomeae.Schlegelia
was separatedfrom Tanaecium(followingMiquel)and the lattermoved from
the Crescentieaeto the Bignonieae.The use of the calyxin genericdelimitation
was given less importanceso that the spathaceous-calyxedmembersof the
Bignonieae,whichhad beenplacedby de Candollein Spathodea,wereput into
Macfadyenawhile Tecomaof de Candollewas split along more naturallines
with its digitate-leavesspecies being moved to a now natural Tabebuiafrom
whichthe membersof the Bignonieaeplacedthereby de Candollehad beenex-
cluded(see Gentry, 1969).Whilethe Monostichaeand Pleiostichaeof de Can-
dolle wereretainedas basic subdivisionsof the Bignonieae,Distictisand Hap-
lolophiumweremoved from the formerto the latter. In the Tecomeae(Catal-
peae of de Candolle)this tribal subdivisionwas abandonedfor a somewhat
morenaturalone basedon severalcharacters,its five majorsubdivisionsbeing
Trichospermeaewith seed wings of fused hairs (Catalpa, Chilopsis, and
Sparattosperma),Pentandreaewith five fertilestamens(Rhigozumand Cato-
phractes(but not Oroxylum),Folliculareswith Amphicome,Incarvillea,and
HistoricalSummary 7

Tourretia,Digitifoliae with simple or digitatelycompound leaves (including

Argylia),and Pinnifoliaewith pinnatelycompound(rarelysimple)leaves. The
Crescentieaewere regardedas no more than a tribe with equal statureto the
Tecomeaeand Bignonieae.The greatflaw in the Bentham& Hookerclassifica-
tion is their final tribe, the Jacarandeae,a very unnaturalassemblagesup-
posedlycharacterizedby basicallyparietalplacentationwith a septumlacking
or incompleteand the capsulevalvesplacentiferousat the middle.This unfor-
tunate conglomerationincluded unrelatedgenera now assigned to the four
tribes Tecomeae (Jacaranda and Digomphia), Crescentieae (Parmentiera),
Coleeae (Colea), and Eccremocarpeae (Eccremocarpus).
The classificationof Bentham& Hookerwas followedverbatimby Hemsley
and later to a large extent by Spragueand his successorSandwith.The pre-
eminenceachievedby the latterin the studyof this familyduringthe first half
of the twentiethcenturywas probablyin part responsiblefor the almost uni-
versalpresent-dayacceptanceof the basic elementsof thissystem.
In Holland, Splitgerber(1842) and Miquel (1844, 1850) made valuable
contributionsto an understandingof the Bignoniaceae,but the majorcontri-
butionsof the Dutch school to taxonomyof the Bignoniaceaewerelimitedto
the Old World species. The many more recent publicationson Australasian
speciesof this family by van Steenis(1927, 1928, 1929, 1978)providemost of
our presentknowledgeof the familyin that geographicarea.
Frenchworkersin the familyachievedtheirgreatestsuccesswith the pub-
lication of Bureau's (1864) Monographie des Bignoniacdes, a compendium of
the taxonomichistory of the family, its taxonomicallyuseful characters,and
most important,a surveyof all generawhichhad everbeen includedin or con-
fused with the Bignoniaceae.He dividedthem into 15 groups, only the first
three of which (correspondingroughly to the tribes Bignonieae,Tecomeae,
and Crescentieaeof Bentham& Hooker) he acceptedinto the Bignoniaceae.
This group includedabout 65 genera.The other45 generawere arrangedinto
12 groupscontainingone to ten generaeach, corresponding to the 12 familesto
which they actually belonged. These families were the Sesamees(4 genera),
Pedalinees(7 genera includingEccremocarpusand Tourrettia),Cyrtandrees
(10 generaincludingSchlegelia),Gentian6es(Kuhlia),Acanthacees(3 genera),
Polemoniac6es(Cobaea), Verbenacees(2 genera), Scrophularinees(9 genera
includingPaulownia), Solanees (2 genera), Apocyntes (2 genera), Oleacees
(Schrebera)and Rubiacees(3 genera).The monographalso includeda nicely
illustratedAtlas encompassingmanyrepresentativespeciesof the family.
Sincethis treatmentwas at the genericlevel, veryfew of Bureau'sideas of
specific relationships were included. Bureau later published revisions of
Lundia (1868a), Saldanhaea (1868b), and Schizopsis (1865, = Tynanthus
Miers(1863))and then, possibly frustratedby duplicationbetweenhis efforts
and those of Miers, apparentlyturnedhis attentionto other areas except for
occasional notes in the Transactionsof the Linnaean Society of Paris.
However, many of the unpublishednames writtenby Bureauon herbarium
specimensat the variousEuropeanherbariahave persisted,being taken up by
laterauthors.AlthoughBureauwas nominallythe seniorauthorof the monu-
mental treatmentof Bignoniaceaefor Martius'Flora Brasiliensis,he appar-
8 FloraNeotropica

entlydid little or none of the actualwork whichwas done ratherby the junior
authorK. Schumann.
Most noteworthyof other Frenchstudentsof the familywas Baillonwho
workedextensivelywith the familyin preparingits accountfor his Histoiredes
Plantes(1888).At the genericlevel, Baillonwas an inveteratesplitter,although
some of his new nameswereattributableto Bureau.Baillonrecognized97 gen-
era arrangedinto the four tribes Bignonieae(44 genera), Tecomeae(39 gen-
era), Crescentieae(13 genera including Schlegelia),and Eccremocarpeae(1
genus). Except for his rejectionof Tourrettia,this tribal alignmentis that in
use today with the generaof Bentham& Hooker'sJacarandeaepartitionedby
their evident naturalrelationships.The excessivegeneric splitting is even in
accord with moderntaxonomythough probablynot reflectingtrue phyloge-
Following Bureau and Baillon as French students of the Bignoniaceae
were Dop (1925, 1929) and Pichon (1945, 1946) who continued the trend
greatlyto the studyof this family.
In Germanyboth Chamisso(1832) and Schlechtendahl(1833, 1853) de-
scribed a great many new species between 1832 and 1854. Later Grisebach
(1866, 1874) describeda number of species from Argentina and the West
Indieswhile Urban(1916)describednumerousWest Indianspeciesand did an
importantstudy of pollen types in the family. But it was K. Schumannwho
mustbe consideredthe greatestGermanstudentof Bignoniaceae.His workfor
Engler & Prantl's NaturlichenPflanzenfamilien(1894) set the stage for his
monumentalstudyin Martius'Flora Brasiliensis(Bureau& Schumann,1896-
97). Schumann'stribeswerethe same as those of Baillonwith the additionof
Tourrettiaas the Tourrettieae.In most cases Schumannalso followedthe gen-
eric delimitationsof Bureau and Baillon rather than those of Bentham &
Hooker. Thus Tecoma included most of the digitate-leavedspecies of
Tabebuia;the names Stenolobium, Tecomaria,Campsis, Campsidium,and
Pandoreawere used for the pinnate-leavedspeciesof Tecoma;and Bignonia
sensuBentham& Hookerwas subdividedinto numeroussegregates.Thanksto
the Flora Brasiliensis,Schumann'sgenerictreatmenthas remainedin general
use in Brazil,althoughmany of his genericconceptshave been greatlymodi-
fied by subsequentauthors.Schumann'ssuccessorsas studentsof Bignoniaceae
at Berlin were Loesner and Melchior, neither of whom contributedsignifi-
cantlyto knowledgeof the family.

Modernperiod(after 1900)

After the turn of the centurythe Englishteam of Spragueand Sandwith,

and later Sandwithalone, becameincreasinglypre-eminentin studiesof this
family,especiallyin the New World,whilethe workof van Steenisin Malaysia
formedthe basis of its taxonomyin the Old World. Sandwith'sgrasp of the
taxonomyof the Bignoniaceaewas the most comprehensiveto date and his un-
derstandingof species delimitationwas unsurpassed.He was an extremely
HistoricalSummary 9

carefulworkerwho went to greatcareto be scrupulouslyaccurate.He assem-

bled an excellentcollection of Bignoniaceaeat Kew includinga remarkably
completeand well-labelledset of types duringhis half centuryof studyingthe
family. There are three importantdrawbacksto Sandwith'swork, however.
His publicationswere scatteredthrough the taxonomic literature,mostly as
short papers in the Kew Bulletin, and he failed to summarizehis extensive
knowledgeinto a comprehensivemonographof the family. Furthermorehis
field experiencewas verylimitedwhichwas a majordifficultyin evaluationof
rangesof intrapopulationaland intraspecificvariation,as he himselffrequent-
ly noted. Finallyhis taxonomywas almosttoo logicalin the sensethat natural
relationshipswereoften obscuredby genericsplittingbasedon logicalbut phy-
logeneticallyunimportantcharacters.Althoughhis specificjudgments(with a
few significantexceptions)were beyond reproach,his genericconcepts were
exceedinglynarrowas has beenpointedout especiallyby Macbride(1961).
Part of the problemof genericdelimitationin this familycan be tracedto
the recenthistoryof its taxonomy. Since 1900a numberof more or less new
taxonomiccharactershave been proposedby a seriesof authorsand each of
these charactershas then been used to split existinggenera. Although all of
these charactershave been of importancein classifyingthe family, little at-
tempt has been made at overallevaluationof the importanceof specific con-
stellationsof charactersin individualcases. Ratherthe tendencyhas been to
acceptthesecharactersas absolutegenericdeterminantsand whengenerahave
been found to have species differingin such charactersthey have been sum-
marilysplit. The only significantgeneric-levelconsolidatorywork in the New
WorldBignoniaceaepriorto my own studieswas the posthumoustreatmentof
Arrabidaeaby Sandwith(1968).
The first of these important20th centurystudiesof the Bignoniaceaewas
that of Urban(1916)who surveyedtendriland pollen variationin the family.
His workwith Bignoniaceaepollenis in fact usuallyconsideredone of the clas-
sical palynologicalstudies. On the basis of these two characters,generallyne-
glectedby earlierauthors,he characterizedeach genus of the family and pro-
posed the segregationof Pseudopaegmawith 5-6-colpatepollen grains from
Anemopaegmawhichhe characterizedas inaperturate,of Roentgeniawith 12-
colpatepollengrainsand bifid or trifidtendrilsfrom Cydistawithinaperturate
pollenand simpletendrils,and of Nestoriawith 3-colpatepollenand trifidten-
drils from Memora with inaperturatepollen and simple tendrils (see also
Gentry& Tomb, 1979). Urban also pointed out several species which were
anomalousin their respectivegeneraon the basis of pollen and most of these
weresoon segregatedby otherauthors.
In 1931 Rohrhoferpublisheda paper discussingtaxonomicimplications
of the staminodein Bignoniaceae,but no significantconclusionsweredrawn
and the only genera characterizedto a large extent by their staminodesare
Jacarandaand Digomphia with elongated more or less glandular-pubescent
In 1948Seibertpointedout the taxonomicutilityof yet anothercharacter
in the Bignoniaceae:the presenceor absenceof glandsor glandularfields and
their distribution.He characterizedten types of glands found in the family,
10 FloraNeotropica

listinggeneracharacterizedby severalof these gland types. Although Seibert

himselfdid not proposeany new generaon this basis, Sandwithgenerally(but
not in his posthumous(1968) paper) consideredthe presenceor absence of
interpetiolarglandularfields as a genericdeterminant,and a numberof his
genera were proposed or maintainedas distinct largely on this basis. Other
taxonomicallyimportanttypes of glands noted by Seibertinclude glandular
fields at the apex of the petiole in Pseudocalymmaand Pachyptera(these are
also presentin Paragonia),calyx glands, neuroaxillaryglands and glandular
fields, glands on the outside of the corolla below the lobes, and the pellucid
leaf glandswhicharethe primarydeterminantof Stizophyllum.
A secondimportantpaperon the use of pollencharactersin the taxonomy
of the Bignoniaceaewas publishedby Gomes (1955). This paperis illustrated
with 29 carefullydrawnfiguresof pollen grainsand includesa key to the Bra-
ziliangenerabasedchieflyon pollencharacters.One otherpaperis noteworthy
as usingpollen charactersin partto segregatenew genera.This is the work of
Pichon (1945)in whichno less than 11 generaweredescribed,mostlyas segre-
gates fromArrabidaea.
Modernneotropicalfloristicwork in the family has includedtreatments
for Guatemala,British Hondurasand southernMexico (Seibert, 1940), for
Peru (Macbride,1961), for Argentina(Fabris,1965),for ItatiaiaPark, Brazil
(Gomes, 1957),for Guatemala(Williams,1973),and by Sandwithfor Surinam
(1938),Trinidadand Tobago (1954),and SantaCatarina(1974, posthumously
revisedby D. Hunt). In the last severalyears I have publishedfloristictreat-
ments for Panama (1973c),Ecuador(1977c),the GuayanaHighland(1978a),
and Venezuela(1979b).
Certainother authorsmeritspecialattentionin a surveyof the taxonomy
of the Bignoniaceae.Rusby, Krinzlin, and to some extent Pittier are note-
worthy for the profligacyof their descriptionsof new species of this family
and for the low percentageof theirnewlydescribedspecieswhichhave turned
out to be actuallynew. Had these systematistsexerciseda bit more caution in
theirdealingswiththe Bignoniaceae,the synonymiesof manyspecieswouldbe
The 1965 death of Sandwithwho had so totally dominatedtaxonomic
workin the Bignoniaceaefor manyyearsleft a vacuumin this family. The re-
sults of my own field-study-basedwork accentuatethe fact that additional
cluesto a taxonomicand evolutionaryunderstandingof this familywill surely
be found in the field ratherthan in the herbarium.The taxonomicframework
establishedby Sandwithis now being correlatedwith the actualplantsas they
occurin nature,settingthe stagefor the monographof the familyof whichthis
treatmentof the tribesCrescentieaeand Tourrettieaeconstitutesthe firstpart.


All genera of Bignonieae are exclusivelyor predominantlylianas, al-

though some generaalso have a few suffruticosespecies, mostly in the Bra-
Morphology 11

zilian cerrado. Most genera of Tecomeae are trees, often canopy giants 30 or
more meters tall, but some are shrubs, a few are lianas (mostly in the Old
World), and two genera-Argylia and Incarvillea-are herbaceous. Crescen-
tieae and Coleeae are trees or shrubs. Schlegelieae are mostly hemiepiphytic
vines or epiphytic shrubs. Tourrettieae and Eccremocarpeae are herbaceous to
wiry vines. Differences in habit are not always taxonomically significant, espe-
cially in suffruticose cerrado forms of species like Arrabidaea brachypoda
(DC.) Bur. which are usually vines, but the dichotomy between lianous
Bignonieae and arboreal Tecomeae and Crescentieae is a fundamental one in
the neotropics. The New World lianas are mostly tendril climbers (see below),
sometimes supplemented by adventitious roots. One climbing genus, Schle-
gelia, is hemiepiphytic and lacks tendrils. Old World climbers lack tendrils and
are twiners or climb by adventitious roots.


A taxonomically neglected character, not amenable to herbarium analy-

sis, bark nevertheless has some taxonomic utility. Many of the species and
genera of lianas can be recognized in the field by presence and type of lenticels
and nuances of bark texture and coloration. In some of the tree genera there is
a marked correlation of thick vertically ridged bark with cerrado or dry forest
species and thinner, smoother, often exfoliating bark in wetter forest species.

Wood and Stem Anatomy

Wood anatomy of Bignoniaceae has been summarized by Record & Hess

(1940) and more recently by van Steenis (1978). Van Steenis' general conclu-
sion is that wood anatomy of Malesian tree genera is rather homogeneous with
aliform or confluent parenchyma, homogeneous rays, fibers with simple to mi-
nutely bordered pits and simple (rarely also reticulate) vessel perforations.
New World tree genera show more variability on the same basic ground plan.
Density and hardness are especially variable from a specific gravity of 0.40 to a
specific gravity of 1.25 (Record & Hess, 1940). Even in a single genus there
may be great variation in the wood. The wood (especially of the roots) of some
Brazilian species of Tabebuia is so exceptionally light and buoyant that it is
widely used for life buoys, fishing floats, and inner shoe soles (Correa, 1926)
while other species of the same genus have some of the heaviest and hardest of
all woods. The temperate zone tree genera have ring porous wood, the tropical
genera do not. Rays vary from uniseriate to eight cells wide and may be homo-
geneous or decidedly heterogeneous, and parenchyma may be paratracheal,
aliform, confluent, or in concentric bands. Wood fibers are occasionally sep-
tate and ripple marks may be present or absent (Record & Hess, 1940). Inter-
generic wood variation is consistent enough that Record & Hess were able to
construct a generic key based on wood anatomy but Tabebuia falls out in no
less than eight different places and spans most of the familial range of
12 FloraNeotropica

The wood anatomyof the lianasof tribeBignonieaeis spectacularlyvari-

able with striking patterns of anomalous structure.The anomalous wood
anatomyresultsfrom failureof certainsegmentsof an otherwisenormalcam-
bium to produce secondaryxylem while continuing production of phloem
(Dobbins, 1971)resultingin a secondaryxyleminterruptedby deep furrowsor
wedgesof phloem. All generaof Bignonieaehave anomalouswood structure
althoughthis is lost in some individualspeciesof severalgenerawhichare suf-
fruticoseor treelets. No genus of Tecomeaehas anomalouswood anatomy,
not even the lianas. Thus the presenceof anomalousstructurein Dolichandra
strongly supports its inclusion in Bignonieae despite perpendicular-to-the-
septum fruit dehiscence. The wood structureof taxonomicallyanomalous
Perianthomegais unknown, but potentiallydefinitivefor tribal assignment.
Should Perianthomegaprove to have anomalouswood anatomy it would be
best retainedin Bignonieae;if not (as I anticipate)then it is best placed in
Anomalousstructureis also an excellentindicatorof genericrelationships
withinBignonieae.Thus Bignonia,Arrabidaea,Adenocalymma,Memoraand
other generahave uniformly4-armedanomalousxylem;large lianas of these
generacan often be recognizedby four conspicuousfurrowsin their trunks
correspondingto the four phloemarms. Manyothergenerahave a patternof
additionalsecondaryphloem arms, nearlyalways in multiplesof four. Thus
generalikeAnemopaegma,Mansoa,Mussatia,and Cydistamay have 8, 16, 32
or more phloemarms, dependinglargelyon the thicknessof the trunk,and to
an unknownextenton the individualspecies.Yet anotherbasicmodificationis
shown by the wood of Macfadyenaand Melloa which have the phloem arms
more numerous,less regular,and less distinctlyseparated.Thereis a general
correlationof increasedtrunk flexibilitywith increasingprominenceof the
anomalousrays so that Macfadyenawhich climbsappressedto tree trunksis
the most flexibleof all the largelianasof the family. Thereare additionaldif-
ferencesin fine structureof the anomalousrays withineach of the threegen-
eralgroupsand many of these modificationsare characteristicof certaingen-
era. Thus Tynanthushas the raysstronglystair-steppedand muchwiderat the
peripherywhile Ceratophytumhas them of nearlyuniformwidth from center
to periphery.Additional study is needed but patternsof anomalous wood
anatomyclearlyhold taxonomicpromise.


Twigsand small branchletsshow severalkindsof taxonomicallyapplica-

ble variation.In Parmentierathe presenceor absenceof nodal spinesis a use-
ful specificcharacter;a similardevelopmentof spinesoccursin Phylloctenium
of Madagascar.Twigs of Stizophyllumare characterizedby being hollow and
most Pleonotomaspecieshavethe extremecentersof largerbranchletshollow.
Conspicuouslyangled branchletsare a notable featureof severalgeneraand
tend to be extremelyuseful in genericdelimitation.For example,all speciesof
Amphilophium,Pithecoctenium,and Haplolphiumhave strikinglyhexagonal
Morphology 13

twigs with ribbedangles. Pleonotoma, Mussatia,and some speciesof Clytos-

tomaand Cydistahave sharplytetragonaltwigswith ribbedangles.
Interpetiolarglandularfields are often presentin tribe Bignonieae.Their
presenceor absenceis frequentlyused as a genericcriterion,with such genera
as Arrabidaea,Pachyptera,Ceratophytum,and Leucocalanthecharacterized
by presenceof interpetiolarglandularfields and Cydista,Mansoa, Paragonia,
Adenocalymmaetc., by theirabsence.Howeverthese glandularfields are not
alwaysconstanteven withinsome speciesand shouldnot be accordedabsolute
valueas a genericcharacter.ThusPetastomaand Cremastuswhichlack inter-
petiolarglands should be combinedwith Arrabidaeawith which they other-
wise agreeand Pseudopaegmawhich often has glandularfields should be re-
tained in Anemopaegmawhich otherwiselacks them. Glandularfields may
also be presentat the apex of the petiole. Petiolarglandularfields are defini-
tive for Paragoniaand also presentin severalspeciesof Pachyptera.
Bignoniaceaelack stipules but the scales of the axillarybuds are often
modifiedinto stipule-likestructuresknown as pseudostipules.Pseudostipules
may be foliaceous(e.g. Mussatia,some speciesof Cydistaand Anemopaegma),
subulate (e.g. Paragonia, Ceratophytum),bromeliad-like(Clytostoma), or
vertically2-3-seriate(Pachyptera).Pseudostipulesare often importantspecific
charactersand sometimesusefulin genericdefinition. In generalpresenceof a
certaintype of pseudostipulemay be definitive for certaingeneraor species
but absenceof pseudostipules,at least on limitedherbariummaterial,is rarely


The leaves of Bignoniaceaeshow a great deal of variability.In the great

majorityof speciesthe leavesare compoundand opposite.This conditionis so
prevalentthat most speciesof angiospermswith oppositecompoundleavesare
membersof Bignoniaceae.However, several genera-notably Amphitecna,
Delostoma, Catalpa,Chilopsis,Astianthus,Deplanchea,most Crescentia,and
the tribe Schlegelieae-are characterizedby simpleleaves and there are a few
simple-leavedspecies or individual variants in normally compound-leaved
genera.Onegenus-Amphitecna of the Crescentieae-has alternateleavesand
Crescentiahas alternatefasciclesof leaves.The predominantcompoundleaves
may be 2-3-foliateor pinnatelyor palmatelycompoundor variouslybicom-
pound. Oroxylumhas leaves with up to four ordersof subsidiarypinnae, the
recordfor any angiosperm.
Phyllarthronof Madagascarhas an unusualleaf formed from a linear
seriesof superimposedarticlesrepresentingthe wingedrachisand petioleof an
ancestrallycompoundleaf; the leaf blade of simple-leavedspeciesof Crescen-
tia is similarlyanalagousto the wingedpetiole of a 3-foliolateleaf like that of
C. alata.
There is a rathersurprisingbut apparentlyfundamentaldichotomybe-
tweenthe leavesof Old and New WorldTecomeae.Old World(andtemperate
zone) generaof the tribe have pinnatelycompoundleaves (or derivedsimple
leaves)and most New Worldgenerahave palmatelycompoundleaves (or de-
14 FloraNeotropica

rivedsimple leaves). This differenceis carriedover to the otherwisesimilarly

derivedherbaceousgeneraIncarvilleaof the Himalayas(pinnateleaves) and
Argyliacenteredin Chile (palmateleaves)and even to the derivedtribesCres-
centieaeand Coleeae. Among neotropicalgenera of Tecomeaeonly isolated
Jacaranda(plus its segregateDigomphia)and Tecomahave pinnatelycom-
pound leaves;the latterjust mightbe derivedfrom a tropicalLaurasianstock
despite its concentrationin the Andes to judge from its occurrenceinto the
southwesternUnited States,the existenceof the relatedmonotypicAstianthus
in Mexico,and the recencyof upliftof the Andes.
Otheraspects of Bignoniaceaeleaves are also taxonomicallyuseful. The
colorof driedleavesis often constantand one of the most obviousdistinguish-
ing charactersof severalspecies. A cartilaginousleaflet margincharacterizes
some speciesof Adenocalymmaand venationwith whitishcartilaginousmar-
gins characterizessome speciesof Amphitecna.Patternof venationand leaf
textureare importantspecificcharactersin severalgenera,especiallyfor spe-
cies havingleaflets noticeably3-veinedfrom the base or those havingwell de-
velopedintricatelyreticulatetertiaryvenationstronglyraisedbeneath.Leaf in-
dument,though frequentlyveryvariable,is also a very useful specificcharac-
ter, especiallythe presenceof branchedor stellatetrichomes.The presenceand
type of lepidotescales or plate-shapedglandsis taxonomicallyuseful in some
genera and one genus, Stizophyllum,is characterizedby glandular-punctate
leaves. Althoughleaflet size and shapetend to be highlyvariable,especiallyif
juvenilefoliage is considered,they are often useful specificcharacters.In sev-
eral general like Distictella, Crescentia,and Antillean Tabebuia,speciation
mostly involves vegetativedifferentiation,especiallyin leaf size, shape, and
The leaves of tribe Bignonieaeare especiallycharacteristicin havingthe
terminalleafletmodifiedinto a tendril.All generaof Bignonieaearecharacter-
ized by these leaf tendrilsalthougha few exclusivelysuffrutescentor treelet-
habitspeciesof generalikeAnemopaegma,Arrabidaea,and Memoralack ten-
drils. Most of these liana generahave basically3-foliolateleaves with both 2-
foliolate tendrillateleaves and 3-foliolate non-tendrillateleaves on the same
plant; a few species or generahave palmatelyor pinnatelycompoundor de-
compoundleaves with terminaltendrilson some of them. Leaf tendrilsare an
extremelyuseful charactertaxonomicallyat the generic level. Many genera
have exclusivelysimpleunbranchedtendrils,othersexclusivelytrifid tendrils,
althougha few genera(Anemopaegma,Memora)may have both types in the
same speciesand even the same plant. Some generahave the tendriltypically
trifurcatedseveraltimes; this is especiallytypicalof Pithecoctenium,Manao-
sella and Bignoniaand occursoccasionallyin Distictis, Pyrostegia,and a few
other genera.Macfadyenaand its relativesare characterizedby shortenedtri-
fid tendrilswith the armsthickenedand bent forminga hooked "cat's claw."
Another interesting and taxonomically useful tendril modification is the
formationof an attachmentdisk at the tendriltip(s). This is usuallyassociated
with compound tendrils (only Pachypteraparvifolia A. Gentry has a disk-
tippedsimpletendril)and often facultative,but is alwayspresentin Glaziovia,
Bignonia, and some species of Distictis. Both Tourrettiaand Eccremocarpus
have much-branchedleaf tendrilsbut climbingmembersof Schlegelieaeand
Morphology 15

Tecomeaelack them. Perianthomega,an anomalousisolated monotypicBra-

ziliangenuswith the fruitdehiscenceof Tecomeaehas a compoundleaf with a


Indument provides a wealth of specific characters but can also be

extremelyvariableintraspecifically.In many speciesthere is a strongpositive
correlationbetween pubescenceand dry habitats. Most of these pubescence
formshave receivedspecificrecognitionand at least one-Levya nicaraguensis
Bur. ex Baill. whichis an extremelypubescentform of variableCydistaaequi-
noctialis (L.) Miers-has been genericallysegregated.In severalcases "spe-
cies" whichlook remarkablydifferentin the herbariumon accountof pubes-
cence differenceshave been found growing on the same plant (e.g. Lundia
densifloraDC. and L. strictaBur. & K. Schum.;Cydistasarmentosa(Bertol.)
Miersand C. aequinoctialis(L.) Miers).Otherpubescence-differentiated spe-
cies pairs like Arrabidaeaegensis Bur. & K. Schum. and A. japurensis(DC.)
Bur. & K. Schum. which I have tentativelyretainedseem likely to prove vari-
antsof the samespecies.
Generallymore fundamentalthan the differencebetween presenceand
absenceof indumentis the distinctionbetweendifferentkindsof indumenttri-
chomes. Especiallysignificantare stellateand dendroidtrichomeswhichare a
constantcharacterfor many species. Differencesin pubescence,especiallyof
the calyx, are very importanttaxonomicallyin Tabebuia,and rathersubtle
variationsbetween simple, forked, stellate, dendroid, and thick-stellatetri-
chomesare majorspecificdeterminantsfor the yellow-floweredspeciesof that
genus. In other genera like Arrabidaeaand Amphilophiumtrichometype is
highlyvariedand taxonomicallyuseful. Lepidotescalesare also taxonomically
important;for examplepresenceor absenceof glandular-lepidotetrichomes
on the corollatube is one of the most importantcharactersfor differentiating
speciesof Anemopaegma.
An interestingaspectof indumentumvariationof Bignoniaceaeis the ten-
dency to have similarpubescencetypes on variousplant parts. Thus if the
leaves of a species have stellate trichomes,the twigs, inflorescence,calyces,
and fruits usually do also, and if calyx pubescenceis restrictedto lepidote
scales, usually so is the indumentof inflorescences,leaves, branchletsand


Inflorescencesof Bignoniaceaerange from terminalto axillaryto rami-

florousto cauliflorous.Especiallystrikingare some of the bat-pollinatedgen-
era whichexhibit flageliflory(Kigelia),"pin cushion" inflorescences(Oroxy-
lum, Haplophragma)and various types of cauliflory. In Amphitecna and
Parmentieraof tribe Crescentieaeplacementof the inflorescence(whichis re-
ducedto one or a few flowersand may be terminal,cauliflorous,or ramiflor-
16 FloraNeotropica

ous) is quite constant for a given species and an extremelyimportanttaxo-

nomiccharacter.In most otherNew Worldgenerathe inflorescencesare typic-
ally both terminaland axillary.Thesetwo conditionstend to intergrade.Thus
an axillaryinflorescencemay include reducedjuvenile leaves which suggest
that it is actually terminal on an immaturebranch. Converselythe lower
branchesof a fundamentallyterminalinflorescenceare often in the axils of the
upper leaves of a branch. Even species which are typically ramiflorousor
cauliflorousoften have some flowersborne terminallyor subterminallymak-
ing inflorescenceplacementan unreliabletaxonomiccharacter,except appar-
ently in the Crescentieae.Some attemptshave been made to segregategenera
on the basis of inflorescenceplacement, notably Saldanhaeawith lateral
inflorescencesfrom Cuspidariawith terminal inflorescences. In this case
inflorescenceplacementis fairlyconstantin a givenspeciesbut does not corre-
late with otherpotentialcharactersfor segregatingtwo genera,and Saldanhaea
is best retainedin Cuspidaria.
Inflorescenceform also providestaxonomicallyuseful characters,espe-
cially for specificdifferentiationin generawhichtend to have largecomplexly
branchedpanicles.For example,some speciesof Arrabidaeahave corymbose
branches,othersmore or less racemosebranches,and othersare variouslyre-
duced.A few generahave strictlyracemoseinflorescencesbut thereis usuallya
pronouncedtendencyfor the lowermostpedicelsof especiallyrobustinflores-
cences to become pedunculatebearingtwo or three flowers. Amphilophium
and Pithecocteniumare good examples.Another kind of intergradationbe-
tween raceme and panicle is shown by genera like Memora and Adeno-
calymmawhichtend to have severalbasicallyracemoseinflorescencesfrom a
commonbase. Anothercomplicationis posedby the fact that nearlyall species
and generacharacterizedby largeinflorescencesare also capableof producing
depauperatefew-floweredinflorescences.In view of the inherentintraspecific
variabilityin Bignoniaceaeinflorescences,I have not attemptedto describe
nuancesof each species'inflorescence,generallyusing "paniculate"in an in-
clusivesensefor any branchinginflorescence.
In some genera inflorescenceform is rather constant. Most species of
Anemopaegmabearracemesso that the inflorescenceof the few specieswhich
have branchingpanicles provides a very good distinguishingmark. Most
Tabebuiaspecieshavebasicallycorymbose-paniculate inflorescences(often re-
duced to capitate or to one or two flowers) lacking a centralaxis. Thus the
pyramidalinflorescencewith a well-developedaxis of Roseodendronis per-
haps the strongestreason for recognizingthat segregate,althoughI preferto
considerit a well-markedsectionof Tabebuiaratherthana distinctgenus.
Othertaxonomicallyuseful aspectsof inflorescencesincludepubescence
type and bracts. Inflorescencepubescencetends to be more constantin a spe-
cies than that of most vegetativeplant parts.Presenceor absenceof an inflor-
escenceindumentis taxonomicallyimportantin Lundiaand whetherthis indu-
ment has gland-tippedtrichomesor not is importantin Mansoa. Bractsand
bracteolesare especially importantin Adenocalymmaand Memora where
their presenceor absenceand placementand size relativeto calyx and pedicel
are among the most importantspecific characters.Occasionalconspicuously
bracteatespeciesin othergeneracan be easilyrecognizedby that character.
Morphology 17


Calyx morphologyprovidessome very importanttaxonomiccharacters.

Calycesof Bignoniaceaeare most frequentlycup-shapedbut are commonly
truncate,5-denticulate,bilabiate,or spathaceous.Severalgeneraor speciesare
characterizedby long filiformcalyx teeth. Most speciesof Lundiaare charac-
terizedby an evenly calypteratecalyx apex. Amphilophium,Haplolophium,
and Glaziovia,have an unusual"double" calyx with a thin frilly submarginal
rim in additionto a thick innermargin.Delostomaalso has a "double" calyx
with five irregularsubmarginalouter calyx teeth and an irregularlybilabiate
inner calyx. Clytostoma pterocalyx Sprague has an additional series of
enlarged wing-like lateral projections. Callichlamysis unique in its large,
thick, spongy calyx. Digomphiaand some speciesof Jacarandarepresentan
oppositeextremewiththin foliaceoussometimescompletelyseparatesepals.
There is often considerableintragenericvariationin calyx structureand
relativelyminor differenceshave sometimesbeen overemphasizedin generic
differentiation.This is especiallythe case with Macfadyenaand its relatives.
Macfadyenasensu stricto has a spathaceousor subspathaceouscalyx with an
inturnedapiculus;Doxantha (=Macfadyena) has a cupularnonspathaceous
calyx;Melloa has a subspathaceouscalyx with recurvedapiculus.Actuallythe
calycesof all threecan be so remarkablysimilarin form and texturethat it is
often difficult to tell them aparton the basis of calyx morphology.The same
kindof variationis found in Amphitecnawheremost speciesare variouslybil-
abiate to irregularlytruncatebut two newly discoveredspecies have spatha-
ceous calyces. Previouslyspathaceouscalyceswere consideredthe best floral
characterfor separatingits relativeParmentierafrom Amphitecna.It is not
even altogethercertainthat a givenspeciesof this groupalwayshas a constant
calyx form. I have seen aberrantspecimensof normallyspathaceous-calyxed
Parmentieraaculeatawith bilabiatecalycesand would not be surprisedto find
similar intraspecificvariationin Amphitecna. Yet this same differenceis a
fundamentalone in such other groupsas Phryganocydiawhich is very clearly
definedby spathaceouscalyces.


Flowersof differentBignoniaceae,all sharingthe common theme of the

sympetalousTubifloraeflower, neverthelessshow much morphologicaldiver-
sification.Flowertype is intimatelycorrelatedwith pollinationsyndromes(see
below). Corollasize rangesfrom the tiny 0.4 cm long flowersof many Tynan-
thus speciesto the up to 35 cm long flower of Tanaeciumjaroba Sw. and in
shape from the small narrowlytubularflowersof SchlegeliasulfureaDiels to
the largebroadlycampanulateflowersof Spathodeaand Kigelia.Most species
have variously tubular-campanulateto tubular-infundibuliformcorollas,
usuallyslightlybilabiatewith a pair of longitudinalridgesin the floor of the
tube and the uppertwo of the five corollalobes slightlysmallerthan the lower
three. Most of the species of Crescentieaediffer in having much reduced
corolla lobes and frequentlya horizontalfold across the floor of the corolla.
18 FloraNeotropica

Rathersubtle differencesin shape differentiatesome genera, but genera like

Tabebuiarunthe wholegamutfrom variouslycoloredtubular-campanulate or
tubular-infundibuliformbee-pollinated flowers to elongate tubular white
moth-pollinatedflowers, large,open-mouthedbat-pollinatedflowersor tubu-
lar redhummingbird-pollinated flowers.
Corollacolor, not preservedin herbariumspecimens,is surprisinglysig-
nificanttaxonomically.The basicdichotomyin coloris betweenpinkor purple
flowers and yellow flowers. Most basicallypurple-floweredgenera have no
yellow-floweredspecies and vice versa. There are hummingbird-pollinated
generaand species with orange and red corollas relatedto both yellow-flow-
ered and purple-floweredgroups. White flower color is even less significant
and white-floweredspecies are found both in otherwiseyellow-floweredand
otherwisepurple-floweredgenera. Even some species have both white-flow-
eredand purple-floweredvariants.
Corollapubescencemay be importantfor differentiatingboth generaand
species.For exampleall speciesof Arrabidaeahave the corolla pubescentout-
side (at least on the lobes) as contrastedto severalrelatedgenerawhich have
glabrouscorollas. Glandulartrichomesare usuallypresentat the level of sta-
men insertionnear the base of the corolla tube and are too constantto be of
much significancein most groups. In Tabebuiathe lengthof trichomesinside
the corolla throat is a useful specific character.Lepidotescales or glands are
usuallypresenton the inside of the lobes but presenceor absenceof lepidote
glandson the outside of the corolla tube is an importantspecific characterin
Anemopaegmaand Clytostomaandthey characterizethe genusMussatia.
Corollatextureis anothercharacterwhichdistinguishessome genera.For
example,Pithecoctenium,Amphilophium,and their relativesall have a very
thick-texturedcorolla (see pollinationecology, below). Howeversome species
of Lundiahavethickcorollasand othersthin corollas.


Thereare severaltaxonomicallyuseful kindsof stamenvariation.Differ-

ences in stamen numberare a very useful feature in genericlimitation. The
overwhelmingmajorityof the familyhas four didynamousstamens(occasion-
al teratologicalflowers may have five or six) but Catalpa (Tecomeae)and
Pseudocatalpa(Bignonieae)have only two stamens and several Old World
genera-Oroxylum, Rhigozum, Cactophractes,some species of Nyctocalos
and one species of Rhodocolea-have five stamens.Most Bignoniaceaehave
bithecateanthersbut one thecais lost in sectionMonolobosof Jacaranda.The
antherthecaemay be parallel,divergent,or divaricate.Cuspidariais uniform-
ly characterizedby havingeach theca bent nearthe base and reflexedforward;
one unrelated species of Pachyptera shares this unusual anther structure.
Antherpubescenceis also useful taxonomicallyand Lundiais best recognized
by its pubescentanthers;unfortunatelyseveralunrelatedspeciesof other gen-
era notably Cuspidaria,and Pachyptera, also have pubescent anthers and
Stizophyllumriparium(HBK.) Sandw. may have either glabrousor slightly
pubescent anthers. Saldanhaeawas formerly (and unnaturally)segregated
Morphology 19

from Cuspidariapartlyon the basis of antherpubescence.Once again a char-

acterwhichis definitivefor some generais highlyvariablein others.
In most Bignoniaceaethe fifth stamenis replacedby a smallvestigialpos-
teriorstaminode.In Jacarandaand Digomphiathe staminodeis greatlyelab-
oratedand usuallylongerthan the fertilestamens.In Digomphiathis reaches
an extremeand the antler-likestaminodeof one speciesis bifurcatedseveral
times; in anotherspecies this is variablewith both once and twice bifurcated
staminodes occurring. A few genera and species are characterizedby the
absenceof even a vestigialstaminode.


The bilamellatestigma and elongate style of most species and generaof

Bignoniaceaeare remarkablyuniform but the ovary provides useful taxo-
nomic characters.Ovaryshapereflectsfruit shapeand is often a good generic
determinant;relativelybroad ovate or ellipsoidovariestend to correlatewith
similarfruit shapesas do linearovarieswith linearfruits. A stipitateconstric-
tion below the ovaryis characteristicof Anemopaegma.Ovarypubescenceis a
very useful specific characterand even some generalike Tynanthusand Lun-
dia have uniformlyvillous ovaries. However,segregationof Neves-Armondia
(glabrousovary giving rise to a non-echinatefruit) from Pithecoctenium(vil-
lous ovarygivingriseto an echinatefruit)does not seemwarranted.
The ovariesof most Bignoniaceaeare bilocularwith a separatingseptum
and axile placentation.In the generawith axile placentation,each locule has
two parallellongitudinalplacentaeand the numberof seriesof ovules on these
is taxonomicallyuseful. Manygenerahave a singlerow of ovules on each pla-
centa, others have exclusivelymulti-seriateovules. However, in genera like
Tabebuiaboth conditionsmay occur. Thereis also a tendencyin multiseriate
generato have fewer ovule seriesat the top of the ovary so that ovary cross-
sections sometimegive very misleadingideas of numbersof series of ovules.
There is also a tendencyto have staggeredovules so that it is often unclear
whethera placentahas two seriesof widelyspacedovules or a single seriesof
closelyspacedovules. Numberof ovuleshas evenbeenused as a majorspecific
characterbut is veryvariable.Despitethe taxonomicsignificancetraditionally
attachedto numberof series of ovules, the characteris clearlynot an abso-
lute one taxonomically.
A few generahave an incompletepartitionperpendicularto the septum
whichgives the appearanceof a 4-loculedovary. This is especiallycharacteris-
tic of Tourrettiaand also occursto a lesserdegreein Astianthusand Hetero-
The most strikingvariationin Bignoniaceaeovariesis found in tribeCres-
centieaewherethe ovaries of Crescentiaare completelyunilocularwith four
parietal placentae while in Parmentierathe ovary is bilocular as in most
Bignoniaceae.Amphitecnais intermediatewith some species unilocularwith
two protrudingparietal placentae and others almost completely bilocular;
most speciesare bilocularat the base and unilocularat the apex dependingon
the degreeof fusion betweenthe protrudingplacentae.Degreeof placentalfu-
20 FloraNeotropica

sion is a useful specific characterbut is too variableto meaningfullyseparate

EnallagmafromAmphitecna.Exactlyparallelsituationsarefoundin Schlegelieae
and Coleeaeboth of whichincludebilocularand almostcompletelyunilocular
extremesplus variousintermediates.The incompletelybilocularplacentalar-
rangementwhich characterizesSchlegeliahas been termed a U-shaped pla-
centa (Leinfellner,1973), but its paralleloccurrencein two other unrelated
groupsclearlybelongingto Bignoniaceae diminishesits taxonomicsignificanceas
an indicatorof familialassignmentto Scrophulariaceae (Gentry,1976b).


A conspicuousnectariferousdisk surroundsthe base of the ovaryin most

Bignoniaceae.The form of the disk is quiteconstantand a good genericchar-
acter.Most generaof Bignoniaceaehave an annular-patelliform disk. In a few
genera-Cydista, Clytostoma,Phryganocydia-this disk is absent;loss of the
disk is correlatedwith multiplebang floweringphenologiesand pollinationby
deception(see below). Lundia and Tynanthuslack well-developeddisks but
have a densering of glandularnectar-producing trichomesaroundthe base of
the ovary.Melloais characterizedby a medialconstrictionin its disk whichhas
beentermeda "double" disk. Anemopaegmahas a conicaldisk contractingto
the stipitateovarybase. Gibsoniothamnusand Schlegeliahave the disk poorly
differentiatedfromthe baseof the ovary.


Fruitingcharactersarethe fundamentalones for subdivisionof the family

and have been emphasizedalreadyin discussingits taxonomichistory. To re-
capitulate,indehiscentfruits characterizeCrescentieaeand Schlegelieae(New
World) and Coleeae (Madagascarplus Kigelia of continental Africa). The
single speciesof Tourrettieaehas a bur-likeexozoochoricincompletelydehis-
cent 4-valvedcapsule.Eccremocarpeae, with a singlegenus, have a thin-walled
ovoid 2-valvedcapsulewithouta septum.Tecomeaeand Bignonieaehave de-
hiscent, usually two-valvedcapsules with the seeds borne on a septum; in
Tecomeaedehiscenceis perpendicularto the septum and in Bignonieaeit is
parallelto the septum(i.e. "marginicidal"),althougha few speciesof Bigno-
nieae (e.g. Melloa, Parabignonia,and MemoratanaeciicarpaA. Gentry)are
"4-valved"with each valvesplit down the midlineand are thus dehiscentboth
parallelto and perpendicularto the septum. At least one monotypicgenus of
Bignonieae,Dolichandra,has its fruitnormallysplit only perpendicularto the
septumbut even in Dolichandraoccasionalindividualshave incompletely4-
valved fruits. Thus tecomoid dehiscencein Dolichandraseems clearlya sec-
ondarymodification,especiallyas the only maturefruit I have seen of closely
relatedParabignoniais clearlyand completely"4-valved," and Dolichandra
should be transferredfrom Tecomeaeto Bignonieae.Converselymonotypic
and isolatedPerianthomegaof Brazilhas fruitdehiscenceperpendicularto the
septumand probablybelongsin Tecomeae.
Morphology 21

More problematicalare the Old World Oroxyleaewith fruit dehiscence

parallelto the septum. I suspect that these clearlyrelict, mostly monotypic,
generaaremorecloselyrelatedto Tecomeaethanto Bignonieae,althoughthey
may well representsurvivorsfrom the commonmatrixwhichgave rise to both
majortribesof the family. A Tecomeaeaffinity is supportedby the phytogeo-
graphicaland other morphologicalinformation. The previouslyunreported
fact that at least one of these species-Nyctocalos cuspidata,Barthes.n. (P)-
has 4-valved fruits also suggeststhat the bignonioiddehiscenceof the other
speciesof Oroxyleaemay not be as fundamentalan indicatorof relationship
with the otherwiseNew World Bignonieaeas has been previouslysupposed.
The discoverythat a numberof speciesof Bignonieaescatteredthroughvari-
ous generahave 4-valvedfruits also supportsthis conclusion, as does dehis-
cence parallelto a "pseudoseptum"in Dolichandroneand Markhamiaof the
At the generic level fruits are also of great taxonomic importance,al-
thoughmore poorly collectedthan flowersand as yet unknownfor many spe-
cies and a few genera. Especiallyimportanttaxonomicallyis the differencein
shape and thicknessbetweenthe relativelybroad woody-valvedfruits of such
generaof Bignonieaeas Pithecoctenium,Amphilophium,Distictis,Distictella,
Clytostoma, Tanaecium, Melloa, and Adenocalymma and the narrow
elongate-linearones of generalike Arrabidaea,Macfadyena,and their allies.
Unfortunatelyfor straightforwardkey construction,othergeneramay encom-
pass both extremes-the same patternof variabletaxonomic significanceof
any given morphologicalcharacterin differentgroupswhichis so prevalentin
the Bignoniaceae.Thus superficiallydifferentfruit forms can best be treated
as congenericin generalike Memoraand Pachyptera,contraryto taxonomic
tradition.Such intragenericvariationbetweenlinear and broad fruits is usu-
ally correlatedwith a switch from wind-dispersedto water-dispersedseeds,
and is clearlynot an indicatorof majorphylogeneticdivergence.
In Crescentieaedifferencesin shape betweenglobose fruits and ovoid or
ellipsoid apiculate fruits are taxonomicallysignificant,especiallyin Amphi-
tecna. Howeverthe taxonomicpictureis somewhatblurredby the occurrence
in the West Indies of ellipsoid subapiculate-fruitedvariantsof Amphitecna
latifolia, along with the globose fruits which distinguishthe species from its
congenerselsewhere. This greatervariabilityis correlatedwith the species'
greaterhabitat range in the West Indies (and Venezuela)-elsewhereit is re-
stricted to coastal swamps and seashores-and is a well known island
Fruit surface featuresalso provide useful taxonomic characters.A few
genera like Pithecocteniumand Clytostoma are characterizedby echinate
fruits. In other generalike Amphilophium,Mansoa, and Arrabidaeatubercu-
late or echinatefruits characterizesome speciesbut not others. Longitudinal
ridges, wings, or furrowsare typical of severalgeneraincludingCuspidaria,
Piriadacus,and Tynanthus.In other genera only isolated species may have
such features(e.g. CydistaheterophyllaSeib. (medianfurrow),Parmentiera
macrophyllaStandl. and AmphitecnacostataA. Gentry(longitudinalridges),
and AdenocalymmapurpurascensRusby (longitudinalwings)). Pubescent
22 FloraNeotropica

fruitsare a good speciescharacterand are typicalof most (but not all) species
of suchgeneraas LundiaandDistictella.


Seedsof the greatmajorityof Bignoniaceaeare winged;even in a few in-

dehiscentfruited generalike Parmentieravestigialseed wings remain. Many
generaof Bignoniaceaehave very characteristicseeds which are distinguished
by presenceor absenceof wings, thicknessof seed body, wing texture(hyaline
membranaceousto opaque and subwoody), wing structure(usually a fused
membranebut composed of distinctcapillaryhairs in Catalpaand Sparatto-
sperma),and wing placement(usuallybialatebut variouslyencirclingthe seed
body in genera like Anemopaegma, Pithecocteniumand Jacaranda).Am-
philophium and Distictis are unusual in a pubescent seed body. Medially
folded cotyledons are an interestingseed modification found in a few Old
It has been taxonomicallytraditionalto considerthe presenceor absence
of seed wings and the concommitantthickeningof the seed body in wingless
seeds as an absolutecharactermandatinggenericdifferentiation.However, I
have demonstrated(1973a)that thesedifferencesmerelyreflectadaptiveshifts
from wind to water dispersalwhich have occurredin numerous unrelated
groupsand may not alwayswarranteven specificrecognition.Thusseparation
of water-dispersedCouralia from wind-dispersedTabebuia,wind-dispersed
Doxantha from water-dispersedMacfadyena, and wind-dispersedPseudo-
calymmafromwater-dispersed Pachypterais unwarranted.

As summarizedby van Steenis (1978) Bignoniaceaeseedlingstend to be
ratheruniform in all tribes. Howeverthere is apparentlya fundamentaldis-
tinctionwithintribes Bignonieaeand Crescentieaebetweenepigealand hypo-
geal germination.Apparentlyall Tecomeaehave epigealgermination.In gen-
eral, speciesor generawith thick-bodiedseedshavehypogealgerminationwith
the cotyledonsremainingenclosedin the seed coat while the majorityof spe-
cies with thin-bodiedseeds have the photosyntheticleaf-like cotyledons ele-
vatedabove groundlevel by an elongatinghypocotyl.In Crescentieae,Amphi-
tecna with relativelylarge thick-bodiedseeds has hypogealgerminationwhile
Crescentiaand Parmentierahave epigealgermination.In BignonieaeTanae-
cium, Distictella,Amphilophium,Memora,Martinella,Melloa, Phryganocy-
dia, Pithecoctenium,and PyrostegiahavehypogealgerminationwhileLundia,
Pleonotoma, and Macranthisiphonare epigeal. This differencemay be some-
what facultativeas indicatedby Callichlamyswhichhas photosyntheticcotyle-
dons whichsometimesremaintrappedin the seed coat and are sometimesele-
vated. In Arrabidaeathe preliminaryevidencesuggeststhat there is a poten-
tially taxonomically significant dichotomy in the genus between relatively
Pollen Morphology 23

large-seeded hypogeal species like A. corallina (Jacq.) Sandw. and A. mollissi-

ma (HBK.) Bur. & K. Schum. and the majority of epigeal species.
The first leaves are nearly always simple. I have seen compound first
leaves only in Pleonotoma, Jacaranda, and Pyrostegia. Most genera of Tecomeae
have the first leaves conspicuously dentate, even when mature leaves are
entire. Bignonieae may have entire or serrate first leaves. Schlegelia seedlings
are especially interesting in having the first leaves distinctly 3-lobed, perhaps
reflecting the old "ontogeny recapitulates phylogeny" theme and supporting
the relationships of this genus of exclusively simple and entire mature leaves
with predominantly compound-leaved Bignoniaceae.


Bignoniaceae include a wealth of palynological diversity, and pollen mor-

phology has long been important for the family's taxonomy, especially in
generic delimitation. Recent summaries of Bignoniaceae pollen and its taxo-
nomic implications have been made by Buurman (1978) focusing on Asiatic
genera, and by Gentry & Tomb (1979) concentrating mostly on New World
genera. Table I (from Gentry & Tomb, 1979) and Figs. 1-2 summarize the pol-
len types of Bignoniaceae. Apertures range from the basic and most prevalent

Table I
New World Bignoniaceae pollen types (from Gentry & Tomb, 1979)
Pollentype Genera
Psilateandmicroperforate, Bignonieae:Arrabidaea,Ceratophytum, Dolichandra,
3-colpate(4-6-colpatein Fridericia,Leucocalanthe,Macfadyena,Manaosella,
Leucocalanthe) Melloa,Parabignonia,Paragonia,Paradolichandra,
Eccremocarpeae: Eccremocarpus.

Psilate,3-colpatetetradsor Cuspidaria

Finelyreticulate,3-colpate(to Bignonieae:Callichlamys,Lundia,Mussatia,Pachyptera
4-colpatein Pyrostegia (exceptP. alliacea),Piriadacus,Pleonotoma,
venustaandRoentgenia) Pyrostegia,Potomoganos,Stizophyllum,Tanaecium
(exceptT. apiculatumand T. nocturnum),Roentgenia
(alsoall Old Worldgeneraexceptsomespeciesof Incarvillea
24 Flora Neotropica

Table I (continued)

Pollen type Genera

Areolate/perisyncolpate Bignonieae: Gardnerodoxa, Mansoa, Memoraflavida,

M. pedunculata, M. cristicalyx, M. imperatoris-maximilianii,
Tecomeae: Delostoma, Perianthomega, Sparattosperma.

Areolate tetrads Catalpa, Chilopsis

Inaperturate, medium-reticulate Bignonieae: Clytostoma binatum, C. pterocalyx, Cydista

aequinoctialis (mostly), C. potosina, Phryganocydia.

Inaperturate, coarse-reticulate Bignonieae: Adenocalymma, Distictella, Distictis, Memora

(except M. flavida group), Pitheococtenium, Tanaecium
apiculatum, Clytostoma costatum, C. sciuripabulum,
C. uleanum, C. convolvuloides (interrupted exine)

Stephanocolpate (zonocolpate), Bignonieae: Amphilophium, Anemopaegma robustum,

coarse-reticulate A. insculptum, Glaziovia, Haplolophium, Urbanolophium.

3-colpate, coarse-reticulate Martinella

Polyporate, finely reticulate Amphitecna

Polyporate, coarse-reticulate Tanaecium nocturnum

Spiroaperturate Neojobertia

Pericolpate (pantocolpate) Anemopaegma (mostly), Cydista decora, C. lilacina,

C. diversifolia, Pachyptera alliacea

Spinulose, inaperturate Cydista heterophylla (several unrelated species of other

genera-Phryganocydia, Roentgenia, Adenocalymma,
Clytostoma, Memora-also show tendency in this direction).

Polyporate, complex exine Macranthisiphon, Saritaea


Ecolpate, 3-porate Schlegelia (small-flowered species)

Not examined: Romeroa, Ekmanianthe, Synapsis

3-colpate condition to monocolpate and inaperturateon one hand and to 4-5-

colpate, zonocolpate, pericolpate, spiroaperturate, and variously syncolpate
types. Various porate forms are also known. The exine ranges from psilate to
finely reticulate, coarsely reticulate, spinulose, or areolate and may be tectate,
semitectate or intectate. Most genera have pollen monads but Catalpa, Chilop-
sis, and Cuspidaria have tetrads and one species of Cuspidaria has polyads.

FIG. 1. Bignoniaceae pollen diversity. A, 3-colpate medium-reticulate (ancestral pollen type),

Pyrostegia dichotoma. B, 3-colpate coarse-reticulate, Martinella obovata. C, polyad, Cuspidaria
bracteata. D, 3-colpate, psilate-foveolate, Xylophragma seemannianum. E, inaperturate coarse-
reticulate, Distictella magnoliifolia. F, inaperturate medium-reticulate, Cydista aequinoctialis. G-
Pollen Morphology 25

~:~~~~~~~~~~~~~~~~~~~ .4
....;~:'~... '~: - .:":;::
:'il...- ,.,.-- - -
: ' ' . '. '
......: : ,? ,

: 't?
..... A-t::'..
': . ...
? <;it}?J i ^
::..: :

: F':~~ :/ ~~4: :, :
:;:.:.,,,~::::,,: ~iii-.':i. :.i:

. .. .?... ~~ r~ij*~: .
? . .....:
?:.''';: :.:... .:.:' ..
=============..: .
. ... -
?: .

: ... ..
----x~F ~~~~~~~4
?: it:

ca ?
i:.':, ' :"~'
~ :::..
'_,':.. :...'~.~~.,
'i::':~ t
7 ' ' i i-' Ili i~~~~~~~~~~~~~~~~? '::i

'A*~ s -,, ;t""


All lines 10 m. (From Gentry & Tomb, 1979).
~ ~ ~ ' . : :::.:::::.
B. :: C~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~<
.?-Y .:::.:::.....

,?, ?:
~~~..:.... ::~'::*':'~:

H, t~pan?olat (zno?lpte)?orsertclt,Alhllh~ ~?ltm ,sboa

view H, ~ e yoraeMacrathisiho v Iop
qaoiscal eou.%o w I J,srit-rt,ei
anthmegavelozoi cano!lana.L, reolte, parttoserm
euatu.Allns? 0 K,spirapcrurae,
~.(Foetr ob,17)Nojoerti

H, stephanocolpate(zonocolpate)coarse-reticulate,
Amphilophium paniculatum;-G, subpolar
view.I, polyporate,
view;H, equatorial longiflorus.J, striate-areolate,Peri-
anthomega candolleana.
vellozoi.K, spiroaperturate,Neojobertia L, areolate, Sparattosperma
26 Flora Neotropica

li~ l. lii!l~~.....

t. ........

FIG. 2. Pollen of Crescentieaeand Tourrettieae.A, Tourrettialappacea,psilate-foveolatewith

tectatecolpi. B, Amphitecnalatifolia,multiporatefinelyreticulate.C-D, Crescentia;C, C. cujete,
6-colpate with equatorial partition of primary colpi; D, C. alata, 3-colpate. E, Parmentiera macro-
phylla, 3-colpate. All lines = 10 nm.

Most of the specialized pollen types have evolved independently in different

unrelated groups within the family, usually in both Bignonieae and Tecomeae.
Individual genera may encompass palynological variability but show single
well-developed palynological trends-e.g. 3-colpate to perisyncolpate to multi-
porate to inaperturate(in Tanaecium), in general exine sculpturing is more con-
servative than number and arrangement of colpi and few taxa, notably Clyto-
stoma, show significant intrageneric exine variation. 3-colpate finely reticulate
pollen seems to be the ancestral type in the family. Pollen is taxonomically ex-
tremely useful in generic delimitation if allowance is made for the possibility of
a certain amount of intrageneric variability.


The chemotaxonomy of the Bignoniaceae has been thoroughly reviewed

by Hegnauer in van Steenis' (1978) treatment of Bignoniaceae for Flora Male-
Chemotaxonomy 27

siana. This summaryis largelyextractedfrom Hegnauer'sreview.Phytochem-

ically the family forms a cohesivegroup (along with severalother Tubiflorae
families). Iridoid glucosides are apparentlypresentin most membersof the
family and iridoid alkaloids are known from several genera of Tecomeae.
Naphthaquinonesand anthraquinonesare prevalentin the family, at least in
Tecomeaeand Coleeae. Lapachol,which forms a conspicuousyellow, some-
times skin-irritatingpowder in the wood, is especially frequent. Complex
esters and glycosides of o-diphenoliccinnamicacid derivativesreplace true
tanninsin Bignoniaceaeand simpleestersof caffeic acid and of derivativesof
cinnamicand benzoicacids are often present,also. Leaf and petal flavonoids
are mostly flavones ratherthan flavonols, a common patternin Tubiflorae.
The common anthocyanincyanidin-3-rutinoside and various carotenoidsare
the major agents of petal coloration of a wide array of genera (Harborne,
1967);Harborne'sdata suggesta basic dichotomybetweenpetal colorationof
Bignonieae and Tecomeae, the former having carotenoids and the latter
and carajuronewhichare responsiblefor the red colorationof driedleaves of
Arrabidaeachica (H. & B.) Verl. Jacaranone,a quinonoidprobablyderived
from tyrosine,has been isolated from leaves and twigs of Jacarandacaucana
Pitt. and is especiallynoteworthybecause of its marked antitumoractivity
(Oguraet al., 1977).
Althoughformalchemicalinvestigationsof Bignoniaceaehave barelybe-
gun, manyotherunstudiedbut phytochemistry-related featuresof certaingen-
era and speciesof Bignoniaceaehavealreadyprovedtaxonomicallyimportant.
For example, many genera, especiallyof Bignonieae,can be recognizedby
their odors. Tynanthuswood and (usually)leaves have a strong clove odor.
Mansoa and Pachyptera(includingPseudocalymma)have many specieswith
strong onion or garlic odors. Tanaecium,which is poisonous to cattle (Du-
gand, 1942) and has hallucinogenicproperties(Prance, 1977) has a strong
almond or cyanide vegetativeodor. Paragoniahas a characteristicsweetish
odor when fresh. Godmaniahas wood and barksmellinglike horseurine,and
some Tabebuiaspecieshavebarkwith a watermelon-likeodor. Odorscan also
be taxonomicallyimportantat the specific level. For examplethe two North
Americanspeciesof Catalpacan be distinguishedby the rankodor peculiarto
C. bignonioides.Closely relatedPachypterahymenaea(DC.) A. Gentryand
P. standleyi(Steyerm.)A. Gentrycan be differentiatedby a garlic-likeversus
an onion-likeodor. Clearlythe family is ripe for additionalchemosystematic


Bignoniaceaeare remarkablyconstantcytologically(TableII). By far the

most frequentbase numberis x = 20 with 2n = 40 the only chromosome
numberknownin tribesTourrettieaeand Coleeaeand the overwhelminglypre-
dominateone in the largerand more diversetribes Bignonieae,Crescentieae
and Tecomeae.The majordiscrepancyfrom 2n = 40 is found in the smallex-
28 FloraNeotropica

Chromosomenumbersin Bignoniaceae.FromGoldblatt&Gentry,1979.
All countsareconvertedinto diploidnumbers.
Species Diploid Number

Argyliauspallatensis 30

Campsisgrandiflora 40
C. radicans 40
C. x tagliabuena 40

Catalpabignonioides 40
C. ovata 40
C. speciosa 40
C. x erubescens 40

Chilopsislinearis 40

Delostomalobbii 42

Dolichandronespathacea 40
D. stipulata 40

Heterophragma 40

Incarvilleacompacta 22
I. delavayi 22
I. maireivar.grandiflora 22
I. olgae 22
I. semiretschenskia 22
I. sinensis 22

Jacarandacoerulea 36
J. hesperia 36
J. mimosifolia 36

Markhamiahildebrandtii 40
M. platycalyx 40

Newbouldialaevis 40 (38)

Pajanelialongifolia 40

Pandoreajasminoides 38

xylocarpa 40

Spathodeacampanulata 26, 36, 38

Stereospermum chelonoides 40
S. kunthianum 40
S. personatum 40

Tabebuiachrysantha (38)
T. guayacan 40
Cytology 29

TableII (continued)

Species Diploid Number

T. heptaphylla 40
T. heterophylla 40
T. impetiginosa 40
T. "pentaphylla" 40
T. rosea 40
T. serratifolia 40
T. "spectabilis" 40

Tecoma garrocha 36
T. sambucifolia 36
T. x smithii 36
T. stans 36
T. stans var. velutina 36

Tecomaria capensis 36

Tecomanthe dendrophila 36
T. speciosa 38


Millingtonia hortensis 30

Oroxylum indicum 28


A mphilophium paniculatum 40

Anemopaegma chamberlaynii 40
A. orbiculatum 40

Arrabidaea corallina 40

Bignonia capreolata 40

Callichlamys latifolia 40

Clytostoma binatum 40

Cydista aequinoctialis 40, 40 + 1-2 B

C. diversifolia 40

Distictis buccinatoria 40

Lundia corymbifera 40

Macfadyena unguis-cati 40, 80

Mansoa difficilis 36 (-38)

Martinella obovata 40
30 FloraNeotropica

TableII (continued)

Species Diploid Number

Memora patula 40

Pachyptera hymenaea 38

Paragonia pyramidata 40

Phryganocydia corymbosa 40

Pithecoctenium crucigerum 40

Pleonotoma jasminifolia 40

Pyrostegia cinerea 40
P. venusta (60)

Saritaea magnifica 40

Stizophyllum riparium 40
S. perforatum 40

Tanaeciumjaroba 40

Xylophragma seemannianum 40


Kigelia africana 40

Phyllarthron comorense 40


Amphitecna latifolia 40
A. montana 40

Crescentia cujete 40

Parmentiera aculeata 40
P. cereifera 40
P. macrophylla 40


Tourrettia lappacea ca. 40


Schlegelia parviflora 40
Cytology 31

clusively Old World group of four genera which are often treated as belonging
to Bignonieae but might better be separated as tribe Oroxyleae. On the basis of
retention of five stamens in two of these genera, they might be suggested as
among the most primitive in the family. Thus the 2n = 28 count in 5-stamened
Oroxylum (Goldblatt, 1976) along with the frequency of n = 20 is highly sug-
gestive of x = 7 as basic to the family, a conclusion compatible with accumu-
lating evidence from other angiosperm families (Raven, 1975). Most of the
family's evolutionary radiation apparently took place subsequent to an initial
aneuploid loss of one chromosome from a palaeohexaploid stock. Only the
Oroxyleae remain at an ancestrally tetraploid level (Millingtonia with 2n = 30
can best be interpreted as an aneuploid addition of a single chromosome pair
and Nyctocalos and Hieris are uncounted.) The two highly derived herbaceous
genera Argylia and Incarvillea, both belonging to Tecomeae, have x = 15 and
x = 11 respectively and might involve aneuploidy at the tetraploid level. Most
of the rest of the reported counts other than 2n = 40 are probably in error. We
have checked several of the anomalous genera and most of them prove to have
2n = 40 (Goldblatt & Gentry, 1979). However, one group of Tecomeae which
includes Tecoma of the New World and Tecomaria, Tecomanthe, and Pan-
dorea of the Old World, clearly has 2n = 36(-38) (Goldblatt & Gentry, 1979),
and the counts of 2n = 36 in Jacaranda are probably also correct. Delostoma
with 2n = 42 may be a relict of the putative palaeohexaploid stock. Polyploidy
is almost unknown in the family with a single reported case in Macfadyena
unguis-cati which has been reported as both 2n = 40 and 2n = 80. Cydista
aequinoctialis apparently contains B chromosomes besides its normal comple-
ment of 40.
The predominence of n = 20 in all tribes except Oroxyleae indicates a
fairly close relationship of these tribes and a common ancestry in a single evo-
lutionary line. The uniformity of 2n = 40 in Bignoniaceae makes chromosome
number a potentially useful taxonomic tool in evaluating the status of such
problematical genera as Schlegelia and Paulownia, both of which are often
placed in Scrophulariaceae. Thus the occurrence in both Paulownia and
Schlegelia of 2n = 40 supports their placement in Bignoniaceae. All apparent
trends-large diploid chromosome numbers, constancy within a family, lack
of polyploidy-are concordant with those expected for woody tropical plants.


Phylogenetic conclusions in Bignoniaceae are rarely obvious and are

fraught with danger because of rampant parallelisms and convergence in
nearly every taxonomically important characteristic of the family. For exam-
ple nearly all of the highly diversified pollen types of the family have evolved
independently in unrelated groups (Gentry & Tomb, 1979) and indehiscent
fruits evolved separately in at least two different lineages (Gentry, 1976b).
A few broad phylogenetic generalizations may safely be hazarded, how-
ever. The Tecomeae and Oroxyleae are closest to the ancestral stock of Bignoni-
32 FloraNeotropica

aceae.Perhapsan arborescentOroxyleae-likeancestorwith five stamens,fine-

ly reticulate3-colpatepollen, pinnatelycompoundleaves and a chromosome
numberof 2n = 28 gave riseto a hexaploidcommonancestorof the restof the
family which would probablybe referredto Tecomeaeif it were alive today.
Delostomawith 2n = 42 may be derivedfrom this stock. This ancestralplant
lost one stamen and a single chromosomepair giving rise to a didynamous
common ancestorof most of the family with 2n = 40. The change from the
fruit dehiscenceperpendicular to the septumwhich characterizesTecomeaeto
the parallel-to-the-septumdehiscence of Bignonieae took place in South
Americasubsequentto separationfrom Africaand was accompaniedby devel-
opmentof lianahabit, leaf tendrils,and anomalouswood anatomy.A second-
aryreversionto 4-valvedfruitsand finallydehiscenceperpendicularto the sep-
tum as in Tecomeaetook place in one group of otherwisetypicalBignonieae.
The SouthAmericanTecomeaegave riseto a palmatelycompound-leaved
form similarto Tabebuiafrom whichmost of the neotropicalmembersof that
alliance are descended. Palmately compound-leavedneotropical Tecomeae
progenitorsalso gave rise to the Crescentieaewith evolutionof an indehiscent
fruitaccompaniedby bat pollination.On Madagascar(or Afromadagascar?)a
pinnately-leaveddehiscent-fruitedancestorreferableto Tecomeaegave rise to
another group of indehiscent-fruitedBignoniaceae,the Coleeae. The two
monogenericAndean-centeredtribes Tourrettieaeand Eccremocarpeaeare
herbaceousor wiry vines with leaf-tendrilsand seem closest to Bignonieae,
despiteverydifferentfruitsand placentation,and may have evolved from the
ancestralplexusof that tribe.No clearaffinitiesare apparentbetweenSchlege-
lieae and the rest of the familyand they presumablyhave a long historyof in-
dependentevolution dating back to the common ancestorsof Bignoniaceae
and Scrophulariaceae.
Withineach of thesemajorgroupsparallelsubsidiarytrendstowardscan-
dent habit (Oroxyleae,Tecomeae, Bignonieae, Crescentieae(if Amphitecna
parviflorais really scandent)),pollen with coarselyreticulateor psilate exine
(Tecomeae,Bignonieae,Schlegelieae,Oroxyleae)and multicolpateor inaper-
turate (and eventually areolate or perisyncolpate)apertures (Tecomeae,
Bignonieae,Crescentieae,Schlegelieae),pollen tetrads(Tecomeae,Bignonieae),
androeciumreductionto two fertile stamens (Tecomeae, Bignonieae),aneu-
ploid loss of chromosomes (Tecomeae, Bignonieae), spines (Crescentieae,
Coleeae), etc. took place independently,sometimes in different alliances
withinthe sametribe.


Bignoniaceaepollination strategiesare extremelydiversifiedand are be-

ginningto be relativelywell understood(Gentry, 1974a, 1976a). Interrelated
aspects of pollination biology of this family include floral morphologyand
type of pollen vector attracted,floweringphenology, and seasonality.More-
over the apparent niche specificity of neotropical Bignoniaceaeflowering
strategiesappearsto have significancefor communityecology. Thesedifferent
aspectsof Bignoniaceaepollinationecologywill be discussedseparately.
Ecology 33

Floralmorphologyand pollenvectors

Bignoniaceaeflowers may be separatedinto ten major morphological

typeswhichcorrelatewith theirpollinationagents(Gentry,1974aand Figs. 3-
4). Most of the speciesof Bignoniaceaehavetubular-campanulate flowerswith
the mouth of the tube oval and open and the anthersincluded. These are
Anemopaegma-typeor "open-mouthedbee flowers." These species are bee-
pollinatedfor the most part and this morphologyis presumablythe basic
flowertype of the family, beingnot only the most widespreadflowerform but
also the only one of the ten morphologicaltypeswhichis found in four differ-
ent tribes.Thereis a significantrangeof variationwithinthis groupof plants.
Some specieshave theircorollaspubescentoutsideand some corollasare glab-
rousor lepidoteoutside. Some specieshaveshortmembranaceouscorollasand
others longer and generallythicker-texturedcorollas. This group thus inter-
gradesat one end of its spectrumwith the "bee/butterflyflowers" and at the
otherend withthe "xylocopidflowers."
Tynanthus-typeor "bee/butterfly flowers" have generallysmall short
corollaswithopen mouthsand subexsertedanthersand tend to be pronouncedly
bilabiate.These flowersare all puberulousor lepidoteoutside and the corolla
lobes tend to be almostvalvatein bud. Theyareveryfragrantand attractlarge
numbersof both small bees and butterflies, both of which are apparently
effective pollinators.The corolla mouth of these flowersis open and the but-
terfliesare apparentlyeffective pollinatorssince both their proboscisand the
upperpartof theirtongueoften makecontactwiththe anthers.
Pithecoctenium-typeor "xylocopid flowers" have a greatly thickened
corolla, especiallyat the base of the tube and have a thickenedridgewith espe-
ciallydensetrichomesat the levelof stameninsertionwithin.All of these flow-
ers are highlypuberulouswithoutand most of them have a bend in the middle
of the corollatube. The calycesof these flowersare also verythickand protec-
tive. Xylocopidsareamongthe most frequentvisitorsto flowersof Bignoniaceae
but aregenerallyrobberspenetratingthe corollatube at its base from outside;
modificationswhich force these bees to become legitimatepollinatorshave
obvious adaptivevalue. Paragoniais of interestin being halfwaybetweenthe
"xylocopidflowers" and the "open-mouthedbee flowers." In Paragoniathe
corollais thickerthan in the lattergroupbut thinnerthan in the other species
of the former group. SignificantlyI have observedvisits by two species of
xylocopidto Paragonia,one of whichpenetratedthe corolla base in the tradi-
tional manner,but the second of whichenteredthe tube as a legitimatepolli-
nator. Paragoniais also intermediatebetweenthesetwo groupsin a numberof
A directmodificationof the xylocopidflowermust have given rise to the
uniquepseudocleistogamousAmphilophium-typeflower. A logical extension
of the trendtowardgreaterprotectionof the nectarmight have led to the un-
opened Amphilophium-typeflower in which not even the pollen is exposed
exceptto legitimatepollinators.CertainlyAmphilophiumand Glazioviahave
the thickestcorollas of all the Bignoniaceaeand their secretorydouble calyx
attracts ants and wasps which presumablygive further protection to the
flower. But the reallyuniquefeatureof this type of floweris its failureto open
34 Flora Neotropica



MX, , NX


FIG. 3. Bignoniaceae floral morphology. A-F, Anemopaegma-type; A-D, Anemopaegma orbicu-

latum; A, corolla in top view; B, calyx and pistil (stigma lobes closed); C, corolla split open to
show stamen placement; D, ovary and disc. E-F, Arrabidaea candicans; E, flower in top view; F,
flower in side view. G-H, Tynanthus-type, Tynanthus croatianus; G, flower in front view; H,
flower in side view. I-K, Pithecoctenium-type, Pithecoctenium crucigerum; I, flower in side view;
J, flower in bottom view; K, pistil, disc, and section through calyx. L-M, Amphilophium-type,
Amphilophium paniculatum; L, flower in top view; M, flower in side view. N-Q, Cydista-type; N-
O, Cydista heterophylla; N, corolla in top view; 0, ovary and part of calyx; P-Q, Arrabidaea ver-
rucosa; P, corolla in top view; Q, corolla in side view. (From Gentry, 1974a).
Ecology 35

y--~ H

FIG. 4. Bignoniaceae floral morphology (continued). A-B, Tanaecium-type, Tanaecium noc-

turnum; A, corolla in bottom view; B, corolla in side view. C-E, Pyrostegia-type; C-D, Martinella
obovata; C, corolla in top view; D, calyx and pistil; E, Pachyptera standleyi, flower in top view;
F-I, Crescentia-type; F-G, Crescentia cujete; F, flower in top view; G, corolla in side view; H-I,
Amphitecna latifolia; H, corolla in side view; I, flower in bottom view. (From Gentry, 1974a).
36 FloraNeotropica

at anthesisuntil the two lips are physicallyforcedapartby a largeand robust

bee, hencethe termpseudocleistogamous.In the herbariumthe two lips of the
corolla are forced open by compressionof the flower duringpressing,so that
this uniqueadaptationhas beengenerallyoverlooked.
Another group of species with "dorsally-compressedbee flowers" or
Cydista-typeflowersshowsa differentset of modificationsfrom the basicpat-
ternof floralmorphology.Theseplantsare evidentlyall bee-pollinated.In this
groupthe corollatube is dorso-ventrallycompressedso that the corollamouth
is loosely closed or very narrow.Manyspeciesof this grouphave conspicuous
nectarguidelines, thesebeingusuallya numberof finelypencilledmoreor less
parallel magenta or brick-redlongitudinallines convergingfrom the lobes
onto the two abaxialfolds of the corollatube. Most of the speciesin this group
havethe corollatube glabrousor only inconspicuouslypuberulouswithout;all
haverelativelythin corollas;the corollais generallymoreevenlyfunnel-shaped
(ratherthan campanulateabove a tubularbase) than in other species of the
family. Most of the genera included here (Cydista, Clytostoma, and Phry-
ganocydia)lack a hypogynousnectariferousdisk and attractvery few visitors
as discussedunderphenologybelow. Arrabidaeaverrucosa(Standl.)A. Gen-
try and its allies are of specialinterestin beingintermediatein severalrespects
betweenthe "dorsally-compressed bee flowers" and other groups. They have
a pubescentcorollatube and somewhatthickertexturedcorollathan the other
One of the most spectacularmodificationsof floral morphologyin the
Bignoniaceaeis that of the "hawkmothflowers." This strikingmorphological
type has been evolved independentlyin the tribes Tecomeaeand Bignonieae
and may be termedthe Tanaecium-type.These flowershave a relativelythick
andrigidcorolla,whichis whiteand has the tubenarrowand greatlyelongated.
The anthersare usually more or less exsertedand the thecae tend to be long
and flexuous. I have not actuallyobservedvisitsby mothsto these flowers,but
their shape, color, and heavy sweet aromaare typicalof the hawkmoth-polli-
nationsyndromein otherfamilies.
Pyrostegia-typeor "hummingbirdflowers" have also evolved independ-
ently in tribes Tecomeae, Bignonieae, Eccremocarpeae,and Tourrettieae.
These flowersare odorlessand brightred-orangeor verydeep violet (in Tour-
rettiaonly the sterileflowersarered). The corollais ratherthick in texture,the
corollamouthis open and the anthersare often exserted.The corollatube (ex-
cept Tourrettia)is glabrousoutside. The flowers are rathernarrowlytubular
althoughin Martinellathe uppercorolla tube is abruptlyexpandedto fit the
headratherthanthe bill of its pollinator.
A very different adaptationto bird pollinationis shown by generalike
Spathodeaand Fernandoa.Nativeto the Old Worldtropicswhereno equiva-
lent of the hover-feedinghummingbirdsexist, they have evolvedeitheran en-
larged, reflexedcalyx or reflexedflower position. The calyx or pedicelserves
as a convenientperch from which a pollinatingbird can reach into the peri-
pherallylocatedopen flowers. In brightred-orangecolor, lack of a noticeable
fragrance,and generallack of corolla pubescence,these plants approximate
their hummingbird-pollinated New World counterparts.Tecomaria,another
Ecology 37

Old Worldbird flower from southernAfrica, is pollinatedby sunbirds(Scott-

Elliot, 1890) and is morphologicallycloser to the "hummingbirdflowers"
thanto those of the Spathodeatype.
The final group of pollen vectorsfor speciesof Bignoniaceaeare bats. In
the New WorldCrescentia-type or "bat flowers"haveevolvedonly in the tribe
Crescentieae,but in the Old Worldthey are presentin Oroxylum,Kigelia,and
speciesof Tecomeae.The New World"bat flowers"have the corollawhite or
greenish-white,thick in texture,and usuallywith a transversefold across the
throat behindwhich is formeda nectar-storingbulge. These flowershave the
characteristicmustyodor, reminiscentof the bats themselves,whichhas often
been remarkedas typical of the bat pollination syndrome. All of the New
World "bat flowers" have the corolla glabrousexcept for some stalked-lepi-
dote trichomes.The flowers of most of these species are cauliflorouson the
trunkor ramiflorouson thickertwigs. The corolla is ratherwidelycampanu-
late withreducedlobes and the anthersand stigmaaresubexserted.
Old World "bat flowers" which are sometimes cultivatedin the neo-
tropics, (Kigelia,Haplophragma),contrastin severalrespectswith the native
New World "bat flowers." They have much largercorollas than their New
Worldcounterpartsas befits theirMegachiropteran pollinators(Gould, 1978;
Baker & Harris, 1959) as contrastedto the much smaller Microchipteran
(familyPhyllostomidae,subfamilyGlossophaginae)flower-visitingbats of the
neotropics.The Old World "bat flowers" also differ in being dully colored
ratherthan whitish and in being borne flagelliflorouslyon danglingleafless
branches(Kigelia)or pin-cushionstyle on thick-branched,stiff inflorescences
projecting out from the canopy (Oroxylum, Haplophragma,etc.). Their
muskyodor is the sameas that of the New World"bat flowers."


Speciessharingthe same patternof floral morphologyand same pollina-

tors may still have different pollination strategies.This is possible through
phenologicalspecializationswhich enable differentialuse of the same polli-
nators (Gentry, 1974a).Thus "steady-state"speciesof Bignoniaceaeput out
one or two flowersa day over a long periodof time and are pollinatedby trap-
liningbees whichfollow fixeddailyforagingroutes(Janzen,1971).In contrast
"big-bang"speciesput out all of their flowersin a few days with all the indi-
vidualsof the speciesbloomingat the sametime. Thesespeciesall bloom in the
dry seasonand attracta variedassemblageof opportunisticallybehavingpolli-
nators, apparentlyincludingbee specieswhich normallyfollow a fixed trap-
line. "Multiple-bang" bignonspecies,like "big-bang"species,put out all their
flowersat once for a very short periodwith floweringof the individualsof a
specieshighly synchronized.They differ in having floweringburstsscattered
sporadicallythroughthe year, and most of them are apparentlynon nectar-
producingmimicswhichdependfor pollinationon the occasionalvisitsof bees
opportunisticallysearchingfor new nectar sources. Most Bignoniaceaespe-
cies, however-including all the speciespollinatedby hawkmoths,humming-
38 FloraNeotropica

birds,and butterfliesand smallbees-bloom for a periodof severalweeksto a

few months buildingup to a distinctpeak of flower productionfollowed by
gradualdecreasesin numberof flowers. These "cornucopia"species bloom
much less synchronouslythan "big-bang"and "multiple-bang"species but
have a definite floweringseason during which all individualsof the species
bloom. Bee-pollinated"cornucopia"speciesdrawthe same diversespectraof
pollinatorsas "big-bang"species.Different"cornucopia"speciesof Bignoni-
aceae flower duringdifferentseasons of the year (Gentry, 1974a,b) and sea-
sonal separationis an importantisolatingmechanismfor them.

constancyand nichespecificity

Althoughdifferentspeciesof Bignoniaceaeoccur in dry, moist, and wet

forest habitats, about the same numberof speciesseem to occur in any low-
land tropicalcore habitat plant community.This is in markedopposition to
the generaltrend shown by most families of increasingspecies diversitywith
increasinglymesic conditions. As a result, Bignoniaceaeare much more con-
spicuous elementsof dry forest than of wet forest plant communities,even
thoughsuchcommunitieshaveno morespecies(about20) thanwet forestones
The ca. 20 strictlysympatricspeciesof Bignoniaceaewhichoccurtogether
in any lowland neotropicalplant communityare highly differentiatedas to
pollinationstrategy.In fact thereis such a strongnegativecorrelationbetween
the pollinationstrategiesof sympatricspeciesthat, in general,in a given com-
munity each species of Bignoniaceaehas a unique and exclusivepollination
niche determinedby pollen vector, phenologicalstrategy(in the case of bee-
pollinated species), and seasonality (in the case of "cornucopia" species).
Four or five of the 20 Bignoniaceaespecies of a given plant communityare
pollinated respectivelyby bats, hummingbirds,butterfliesand small bees,
and hawkmoths.About five of the remaining15 large bee-pollinatedspecies
utilize "steady-state," "big-bang" and "multiple-bang"phenologies, while
about ten are "cornucopia"flowerers.Each of these about ten strictlysym-
patric"cornucopia"speciesof Bignoniaceaehas its floweringpeakin a differ-
ent month. In general,in any givenplantcommunity,only a singlebignonspe-
cies usinga givenphenologicalstrategywill be in flowerat any time.


I have summarizedvarious aspects of Bignoniaceaedispersal ecology

from several perspectives(Gentry, 1973a, 1974a, 1976b, 1977a, b, 1978b,
1979a). Most species of the family have dehiscentfruits and winged wind-
dispersedseeds. There are two basic modificationsof this pattern. Water-
dispersedseedshave evolvedfrom wind-dispersedones independentlyin many
taxa of Bignoniaceae.These water-dispersedspecies have thickercorky seed
bodies with the wings reducedor lacking. The differenceshave been over-
Ecology 39

emphasizedtaxonomicallywith manygenerasegregatedon accountof the dif-

ferencebetweenwind-dispersedand water-dispersedseeds (Gentry,1973a).In
some cases a single species may even have both wind-dispersedand water-
The second major trend in Bignoniaceaeseed dispersalis a switch from
dehiscentto indehiscentfruits. Indehiscentfruitshave evolved from dehiscent
fruitstwice,givingriseto the Crescentieae in the New Worldand the Coleeaein
Afromadagascar(Gentry, 1976b). IndehiscentBignoniaceaefruits are fleshy
and mammal-dispersedand the evolutionarysequenceleading to neotenous
fruit maturationto make use of the mammalianpredatorsof immaturede-
hiscentfruitsas dispersalagentsis clear (Gentry,1974a, 1976b).Severalinde-
hiscent-fruitedspecies still retain vestigialseed wings, indicatingtheir evolu-
Schlegelieaealso have indehiscentfruits although their relationshipto
other Bignoniaceaeis unclear.The fruits of most Schlegelieaeare small bird-
Bignoniaceaedistributionpatternsareintimatelycorrelatedwith dispersal
ecology. Most wind-dispersedand water-dispersed species are wide-ranging.
In contrastthe mammal-dispersed indehiscent-fruitedspeciesmostlyhavevery
restrictedrangesand high local endemism(Gentry,1974a, 1979a).Apparent-
ly this distributionaldichotomyis relatedto more localizeddispersalby mam-
mals. The few wide-rangingspeciesof Crescentieaehavehardthick, protective
shells outside the pulp in which the seeds are embeddedand the whole fruit
secondarilywater-dispersed.Bird-dispersedSchlegelieaehave an intermediate
distributionalpatternwith some speciesratherwide-rangingand some strictly
Water-dispersedspecies tend to be almost as wide-rangingas wind-
dispersedspecies but are mostly edaphic specialists confined to mangrove
swampsor their fringes(Phryganocydiaphellosperma,Tabebuiapalustris, T.
aquatilis, T. obtusifolia, T. cassinoides), seashores Amphitecna latifolia
(except in the West Indies)), freshwaterswamps, streamsides(e.g. Anemo-
paegma chrysoleucum(HBK.) Sandw., Phryganocydiauliginosa Dugand,
Tanaeciumjaroba Sw., T. crucigerumSeem., Macfadyena uncata (Andr.)
Sprague& Sandw., Clytostomabinatum(Thunb.)Sandw.), or the seasonally
inundatedvarzeaand tahuampaforests along the Amazon and Orinoco and
their major tributaries(e.g. Anemopaegmaparaense Bur. & K. Schum.,
Memoraschomburgkii(DC.) Miers, Arrabidaeabilabiata(Sprague)Sandw.,
A. bracteolata(DC.) Sandw.,A. revillaeA. Gentry).
Bignoniaceaedispersalecology seemsto be correlated,in a somewhatun-
expectedway, with the emergingphytogeographicpatternsrelatedto and de-
rived from Pleistoceneforest refugia. The prevalentpatternof wide-ranging
ecologicallyrestrictedcore habitat Bignoniaceaespecies may reflect cyclical
range extensionsof easily dispersedspecies, dry forest ones alternatingwith
mesicones as theirrespectivehabitatsexpandedand contracted,with capacity
to partitionlocal pollinatorresourcesthe criticalfactor throughevolutionary
time in limitingintracommunitydiversityand determiningwhich new arrivals
would successfullyenterexistingcommunities.The intricatediversebiotically
40 FloraNeotropica

regulatedBignoniaceaecommunityends abruptlynear the Tropicsof Cancer

and Capricornand the few temperatezone speciesseemby comparisonalmost
randomcomponentsof their communities.In the rich equilibratedcore habi-
tat tropical communities,evolutionarychange may largely involve replace-
ment, throughouta region and habitat, of speciesoccupyingparticularpolli-
nation niches by easily dispersedsuperiorcompetitors.Successfulspeciation
leadingto overallincreasesin numberof speciesof neotropicalBignoniaceae
probablymost frequentlyinvolvesenteringfloristicallyimpoverishedmarginal
habitats. Active diversificationof populationsisolated in refugia may have
lead largelyto evolutionarydeadendsunlessthe end productsof this evolution
wereable to enterand specializein peripheralhabitats.Thus it is no doubt sig-
nificant that most of the bignon specieswhich are restrictedto specific refu-
gium-correlatedphytogeographicregionsseem to be edaphicspecialists.Effi-
cient dispersalseemsto have guaranteedthat any superiorcompetitorfor one
of the availablecore habitat pollinationniches which might have arisenin a
refugialregionwould have spreadwidelyin suitablehabitatsas refugialforest
patchescoalescedduringperiodsof decreasingaridity(Gentry,1979a).


Although Bignoniaceaeare pantropicalin distributionand even include a

few temperatezone representatives,they are predominantlyneotropicalwith
620 or 78%oof a worldtotal 800 speciesnativeto the New Worldtropics.In the
processof mongraphingthe family for Flora Neotropica,I have been able to
studyBignoniaceaeextensivelyin the field overmost of LatinAmerica;as a re-
sult, its distributionis perhapsbetterknown in severalcountriesthan that of
any other plant family. An overviewof distributionalpatternsof neotropical
Bignoniaceaeconcentratingon some of their phytogeographicimplications
has been presentedelsewhere(Gentry, 1979a).A more detailed summaryof
distributionsis givenbelow.
Dividing the family into tribes gives a clearerpicture of its distributional
patterns(Fig. 5). For the familyas a whole Brazilis clearlythe centerof diver-
sity. However, each tribe has a different, and unique, center of diversityif
numberof speciesis takenas a diversitycriterion.


Bignonieaeis by far the largesttribewith 349 speciesdistributedamong50

genera, 56%oof the Americanspecies. All generaare exclusivelyor predomi-
nantlytendrillatevines, fruitdehiscenceis generallyparallelto the septumand
the seeds are wind (or water)dispersed.Most speciesare wide-ranging.Brazil
has by far the most species(269, 74%oof the tribe'stotal) and genera(45). All
generaexcept five monotypicones-Saritaea in Colombia, Macranthisiphon
in coastal Ecuador and adjacent Peru, Bignonia in the southeasternUnited
States, Pseudocatalpain northern Central America, and Potomoganos in
Guayana-are representedwithinthe limitsof Brazil;37 generahave as many
or more Brazilianthan extra-Brazilianspecies. Diversitydeclinesrapidlyand
Distribution and Phytogeography 41

c I
n/' o

a\~ 4 t -

Wi'---'^ AT /-

/ ?
SM-w/y C A^


countries, West
FIG. 5. Distribution of neotropical Bignoniaceae by country. Central American countries,
(coastal islands excluded), and Guianas treated as single
Indies (coastal genera
single units. Total species/total genera
species per tribe for Bignonieae
plus number of species
for each country plus (T), and Crescen-
Bignonieae (B), Tecomeae (T),
(C). Species
tieae (C). Species breakdowns for Schlegelieae with 23 species Panama,
species centered in Colombia and Panama,
Eccremocarpeae with 6 species mostly in Peru, and Tourrettieae are not listed separately. Many
species of West Indian Tecomeae have not yet been critically evaluated. (From Gentry, 1979a).

rather uniformly
rather away from
uniformly away (Fig. 5).
Brazil (Fig.
from Brazil 5). Only nine species
Only nine species are endemic to
are endemic to
America and
Central America
Central to the
three to
Mexico, three
and Mexico, West Indies,
the West monotypic
and monotypic
Indies, and
Bignonia to to the
the southeastern United States.
southeastern United Old World
The Old
States. The often
trees often
World trees
to Bignonieae
assigned to
assigned are probably
Bignonieae are probably unrelated 1976b, and
(Gentry, 1976b,
unrelated (Gentry, below).
see below).
and see
very natural
this very
Clearly this
Clearly natural tribe, thus the
and thus
tribe, and the major part of
major part of the
the family, evolved
family, evolved
probably in
America, probably
South America,
in South in Brazil,
Brazil, subsequent to the
subsequent to breakup of
the breakup Gond-
of Gond-


the second
Tecomeae, the
Tecomeae, major tribe
second major tribe of includes 34%
Bignoniaceae, includes
of Bignoniaceae, of
(215) of
34o (215)
New World
the New
the species distributed
World species among 20
distributed among genera. Most
20 genera. genera are
Most genera or
trees or
are trees
few-north temperate
shrubs, a few-north
shrubs, south temperate
Campsis, south
temperate Campsis, temperate Campsidium, and
Campsidium, and
42 FloraNeotropica

the problematicaland isolated BrazilianPerianthomega-are vines without

terminalleaflets modifiedinto tendrils.Argyliaof Chile and adjacentArgen-
tina and Peru is herbaceous.Fruitdehiscencein Tecomeaeis perpendicularto
the septum and the seeds are wind (or water) dispersed.The distributional
patternof Tecomeae is not as clear cut as for Bignonieae. Tabebuiais the
largestgenus and dominatesthe statistics. If the 58 Tabebuiaspecies recog-
nizedby the Floraof Cubaare accepted,Cubahas moreTecomeaespecies(71)
than any other countryand mightbe consideredthe tribe'scenterof distribu-
tion. Howeverall of the West Indianspeciesof Tabebuia(excludingthose of
the coastal islandsoff South America)as well as those of its segregategenera
Ekmaniantheand Spirotecomaare very closely relatedto each other and to
mainlandT. rosea (Bertol.)DC. Despiteits many species,this groupincludes
only a small fractionof the specific variabilityof South AmericanTabebuia
whichis actuallycenteredin Brazil.Moreovergenericdiversityin Tecomeaeis
greatestin Brazil (9 genera)and Peru (8 genera)and Jacaranda,the second
largestgenus of the tribe, has 30 of its 46 speciesin Brazil. Brazilseems the
apparentdistributionalcenterfor Tecomeaeas for Bignonieae;the explosive
adaptive radiation of leaf size and texture which produced so many West
Indian species from one otherwisesmall section of Tabebuiais a secondary
Tecomeae, perhaps the oldest extant group of Bignoniaceae(Gentry,
1976b)almostcertainlypredatethe breakupof Gondwanaland,and the Ameri-
can genera appearto derive from severalsources. While Jacarandaand the
basically palmately compound-leavedAmerican genera centered around
Tabebuiaare autochthonouslyevolved and not closely relatedto Old World
Tecomeae,thereis a distinctivetemperateNorth Americanelementconsisting
of Chilopsis, Catalpa,and Campsiswhich is more closely relatedto the Old
World Tecomeae than to the South Americanones. Catalpa and Campsis,
each disjunctbetweenthe southeasternUnited States and southeastAsia (the
former also in the West Indies), were historicallypart of the well-known
Arctotertiaryflora. Chilopsisis closerto Catalpathan to any othergenus and
derivedfrom the sameancestralstock. It seemsprobablethat tribeTecomeae,
perhapsthe moderngroup nearestto the ancestralstock of Tubiflorae,stem
from a West Gondwanalandorigin (cf. Raven & Axelrod, 1974) with the
North American representativesof Tecomeae arriving later via Eurasia
(Catalpais presentin the EoceneLondonclay flora (Paclt, 1952)and has scat-
tered fossils elsewherein Europe and westernNorth America) and derived
from palaeotropicalancestors.The pinnatelycompound-leavedSouth Ameri-
can genera are also problematicalphytogeographically.South temperate
Campsidiumis unrelatedto other South Americanspecies but is similarto
Campsisand to Tecomantheof Australia.Perhapsit is a remnantof the Old
West Gondwanalandbignoniaceousstock; this would be supportedby the
strikingresemblanceof Campsidiumto monotypicPerichlaenaof Madagascar
which may prove more than superficialafter all. SimilarlyTecoma,with ten
species of the Central Andes plus a single wide-rangingspecies distributed
from southwesternUnited Statesand the West Indiesto Argentina(but absent
from Braziland Guayana),is more closely relatedto its South Africansegre-
gate Tecomariathan to any New World genus. Their relationshipis so close
Distributionand Phytogeography 43

that it has been argued(Seemann,1863)that Tecomariamust reallybe native

to SouthAmerica(whereit is widelycultivatedbut rarely,if ever, escaped)and
introducedin Africa. New evidencefrom cytology(Goldblatt& Gentry, 1979)
and experimental greenhouse crosses supports reuniting Tecomaria and
Tecoma,a suggestionalreadyfollowedby van Steenis(1978).Relationshipsof
threeother New Worldgeneraare unclearand may date from ancientstocks.
Monotypic Astianthus of southern Mexico and northern Central America
could be derivedfrom Tecomaor mightrepresenta remnantof an old tropical
North Americanstock derivedfrom the same ancestorsas Chilopsis.Argylia,
centeredin centralChilewith a singlespeciesreachingthe southernmostPeru-
vian lomas, is the only herbaceousgenusof Tecomeaein the neotropics.Its re-
lationshipto similarlyherbaceousand superficiallyvery similarIncarvilleaof
the Himalayashas been suggested.However,Argyliahas the palmatelycom-
pound leaves of most neotropicalTecomeaeand Incarvilleathe pinnateones
typical of Old Worldgenerawhich may indicateconvergence,instead. Delo-
stoma (four speciesof the PeruvianAndes, one of them rangingnorth to the
subparamosof Colombia and extreme western Venezuela, has no obvious
affinitieswitheitherOld or New Worldgenera.


In the narrowsenseusedhere,Crescentieaeincludesonly threegeneraand

33 species. All are trees or shrubs with large indehiscent fruits, mostly
mammal-dispersed seeds, and bat-pollinatedflowers. In contrastto Tecomeae
and Bignonieae,Crescentieaeare prone to local endemismand most species
have highly restricteddistributions,often on single mountains.The distribu-
tionally exceptionalwidespreadspeciesare those few which have secondarily
revertedto water dispersal-of the entire fruit (Gentry, 1974a). Crescentieae
diversityis centeredin CentralAmericaand southernMexico where29 (88%)
of its speciesoccur. Crescentia,the most highlyderivedgenus (Gentry,1974a),
is also the most widespreadwith three of its six speciesendemicto the West
Indies. Except for the cultivatedcalabash(Crescentiacujete) and the rather
widespreadcoastalspeciesAmphitecnalatifolia,the tribebarelyreachesSouth
America-two species, one endemic,are in extremenorthwestColombianear
the Panamaborderand Crescentiaamazonica,possiblya small-fruitedrecent
derivativeof introducedC. cujete,is along the upperand CentralAmazonand
Orinoco. The indehiscent-fruited Madagascar Bignoniaceae, separately
evolved from a differentTecomeaestock, representconvergenceratherthan
commonoriginand shouldbe treatedtaxonomicallyas tribe Coleeae(Gentry,


Schlegelieae,consisting of three genera-Schlegelia, Gibsoniothamnus,

and Synapsis-are simple-leavedlianas and shrubs, usually hemiepiphytic,
with small to medium-sized, usually berry-likebird-dispersedindehiscent
44 FloraNeotropica

fruits, and bee- or hummingbird-pollinated flowers; traditionallythey have

been treated under Crescentieae.Some recent studies have suggested that
Schlegeliaand its relativesmay be closerto Scrophulariaceae than to Bignoni-
aceae although their chromosomenumberand 3-partedseedlingleaves sup-
port retentionin Bignoniaceae.At any rate they are not closely related to
Crescentieaeand merit at least tribal recognitionin either family. The taxo-
nomic problemsin familialplacementof this group suggestthat it may be an
old one, representinga survivingremnantof the ancestralstock common to
both Scrophulariaceaeand Bignoniaceae.Schlegelieae-exclusivelyneotropi-
cal-are completelyunrelatedto southeastAsian Paulownia, althoughPaul-
owniais similarlyintermediatebetweenBignoniaceaeand Scrophulariaceae.
Species diversityof Schlegelieaeis centeredin Colombia and southern
CentralAmerica.Eight of the 14 speciesof Schlegeliaoccur in Colombiaand
five of the eight speciesI recognizein Gibsoniothamnus (threeout of threeby
D'Arcy's (1979)overbroadconcepts)occurin Panama,none in South Ameri-
ca or the West Indies, and only a single speciesnorth to Mexico. Monotypic
Synapsisis endemicto Cuba'sOrienteProvince.

Tourrettieaeand Eccremocarpeae

The final two tribesof AmericanBignoniaceaeare both small and mono-

generic. Tourrettieaewith the single species Tourrettialappaceais montane,
occurringthe length of the Andes in South Americaand north in the Central
Americancordillerasto Mexico. It is an annualherbaceousvine with tendril-
late leaves, a 4-loculedovary and spiny bur-likeincompletelydehiscentexo-
zoochoricfruit. Eccremocarpus,the only genus of Eccremocarpeae,is central
Andean with five species in Peru, one of these reachingnorth to southern
Colombia, and a sixth in Chile and Argentina.Eccremocarpusis a wiry vine
with multifoliolateusuallytripinnatisecttendrillateleaves, tubularhumming-
bird-pollinatedflowers, a 1-celledovarywith parietalplacentationand a thin-
walledovoid capsulewithouta septum. Tourrettiaand Eccremocarpusare re-
lated to each other but otherwiseisolated in the family; the formersuggests
Pedaliaceaeand could providea link withthat Old Worldgroup.


Accordingto van Steenis(1978)"thereare no outstandingqualitiesmark-

ing (Australasian) Bignoniaceae as useful plants. . . ". The situation is decid-
edly differentfor New WorldBignoniaceae.Probablythe majoreconomicim-
portanceof the familyis in horticultureand some of its speciesare amongthe
most widelyplantedin the tropicalworld.JacarandamimosifoliaD. Don may
well be the world'ssinglemost widelyplantedornamentaltropicaltree and its
profusion of purplish-blueflowers are an even better known feature of
southernAfricanand Australiancities than of theirNew Worldcounterparts.
Uses 45

In Pretoria, South Africa, for example, nearly every street tree is of this spe-
cies (Goldblatt, pers. comm.). Spathodea campanulata Beauv., native to tropi-
cal Africa, is another of the most widely planted ornamental tropical trees,
and is especially frequent in neotropical cities. Campinas, Brazil, represents an
extreme case: Spathodea is so esteemed that a municipal policy has been imple-
mented to encourage its propagation-homeowners are provided free saplings
of Spathodea but not of any other tree species (L. El-Dash, pers. comm.).
Podranea ricasoliana (Tanfani) Sprague, native to Africa, and Pyrostegia
venusta (Ker.) Miers, native to South America, are among the most attractive
and most widely cultivated tropical ornamental vines and Campsis radicans
(L.) Seem. (trumpetvine) claims a similar distinction in temperate North
America. Other very widely cultivated ornamental species of Bignoniaceae in-
clude Saritaea magnifica (Sprague ex van Steenis) Dugand, Tecomaria capen-
sis (Thunb.) Spach, Pandorea jasminoides (Lindl.) K. Schum., and several
species of Tabebuia. Menninger (1970) lists 34 species of Bignoniaceae vines
which he considers of horticultural importance.
The esthetic qualities of showy-flowered Bignoniaceae are also appreci-
ated in their native countries. Many Latin American countries have selected
species of Bignoniaceae as their national tree or national flower. These include
El Salvador (Tabebuia rosea (Bertol.) DC.), Venezuela (Tabebuia billbergii
(Bur. & K. Schum.) Standl.), Brazil (Tabebuia serratifolia (Vahl) Nichols.),
Ecuador (Tabebuia chrysantha (Jacq.) Nichols.) and Paraguay (Tabebuia sp.);
Jacaranda mimosifolia was the runnerup for this designation in Argentina
(Fabris, 1965).
Bignoniaceae are also among the most important neotropical timber trees.
The species of the most widespread section of Tabebuia have perhaps the
hardest, heaviest, most durable wood of any neotropical tree, a distinction
shared with lignum vitae (Guaiacum). The wood of Bignoniaceae is much
prized for construction because of its strengths and durability. Beams of
Tabebuia wood, known to have been exposed to the elements for 400 years in
Panama, are still free of decay (Record & Hess, 1940). The few sound trees
left standing in Gatun Lake from the forest inundated over 50 years ago by
construction of the Panama Canal are mostly Tabebuia guayacan (Seem.)
Hemsl. (Southwell & Bultman, 1971). Resistance of T. guayacan wood to
marine wood-boring organisms is among the highest for any Panamanian spe-
cies, and it is the most termite resistant of the 112 species tested by Bultman &
Southwell (1976). The wood of many Tabebuia species is further esteemed for
the esthetic appeal of the contrasting very dark heartwood and light sapwood.
For example, Tabebuia chrysantha (Jacq.) Nichols., called "guayacan," is the
most important timber tree of western Ecuador with much of its lumber being
converted into knickknacks like salad and sugar bowls and carved statues as
well as furniture. One indication of the esteem in which "guayacan," now be-
coming progressively rarer in Ecuador (Gentry, 1977a), is held is the preva-
lence on the tourist market of items of imitation guayacan made by painting
dark brown bands or patches on an article made from a light-colored wood.
The very different medium weight wood of Paratecoma peroba (Record)
Kuhlm. and of Tabebuia rosea, T. heterophylla (DC.) Britton, and their rela-
tives is also an important commercial timber. Tabebuia rosea, called "roble"
46 FloraNeotropica

(Spanishfor oak), is one of the major timberspeciesin CentralAmerica;in

Panamathis is the fourth speciesto be selectivelycut by timbercruisers(after
mahogany(Swietenia),spanish cedar (Cedrella),and "cedro espino" (Bom-
bacopsis), before general lumberingof an area begins. Paratecomaperoba
("peroba" or "ipe peroba"), now threatenedwith extinction,was one of the
most importantwoods of coastal Braziland used to finish nearly all of the
betterclassof housesand commercialbuildingsof Rio de Janeiroas well as for
muchof the best qualityfurnitureof that region(Record& Mell, 1924).Many
additionaluses for the wood of variousBignoniaceaeare listed by Record&
Hess (1940),Standley(1926),and otherauthors.
Bignoniaceaeare also ethnobotanicallyimportantand constitutean im-
portantelementof many native pharmacopeias.For example,the onion and
garlic-smellingspecies of Pachypteraare widely used in tropical folk medi-
cines. "Clavohuasca,"one of the best known ingredientsof folk medicines
and tonics in AmazonianPeru, comes from clove-smellingspeciesof Tynan-
thus. Martinellaobovata (HBK.) Bur. & K. Schum.is used and cultivatedby
diversegroupsof SouthAmericanIndiansin severalcountriesas an eye medi-
cine. Cyanide-containingTanaeciumnocturnum (Barb. Rodr.) Bur. & K.
Schum. is used as a love potient (Gentry, 1973c)and hallucinogen(Prance,
1977)by variousprimitivepeoples. The barkof some speciesof Tabebuia,es-
pecially T. impetiginosa(Mart. ex DC.) Standl. and T. serratifolia (Vahl)
Nichols. is widelyremarkedas a cancercure, so muchso that in muchof Ama-
zonia these species have been seriously decimatednear population centers
(Tyson, pers. comm.; Plowman, pers. comm.). Interestinglythis folk medi-
cinal anti-canceruse proves to have a strong factual basis; Bignoniaceaehas
provedto be among the most promisingfamiliesin anti-canceractivityin the
currentUSDA screeningprogram, along with Apocynaceae, Leguminosae,
Euphorbiaceae,and Rutaceae(Hudson,pers.comm.; Oguraet al., 1977).
Bignoniaceaespeciesare also the sourceof severaldyes. Arrabidaeachica
(H. & B.) Verl. producesa red dye which was formerlymuch used as a cos-
meticto color the skin;Humboldt(Humboldt& Bonpland,1808)reportedthis
use so prevalentalong the Orinocothat the local equivalentof "He is so poor
that he can't clothe half his body" became"He is so poor he can't paint half
his body." Cybistaxantisyphilitica(Mart.)Mart.ex DC. providesa blue dye,
formerlymuchusedin AmazonianPeru(Spruce,1859).Othermembersof the
Bignoniaceaeare notoriouslytoxic to livestock. TanaeciumexcitiosumDu-
gand of the MagdalenaValley of Colombia, universallyknown to ranchers
and campesinosas "mataganado," has beenpracticallyeradicatedin orderto
protect cattle from it (pers. obs. and Dugand, 1942). Arrabidaeabilabiata
(Sprague)Sandw. enjoys similarnotorietyin the Orinocovalley of Venezuela
as does Arrabidaeaelegans (Vell.) A. Gentry of coastal Brazil (Kuhlman,
1941;Tokarniaet al., 1969). On the other hand, many of the vines of tribe
Bignonieaeare important"bejucos de agua" from the cut sectionsof which
campesinosand woodsmenget potablewater.
Several membersof tribe Crescentieaehave useful fruits. These range
fromthe ediblefruit, similarin textureto a cucumber,of Parmentieraaculeata
(HBK.)Seem. to the hard-shelledcalabashproducedby speciesof Crescentia.
Systematic Treatment 47

The hollowed shell of the calabash or tree gourd is used throughout the neo-
tropics as a household utensil and Crescentia cujete is one of the most widely
cultivated Latin American species. The calabash was widely cultivated long be-
fore the Spanish conquest. In fact one of Columbus' first journal entries on
the day he discovered the New World, October 13, 1492, is a commentary on
the use of calabashes to bail the natives' dugout canoes (Sauer, 1969). Another
interesting use of calabashes by the West Indian natives noted by Columbus
and his contemporaries was in hunting waterfowl. Indian swimmers hidden
under a floating calabash with eyeholes cut in it would approach a flock of
birds and pull individual birds under water without disturbing the flock. The
earliest accounts of Panama record a similar practice (Sauer, 1969). The
smaller fruit of C. alata is widely used for "maraca" rattles and as ladles and
dippers. The pulp of Crescentia fruits, though insipid, is occasionally eaten as
is the waxy fruit of Parmentiera cereifera (Wafer, 1699). The latter has been an
important cattle food (Seemann, 1851) and was dried and used for candles by
pre-conquest Panamanian Indians (Wafter, 1699). The seeds of Crescentiaalata
are used to make a beverage called "semilla de jicaro" in Nicaragua; although
much esteemed and popular locally, this use does not appear to have caught on
in other countries.


BIGNONIACEAE A. Jussieu, Gen. 137. 1789 ("Bignoniae"); de Candolle,

Prodr. 9: 142-248. 1845; Bureau, Monographie des Bignoniacees 1-215.
1864; Bentham and Hooker, Gen. P1. 2(2): 1026-1053. 1876; Baillon,
Hist. des Plantes 10: 1-58. 1891; K. Schumann in Engl. and Prantl., Nat.
Pflanzenf. 4(3b): 189-252. 1894; Bureau and K. Schumann in Martius,
Flora Bras. 8(2): 1-452. 1896-97; van Steenis, Flora Malesiana, ser. I,
8(2): 114-186. 1977.

Woody trees, shrubs, or vines, rarely herbaceous; outer scales of axillary

buds often pseudostipular and sometimes foliaceous; glandular fields often
present between the petioles and rarely at the petiole apex. Leaves usually op-
posite, usually palmately or pinnately compound, sometimes simple, the ter-
minal leaflet often replaced by a tendril. Inflorescence terminal or axillary or
cauliflorous, paniculate or racemose or reduced to a fascicle or a single flower.
Flowers with the calyx synsepalous, usually more or less cupular, truncate to
variously lobed or spathaceously split; corolla sympetalous with a conspicuous
tube and 5 usually imbricate lobes; stamens inserted on corolla tube, didyna-
mous, usually 4, rarely 2 or 5, a posterior staminode usually present, anthers
with 1 (rarely) or 2 thecae, these usually divergent or divaricate and held
against the adaxial side of corolla tube, the filaments slender, the pollen 3-
colpate with a finely reticulate exine to inaperturate, multicolpate, or areolate,
with psilate to coarse-reticulate or spinulose exine, the grains simple, in
tetrads, or rarely in polyads; ovary superior, 1 or (usually) 2 locular, the ovules
48 Flora Neotropica

numerous, attached to 4 vertical placental ridges, these axile in 2-locular

ovaries and parietal in unilocular ovaries (a single placental ridge in each locule
in many species of Schlegelieae), style slender, stigma bilamellate; a nectarifer-
ous disc usually conspicuous at base of ovary, sometimes lacking. Fruit a cap-
sule with dehiscence perpendicular or parallel to the septum or a berry or hard-
shelled pepo (calabash); seeds without endosperm, somewhat flattened,
usually winged, the cotyledons foliaceous.
Type genus. Bignonia L.
A family of 113 genera and 800 species, mostly tropical with a few genera
in warm temperate regions. The family is divided into eight tribes, two of
which are monogeneric. The Tecomeae are found in both the Old and New
Worlds. The Bignonieae sensu stricto, Crescentieae sensu stricto, Tourrettieae,
Eccremocarpeae,and Schlegelieae2are restrictedto the New World. Oroxyleae3
are in Southeast Asia; Coleeae are on Madagascar with a single monotypic
genus on continental Africa.

Key to the tribes of Bignoniaceae

1. Fruitdehiscent;trees,shrubs,herbs,or compound-leaved lianas.
2. Placentationparietal;fruit withouta septum;wiryvines;above 3000 m altitude(unless
in Chileor Argentina)in the Andes. Eccremocarpeae.
2. Placentationaxile; fruit with a septum;habit various,if scandentthen succulent-herba-
ceous or woody; below 2800 m altitude(excepta few speciesof herbaceousArgyliain
ChileandArgentinaandIncarvilleain the Himalayas).
3. Herbaceousrathersucculentvines;inflorescencesubspicate,the upperflowersmostly
sterile;ovary4-locular;capsuledenselyuncinate-spiny,bur-like,the 4 valvesnot split-
tingto the base. Tourrettieae.
3. Woodyvines, trees,shrubs,or erectherbs;inflorescenceneverspicate,withoutsterile
flowers;ovarybilocular;capsulespinyor not, if spiny,not bur-likeand the two valves
dehiscentto base.
4. Lianaswith the terminalleafletof a compoundleaf frequentlymodifiedinto a ten-
dril (or derivedsuffrutescentor treeletforms withouttendrils);stemswith anoma-

2Schlegelieae tribus nov.-A tribu Crescentieae foliis simplicibus oppositisque, fructibus baccatis
parvioribus, seminibus minutis angularibus, floribus tubularibus lobis parvis reflexis, disco nec-
tarifero plus minusve carenti, habito plerumque epiphytico differt.
This tribe includes two genera formerly placed in Crescentieae and one recently described in
Scrophulariaceae. They are characterized by opposite simple leaves, relatively small baccate, inde-
hiscent fruits with tiny wingless angular seeds, reduction of the style and stigmas, and usually
epiphytic or hemiepiphytic habit and parietal placentation. Type genus Schlegelia. Also included:
Gibsoniothamnus and Synapsis. The familial placement of this clearly natural aggregate is unclear
but it is not closely related to Crescentieae and would deserve tribal recognition just as much in
Scrophulariaceae as in Bignoniaceae.

3Oroxyleaetribus nov.-A tribu Tecomeae fructibus parallelibus ad septis dehiscentibus recedit. A

tribu Bignonieae foliis non cirrhis, ligno sine anatomia anomala, chromosomatum numero 14 vel
15, staminum numero plerumque 5 differt.
This tribe includes those Old World genera with fruit dehiscence parallel to the septum which
were formerly placed in Bignonieae. Type genus Oroxylum. Also included: Millingtonia, Nycto-
calos, and Hieris (fide van Steenis, 1978). It is not altogether certain that these genera are suffi-
ciently distinct from Tecomeae for tribal recognition, despite the mode of fruit dehiscence, but
they are clearly not closely related to Bignonieae.
SystematicTreatment 49

lous phloem arms (exceptin a few suffrutescentcerradospecies);fruits dehiscent

parallelto the septum(or occasionally4-valved)exceptDolichandracynanchoides;
New World. Bignonieae.
4. Trees, shrubs,herbs, or non-tendrillatelianas (Perianthomegahas a twiningpeti-
ole); stems withoutanomalousphloemarms;fruitsdehiscentperpendicularto the
septumexceptin a few OldWorldspecies;OldandNew Worlds.
5. Fruitsdehiscentparallelto the septumor 4-valved;tropicalAsia. Oroxyleae.
5. Fruitsdehiscentperpendicular to the septum;widespread. Tecomeae.
1. Fruitindehiscent;trees,shrubs,or simple-leavedhemiepiphyticlianas.
6. Leavespinnatelycompound(or reducedto superimposedarticlesor rarelysimple);Mada-
gascar and Africa; terrestrial;flowers mostly bee-pollinated(flagelliflorousand bat-
pollinatedin Kigelia). Coleeae.
6. Leavessimpleto palmatelycompound4;neotropics;terrestrialor epiphytic;flowerscauli-
florousto terminal,bat-pollinatedif trees.
7. Flowersinsect-or hummingbird-pollinated; usuallyepiphyticshrubsor hemiepiphytic
lianas;leavessimpleandopposite;fruitsless than4 cm long; seedswithnon-foliaceous
cotyledons. Schlegelieae.
7. Flowersbat-pollinated;terrestrialtreesor shrubs;leavesalternateor palmatelycom-
pound(orboth);fruitsmorethan(3-)4cm long;seedswithfoliaceouscotyledons.

I. BignoniaceaetribeCrescentieaede Candolle,Prodr.9: 240. 1845.

Trees or shrubs (possibly scandent in Amphitecnaparviflora). Leaves

oppositeor alternate,simpleor palmately3-7-foliolate(one collectionof Par-
mentieraaculeatawith some leavespinnately5-foliolatehas been seen). Inflor-
escenceone to a few flowers, terminal,axillary,or in fasciclesfrom ramiflor-
ous or cauliflorousbracteateshort shoots. Flowers bat-pollinated,with the
calyx irregularlyrupturingapicallyto spathaceous,bilabiate,or irregularly3-
labiate;corollatubular-campanulate or tubular-infundibuliform, usuallywith
a bulgeor transversefold in throat,the lobes frequentlyreducedor essentially
absent;glabrousor with scatteredstalked-lepidoteglands;stamens4, didyna-
mous, usually subexserted,a posteriorstaminodeusually present,the anther
thecae thick, divergent, the pollen grains simple, 3-colpate to multiporate,
finely reticulate;ovary unilocularwith 2 or 4 parietalplacentae,or bilocular
with 2 axile verticalplacentalridgesin each locule, often bilocularat base and
unilocularat apex, the stigmabilamellate;nectariferousdisc at base of ovary
large,conspicuous.Fruitindehiscent,baccatewith a fleshy cortexand fibrous
core or a hard-shelledpepo or calabashwith the seeds embeddedin the soft
pulp;seedsflattened,winglessor witha vestigialmucilaginouswing.
Typegenus. CrescentiaL.

Keyto the Generaof Crescentieae

1. Leavesopposite,3-7-foliolate,the petiolewinglessor narrowlywinged;calyx spathaceously
split;fruitwithouthardshell. 3. Parmentiera.

4Onecollectionof normally3-foliolateParmentieraaculeatawitha pinnately5-foliolateleaf (and

one witha palmately5-foliolateleaf) hasbeenseen.
50 FloraNeotropica

1. Leavesalternateor fasciculate,usuallysimple(if compound,the wingedpetiole almost as

wide as the leaflets);calyx various, usuallybilabiate,rarelyspathaceous;fruit with hard
2. Leavesalternate,not fasciculate;placentationaxile at least at extremebase of ovary,
aboveparietalon two placentae;seedslarge(greaterthan8 x 10mm). 1. Amphitecna.
2. Leaves fasciculatein alternateclusters;placentationparietalwith 4 placentae;seeds
small(lessthan8 x 9 mm). 2. Crescentia.


1. AmphitecnaMiers,Trans.Linn. Soc. ser. 1, 26:163. 1870.

DendrosicusRafinesqueSylvaTell. 80. 1838.Type: D. saxatilisRafinesque= A. latifolia
(Miller) A. Gentry, nom. rej.
Crescentiasect. EnallagmaMiers,Trans.Linn.Soc. London,ser. 1, 26:174. 1870.
Enallagma (Miers) Baillon, Bull. Mens. Soc. Linn. Paris 85: 679. 1887. Type: E. cucurbitina
(Linnaeus) Baillon = A. latifolia (Miller) A. Gentry.

Smallto medium-sizedtrees. Leavessimple,alternate,bornesinglyon the

twigs. Inflorescencea solitary flower or a few-floweredfascicle terminalor
cauliflorouson bracteateshortshoots from oldertwigs and branches.Flowers
with the calyx large, rupturingirregularly,usually bilabiate, sometimessub-
truncate, rarely spathaceous; corolla greenish-white, variously tubular-
campanulate,usually with an inconspicuoustransversefold in the throat,
thickish;stamenssubexserted,the anthersglabrous,the thecae thick, some-
what divergent;pollen grains single, inaperturateto multiporate,the exine
microreticulate;ovary ovoid-elliptic,more or less lepidote, incompletely2-
locular. Fruita pepo or calabash,sphericalto ellipsoidal,indehiscent,with a
relativelyfragilewoody shell and soft pulp; seeds thick, large, more than 1.3
cm long and 1.4 cm wide, not winged,embeddedin the pulp.

Typespecies.A. macrophylla(Seem.)Miersex Baill.

Eighteen species ranging from Mexico and the West Indies to coastal
Venezuelaand Ecuador.Centeredin nuclearCentralAmericaand restrictedto
continental Central America from Mexico to extreme northwest Colombia
speciesA. latifolia.
exceptfor the wide-rangingwater-dispersed
Changesin the Code of Nomenclatureadoptedat the LeningradCongress
changethe status of nominarejiciendaand necessitatethe use of Amphitecna
ratherthan Dendrosicuswhen the formerEnallagmais mergedwith Amphi-
tecna(Gentry,1973b, 1976c).

Keyto Speciesof Amphitecna

1. Widespread from Mexico and the West Indies to Venezuela and Ecuador; fruit more or less
globose (sometimes ellipsoid in West Indies but less than 1.7 x as long as wide), apex
Amphitecna 51

rounded(sometimesbluntlyapiculatein WestIndies);leavescoriaceous;in CentralAmerica

alwaysnearsea level, mostlyin swampyplacesjust inlandfrom the mangroveassociation.
7. A. latifolia.
1. ContinentalCentralAmericato extremeNW Colombia;fruitellipsoidto ovoid (morethan
1.4 x as long as wide), moreor less pointedat apex;leavesmostlymembranaceous to sub-
coriaceous;not in swamps.
2. Calyxspathaceouswitha sharpacumen;budspointed.
3. Leavesoblanceolate,the tertiaryvenationnot obvious;ovarybilocularto above mid-
dle;GuatemalaandMexico. 17.A. steyermarkii.
3. Leavesnarrowlyelliptic, the tertiaryvenationintricatelyimpressedbelow; ovary bi-
locularonly at extremebase;Panama. 16.A. spathicalyx.
2. Calyxbilabiateor trilabiateor irregularlysubtruncate(occasionallymore or less spatha-
ceousbut withoutan acumenin A. montana);budsround-tipped.
4. Cauliflorous;leaves mostly more than 20 cm long (12-20cm long in A. parviflora);
fruitssometimesangledor costate.
5. Leavesmostly50-100cm long, oblanceolate;fruit(whereknown)angled.
6. Corollaevenlyinfundibuliform,not constrictedat middle,morethan 5 cm long;
Mexico. 13. A. regalis.
6. Corollacampanulateabovea tubularbase, constrictednearmiddle,2-4 cm long;
Guatemala. 9. A. megalophylla.
5. Leavesmostly20-50cm long (12-20cm long in A. parviflora),ellipticto narrowly
7. Leaves all oblanceolate, not intricatelyreticulate;corolla radiallysymmetric;
Mexicoabove 1000m. 8. A. macrophylla.
7. Leavesmostlyellipticto obovate,moreor less intricatelyreticulate(shagreened);
corolladistinctlybilabiatewitha transversefold or bulgein throat;Guatemalato
8. Corollaless than2.5 cm long, coveredby calyx for 2/3 to 3/4 its length;scan-
dentshrub;Panamaabove 1000m. 12.A. parviflora.
8. Corollamorethan3 cm long, coveredby calyxca. half its length;trees;Guate-
malato Panama,below 1000m.
9. Fruitlongitudinallycostate;corollaca. 3.5 cm long;Guatemala. 3. A. costata.
9. Fruitunribbed;corollamorethan4 cm long;Hondurasto Panama.
6. A. kennedyi.
4. Flowersterminalor on smallbranches;leavesless than 38 cm long; fruitsneverridged
or angled.
10. Corollatubular,less than 1 cm wide at mouthof tube, the rim hardlyexpanded;
below350m, Veracruzto Guatemala. 1. A. apiculata.
10. Corollacampanulateto infundibuliform-tubular, morethan 1 cm wideat mouth
of tube, the rim or lobes usuallyobviouslyexpanded;near sea level to 2000 m,
Veracruzto Colombia.
11. Pedicels6-10cm long; flowerspendent;calyxcircumscissilydeciduous,3-4cm
long;fruitellipsoid,veryinconspicuouslyshort-apiculate. 11.A. montana.
11. Pedicelsless than 4 cm long; flowers not pendent;calyx either not circum-
scissileor less than2.5 cm long; fruitovoid to narrowlyellipsoid,taperingto a
conspicuouslyapiculateor acuteapex.
12. Corolla less than 2.8 cm long, evenly and broadlyinfundibuliform-cam-
panulatewithout a basal tube; calyx less than 1.6 cm long; leaves mem-
branaceouswith conspicuouslyprominuloustertiaryvenationbeneath,less
than5 cm wide;lowlandsof east centralGuatemala. 4. A. donnell-smithii.
12. Corollamore than 2.5 cm long, campanulateor infundibuliformabove a
pronouncedbasal tube; calyx more than 1.4 cm long; leaves chartaceous
to coriaceous,the tertiaryvenationnot prominulous,relativelyobscure,at
least the largerleavesmorethan (4-)5cm wide; Mexicoto Colombia,sev-
eralspeciesonly above 1000m.
13. Calyx2.3-3.0cm long;CostaRicato Colombia.
14. Calyx thick, irregularlyand shallowlysplit; leaves dryinggray or
52 Flora Neotropica

brownishwithdarkermidvein;over 1000m elevation.

14. A. sessilifolia.
14. Calyxthin, splitto nearbase;leavesdryinggrayor olive withlighter
midvein;under1000m. 5. A. isthmica.
13. Calyx1.4-2.5cm long;Mexicoto Honduras.
15. Leaf tertiaryvenationconspicuouslywhitish-margined and intri-
catelyreticulate;extremeeast centralChiapasand northerncordil-
leraof Guatemala(800-)1000-1400 m. 15.A. silvicola.
15. Leaf tertiaryvenationnot whitish-margined nor conspicuouslyre-
ticulate;Veracruzto Honduras,below 1000m or above 1500m.
16. Corollaradiallysymmetric,4.5-5.5 cm long; calyx circumscis-
sile, caducous;Veracruz. 18.A. tuxtlensis.
16. Corollabilabiatewithventricosebulgein throat,3-4.5cm long;
calyxpersistent;centralChiapasto Honduras.
17. Calyxusuallysplit to nearbase; leaves coriaceousto sub-
coriaceous,secondaryveins7-14on a side, mostly 1-2.5cm
apart;centralChiapasto Belize,below 1000m.
2. A. breedlovei.
17. Calyx split less than halfway;leaveschartaceous,second-
ary veins (14-)15-18on a side, 0.5-1 cm apart;Honduras,
ca. 1600 m. 10. A. molinae.

1. Amphitecna apiculata A. Gentry, Rhodora 79: 435. 1977. Figs. 6, 7, 8.

Small tree 4-13 m tall; branchlets conspicuously angulate when young,

subterete when older. Leaves simple, alternate, narrowly obovate, acute to
acuminate at apex, more or less cuneate at base, (10-)15-38 cm long, (3-)5-11.8
cm wide, (some leaves of every collection at least 20 x 6 cm), subcoriaceous to
chartaceous, glabrous throughout except for a very few minute scattered lepi-
dote scales, midrib raised slightly above and prominently below, secondary
veins 10-15 on each side, usually separated by 1.5-2 cm, prominulous above
and prominent below, the main veins usually drying tannish against a grayish-
olive surface, petiole essentially absent. Flowers mostly terminal, sometimes in
part ramiflorous from bracteate short-shoots, borne singly or in fascicles of 2
or 3, the pedicels slender, 3.5-5 cm long; calyx campanulate, 16-20 mm long, 9-
10 mm wide, glabrous, irregularly 2-3-labiately split in upper third or half;
corolla white, more or less tubular, barely exceeding calyx, 2.2-2.8 cm long,
ca. 1 cm wide at top of tube, the lobes fused into an inconspicuous rim, lobes
stalked-lepidote outside, tube glabrous outside, glabrous inside except a few
stalked-lepidote glands near level of stamen insertion; stamens inserted ca. 10
mm from base of tube, the filaments 1-1.3 cm long, the thecae thick, hardly
divergent, 3 mm long; ovary ovoid, 2 mm long, 1.5 mm wide, densely lepidote;
unilocular except at base with the ovules on two parietal placentae; disc ca. 1
mm long and 3 mm wide. Fruit ovoid, more or less apiculate-acuminate at
apex, rounded at base, smooth or rough-surfaced, 6-11 cm long, 4-5 cm wide;
seeds thick, slightly bilobed, 11-14 mm long, 12-16 mm wide.
Type. Mexico. Tabasco: San Isidro, Balancan, Matuda 3388 (holotype,
MO 1172883; isotypes, MO 1198258, A, F, K, MICH,NY).
Distribution. From about 21? N latitude in Veracruz to Peten, Guate-
mala, in lowland moist forest mostly below 500 m altitude.
Amphitecna 53

MEXICO. Sin. loc.: Sesse & Mocino 2228 (F). Chiapas: 6-12 km S of Palenque, 300 m,
Breedlove 28886 (DS); Yaxchilan, 300 m, Breedlove 33928 (DS); Lacanja, 350 m, Breedlove 34472
(DS); mouth of Rio Ixcan, 300 m, Breedlove & McClintock 34091 (DS). Tabasco: Teapa, Linden
5a (P), 1608 (K). Veracruz: Sin. loc.: Hahn 120 (P); Estaci6n Biologica de Los Tuxtlas, Calzada
158 (GH); Cedillo & Calzada 66 (BM, MO); Catemaco, Cedillo 221 (F); entre San Rafael y
Nautla, 80 m, Gomez-Pompa & Riba 20 (MEXU); Colipa, Liebmann 587 (C, US); entre Yecuatla
y Colipa, 250 m, Martinez 54-4 (MEXU); Los Tuxtlas, Sousa 3298 (US), 4427 (MEXU); Hidalgo-
titlan, 200 m, Vazquez 67 (F); La Martinica, 50 m, Ventura3341 (F); Fortuno, 30-50 m, LI. Wil-
liams 8394 (F, MICH), 8580 (F).
GUATEMALA. Alta Verapaz: Rio Icvolay, 200-210 m, Steyermark 45080 (A, F). Peten: Rio
Chinaja, 50-70 m, Steyermark 45458 (A, F, MICH, NY); Rio Santa Isabel, 100 m, Steyermark
45819 (A, F, US).

Localname. Mexico:malocacao,tecomatillo.Guatemala:a-jo-nocht.

Two questionablecollectionswhichmay representaltitudinalextremesof

A. apiculataare not includedin the descriptionand distributionof this species.
Beaman 6404 (MO) from the summit of Cerro Vigia, Municipio Santiago
Tuxtla,Veracruz,at 950 m, has largermorerigidcoriaceousleavesthan other-
wise knownfor A. apiculata.This could provea distinctspeciesalthoughonly
buds havebeen seen to date. A secondproblematicalhigh altitudecollectionis
Breedlove34896(DS) from 1200m nearIxhuatan,MunicipioSolosuchiapa,in
northwesternChiapas.This collectionhas unusuallylargeleavesand an imma-
ture fruit. Both of these were previouslylisted as perhapsattributableto A.
steyermarkii(Gentry,1977b)but lack the oblanceolateleaveswith a gradually
long-taperingnarrowbasewhichprovedefinitivefor that species.A sterilecol-
lection from Alta Verapaz(Steyermark45080)whichwas listed as a paratype
of A. steyermarkiiis clearlyA. apiculataon the basis of its lowland habitat
and broaderleaf base.
Acceptanceof A. apiculatais contingentupon rejectionof Amphitecna
nigripesas a competingname for this species. Crescentianigripeswas first
listed by Lindenin 1863in his Cataloguedes plantes exotiques, No. 18. Lin-
den's inadequatedescription,entirelyof sterilematerial,describesthe leaves
as reaching75 cm in length and 25 cm in width, dimensionsfar in excess of
those of A. apiculata. These dimensionssuggest that Crescentianigripesis
synonymouswithAmphitecnamacrophylla.Althoughtherearetwo otherspe-
cies of Amphitecnawith leaves this large, one (A. megalophylla)was not
knownin Europeand the other (A. regalis)was describedin the samepublica-
tion as havingeven largerleavesover 1 m long and 30-35cm wide. Crescentia
nigripeswas duly recordedin IndexKewensis(incorrectlyciting Linden's1862
Catalogue(No. 17) ratherthan the 1863one) and has since been regardedas a
synonymof A. macrophylla.
Thereis a single leaf 44 cm long and 13 cm wide with a distinct,thick, 5
mm long petiolepreservedat BM as Crescentianigripes.This leaf derivesfrom
materialcollectedby Ghiesbrechtin Chiapasand its identificationis uncertain.
In size it fits A. macrophyllabetterthanany otherspeciesalthoughnone of the
(only six!) leaves I have seen of A. macrophyllahas nearly such a well-
demarcatedpetiole. One atypicalcollectionof A. megalophylla(see below)has
54 Flora Neotropica

B |!.ID1


:\ u



FIG. 6. Flowers of Amphitecna. A-B, A. apiculata (Steyermark 45508); A, corolla; B, flower. C-

D, A. parviflora (Hammel 2236); C, corolla; D, flower. E, A. costata (Steyermark 39130), flower,
F-G, A. donnell-smithii (Tuerckheim 11221); F, corolla; G, flower. H, A. montana (Breedlove &
Thorne 31010); I, A. steyermarkii (Steyermark 51252). J-K, A. megalophylla (Steyermark 44780);
J, calyx; K, corolla. L-O, A. silvicola (Breedlove 35106); L, calyx; M, corolla; N, x.s. near base of
ovary; 0, x.s. near top of ovary. (All x 5/7).
Amphitecna 55

a similarroundedleaf base but is otherwiseunlikethe "C. nigripes"leaf. This

leaf may be of A. apiculatawhichcan havea similarleaf base (especiallyin the
uplandform) but has smallerleaves. Althoughthis leaf probablyrepresentsa
plant distributedby Lindenas C. nigripes,it is probablynot the much larger-
leavedplant he earlierdescribedunderthat name. Its lack of agreementwith
the protologuesuggeststhat it shouldnot be consideredto typify C. nigripes.
Subsequently Baillon (1882) published an illustration and complete
descriptionof a plant cultivated at Paris as Crescentianigripes. Baillon's
plant, describedas havingleavesless than a foot long, is quite unrelatedto C.
nigripesLinden and is, in fact, A. apiculata, as proved by the illustration.
Thereare severaldiscrepanciesin Baillon'sdescription,notably his emphasis
of cauliflory,which suggestthat he may have confusedA. apiculataand A.
macrophylla.Cultivatedspecimensof both species are filed together as A.
nigripesat Paris, furthersuggestingconfusion. Additionallythere is a speci-
men in the Parisherbarium,annotatedby Sandwithas the type of Amphitecna
nigripes,whichis from Teapa, Chiapasand belongsto A. apiculata.A second
Linden collection of A. apiculata from Teapa, Chiapas, at Kew, has been
designatedby Sandwithas an isotypeof A. nigripes.Howeverit seemsunlikely
that Linden's own collections from Mexico, made in 1838 to 1840 (Anony-
mous, "Les ExplorationsBotaniquesde la Colombie," Belg. Hort. 17. 1867),
have any relevanceto the plant he introducedinto cultivationmore than 20
yearslater. In any event Baillon apparentlyconsideredthe plant he described
to be the same as CrescentianigripesLinden,though failingto cite an author,
as was his custom.
To summarize,it seemsprobablethat threeseparategatheringsbelonging
to two speciesare involved:An earlycollectionof A. apiculataby Lindenhim-
self representsa plantnot introducedinto cultivation.Two latercollectionsby
Ghiesbrechtwere introducedinto cultivationby Lindenthe first describedas
C. nigripesand belongingto large-leavedA. macrophylla,the second belong-
ing to A. apiculatabut also distributedas C. nigripes.
Rejectionof Crescentianigripesas applicableto A. apiculatadoes not
automaticallysolve the nomenclaturalproblem. Baillon's article also sug-
gestedthat Crescentianigripesbelongsto Amphitecna.An additionalquestion
must be resolved:Is Baillon'sAmphitecnanigripesintendedas a new combi-
nation based on Linden'sCrescentianigripesor should Amphitecnanigripes
be considereda validly publishednew name based on the plant cultivatedat
Paris?In the formercase, A. nigripes(Linden)Baill. is automaticallynomen-
claturallysynonymouswith A. macrophylla.In the lattercase Amphitecnani-
gripesBaill. wouldbe the oldest namefor A. apiculata.The secondinterpreta-
tion is furthercloudedby Baillon'sfailureto clearlyindicatewhetherhe con-
sideredCrescentianigripesor Amphitecnanigripesto be the correctname for
the taxon. Both the article and the illustrationare entitled "Crescentiani-
gripes."Amphitecnanigripesis used twice in the text and Crescentianigripes
once. In this context, Baillon's: "C'est probablementau genre Amphitecna
que doit etre egalmentrapportele pretenduCrescentianigripes(Fig. 99); il
devra donc prendrele nom d'Amphitecnanigripes" does not seem strong
enough to preventrejectionof A. nigripeseither under Article 34(1) of the
Code as not acceptedby its authoror Article34(3) as merelymentionedinci-
56 Flora Neotropica

1 ???-- ~'
~~.?:.. . ? : &... -

:I' ' ? ?1?


r~~. ~ ~ .. .. ''
? .I

t': :3:!iiiiiiii~

~:"-:! , ~~~~c
Amphitecna 57

FIG. 8. Map 1. Distribution of Amphitecna. A. apiculata (star in circle), A. breedlovei (closed cir-
cles), A. costata (open circles), A. isthmica (crosses), A. kennedyi (open stars), A. macrophylla
(closed squares), A. megalophylla (triangles), A. montana (closed stars), A. parviflora (open

dentally. Despite Baillon's omission of an author citation for Crescentia ni-

gripes, I opt to treat A. nigripes as based on the Linden basionym and thus
nomenclaturally a synonym of A. macrophylla.

2. Amphitecna breedlovei A. Gentry, Rhodora 79: 432 1977.

. Figs. 7, 8.

Small tree 7-8 m tall; branchlets conspicuously angulate when young, sub-
terete when older. Leaves simple, atnt, e, obovate to oblanceolate, acumi-
nate to rounded at apex, more or less cuneate at base, 5-18 cm long, 1.7-6 cm
wide, chartaceous to coriaceous, glabrous throughout except for scattered
lepidote scales, midrib noticeably raised above and especially below, second-

FIG. 7. Fruits of Amphitecna. A, A. costata (Steyermark 39130). B, A. montana (Breedlove

22260). C-D, A. breedlovei (Breedlove 24946 and Breedlove 38684, respectively). E, A. silvicola
(Breedlove 38800). F-G, A. apiculata; F, normal fruit (Breedlove 33928); G, fruit with diseased
surface (Breedlove & McClintock 34091). H, A. megalophylla (after Griggs, 1904). I, A. latifolia
(Liesner 1384). (From Gentry 1977b. All x 3/8).
58 FloraNeotropica

ary veins 8-14 on a side, prominentbelow, planeand hardlyvisibleabove, the

main veins below dryingtannishagainstan olive-graysurface, petiole essen-
tially absent. Flowersterminal,borne singly, the calyx 1.5-2 cm long, bilabi-
ately splitto nearbase, pedicel2.5-4.5 cm long; corollawhite, campanulate,3-
4 cm long, 1-1.8cm wide at mouthof tube, the lobes completelyfused;ovary2
mm long, 2 mm wide, disc 2 mm long, 7 mm wide. Fruit oblong-ellipsoidor
subglobose,more or less apiculate,smooth-surfaced,7-10 cm long (including
the up to 1.5 cm long apiculation),4.5-6 cm in diam.; seeds 1-1.6cm long, ca.
1 cm wide, distinctlylongerthanwide.
Type. Mexico. Chiapas: Cerro Tres Picos, 1000 m, Breedlove 24946
Distribution.A ratherpeculiardisjunctdistribution:A) between550 and
1000m on the southwestside of the Rio Grijalvadrainagefrom south central
to extremewesternChiapas.B) The MountainPine Ridgearea of west central
Belize.C) ExtremeeasternGuatemalanearbase of CerroSan Gil. Most collec-
tions are frompine or pine/oak forest.
MEXICO. Chiapas: 45 km N of Ocozocoautla, 550 m, Breedlove 32819 (DS, US); Tuxtla
Gutierrez-Nueva Concordia road, 850 m, Breedlove 34432 (DS); 46 km N of Ocozocoautla, 700 m,
Breedlove 38684 (DS); Rancho Concepci6n y Cerro Bruja, Miranda 5154 (US); cerca Tigrillo,
Miranda 6916 (MEXU, US).
GUATEMALA. Izabal: Rio Frio, 75 m, Steyermark 39944 (F, US), 75-150 m, 41587(F).
BELIZE. Without district: Peck 969 (GH), Schipp 867(F). El Cayo: Mt. Pine Ridge, Bartlett
11769 (F, MICH, US); Vaca, Gentle 2605 (MICH); Augustine, Hunt 405 (US); S of Puente, Lamb
107 (F); Augustine, Lundell 6618 (F, MICH, NY); Augustin, 650 m, Molina 189 (F, GH); All
Pines, Schipp 734 (A, F, GH, MO), Blancaneaux 1864 (A, MICH, NY); Silk Grass Creek Reserve,
Gentle 3093 (MICH); Baboon Ridge, Gentle 3218 (MICH).


It is possible that more than one species is included here. The lowland
Chiapas collections (below 800 m) match those from lowland Izabal,
Guatemala,in thinnerchartaceous,acuminateleaveswhichapproachthose of
A. donnell-smithii,while the Belizecollectionsare similarto the typicalform
from uplandChiapaswith coriaceousmostlysubacuteleaves. The acuminate-
leavedform is very like A. donnell-smithiibut has slightlythickerleaves with
less prominuloustertiaryvenationbeneaththan does that species. In fruit A.
breedlovei is similar to A. apiculata, but the fruits are usually shorter, the
leavessmaller,and the flowerslargerand morecampanulate.

3. AmphitecnacostataA. Gentry,Rhodora79: 431. 1977. Figs. 6, 7, 8, 9.

Small tree 3-14 m tall; branchletsangulate,the surfacesmooth, tannish.
Leaves simple, alternate, usually clustered near tips of twigs, obovate to
obovate-elliptic,sharplyacuminate,the acumen1-3 cm long, cuneateat base,
21-50 cm long, 6.5-18 cm wide, coriaceous, glabrous throughout, midrib
FIG. 9. Leaf variation in Amphitecna. A, A. regalis (Vazquez 105); B, A. costata (Steyermark
41796); C, A. donnell-smithii (Tuerckheim 2278); D, A. molinae (Molina & Molina 24295); E, A.
steyermarkii (Steyermark 51252). (All x 1/2).
Amphitecna 59

D^^ DB
60 FloraNeotropica

raised slightly above and conspicuouslybelow, secondaryveins 10-16 on a

side, plane above, prominulousbelow, tertiaryvenation perfectlyplane and
conspicuouslywhitish-marginedbelow (cf. A. silvicola), dryinggray or olive-
gray with the main veins darkerbelow, petiole poorly differentiated,occa-
sionally to 1 cm long but winged at top and mergingwith leaf base. Flowers
cauliflorous,mostlynearbase of trunk,mostlybornesinglyat the nodes from
fasciclesof smallbracts,the pedicels2-2.5 cm long, the calyxbilabiate,1.3-1.5
cm long, the lobes obtuseto subapiculate,glabrous;corollapale greenwith the
lobes completelyfused; stamenssubexserted,the antherthecae divergent,4-5
mm long; pistil ca. 3.5 cm long, the ovary oblong, 3-4 mm long, prominently
longitudinallystriate;disc annular-pulvinate,ca. 6'mmwide. Fruitcylindrical,
tapered to both ends, prominentlylongitudinallycostate, 22-36 mm long, 6-7
cm wide, the seedsembeddedin the whitishpulp.
Type. Guatemala.Izabal:Rio Juyama,50-100m, Steyermark39130, fl,
fr (holotype,F 1042987;isotypes,F 1042929,1055459).
Distribution.Apparentlyendemicto the lowlandsaroundCerroSan Gil
in extremeeasternGuatemalabetween50 and 100m altitude.
41796(F, US).
GUATEMALA.Izabal:Rio Tameja,50 m, Steyermark

Amphitecnacostatais most closelyrelatedto A. megalophylla.Theseare

among the largest-leavedmembersof the genus and are the only two Amphi-
tecnaspecieswith angulateor ridgedfruits. The two specieswereconfounded
in the Flora of Guatemala(underA. macrophylla)and the fruit illustratedfor
A. macrophyllais actuallythat of A. costata. The very differentleaves of A.
megalophyllaare much largerand oblanceolatewhile its fruit is shorterand
angulateratherthan long, cylindrical,and costateas in A. costata. Thesetwo
species are geographically, ecologically, and altitudinally separated, A.
megalophyllaoccurringfurtherwest at higheraltitudesand perhapsrestricted
to limestonesubstrates.

4. Amphitecnadonnell-smithii(Sprague)L. O. Williams,Fieldiana,Bot. 36:

22. 1973. Figs. 6, 9, 10, 11.
Crescentiadonnell-smithiiSprague,Bull.Herb.Boissier,ser. 2, 6: 376. 1906.
Enallagmadonnell-smithii(Sprague)Standley,Field Mus. Nat. Hist., Bot. Ser. 12: 361.
AmphitecnaoblanceolataL. O. Williams,Fieldiana,Bot. 36: 23. 1973. Type. Tun Ortiz
1459,Guatemala,Peten,fl, fr (holotype,F; isotypes,F, EAP, US).
Dendrosicusdonnell-smithii(Sprague)A. Gentry,Taxon22:640. 1973.

Smalltree 3-10 m tall; branchletsmoreor less terete.Leavessimple,alter-

nate, oblanceolate,acuminate(the acumento 3 cm long) acute at base, 4.5-15
(-22 in juveniles)cm long, 1.3-5 cm wide, membranaceous,glabrousthrough-
out, midrib prominulous above, prominent below, secondary veins plane
above, prominulousbelow, tertiaryvenation prominulousand reticulatebe-
low, petiolepoorlydemarcated,less than 1 cm long. Flowersmostlysingleand
terminal,the pedicels1.4-3 cm long, the calyxbilabiatelysplit to base, 1.2-1.6
cm long, the lobes obtuse to acutish,glabrous;corollagreenish-cream,evenly
Amphitecna 61

A B c


" 'i


FIG. 10. Fruits and flowers of Amphitecna. A, A. donnell-smithii (Tun 1459); B, A. steyermarkii
(Matuda 17649); C, A. isthmica fruit (Mori & Kallunki 2234); D, A. regalis flower (Vazquez 105);
E, A. macrophylla flower (Conzatti 8). (All x 3/4).
62 Flora Neotropica

and broadly infundibuliform, campanulate, the lobes almost completely

fused, 1.5-2.8 cm long, 1.1-2.3 cm wide at mouth of tube, essentially glabrous,
stalked glandular near base of lobes; stamens included, the anther thecae
slightly divergent, 2.5-3 mm long; pistil with the ovary ovoid, 2-3 mm long, 2
mm wide; disc annular, 1 mm long, 2-3 mm wide. Fruit narrowly ovoid, acute
to short-acuminate at apex, smooth, 13-14 cm long, 4.5-5 cm in diam., the
seeds thick, oblong, 10-11 mm long and broad, embedded in the whitish pulp.
Type. Guatemala. Alta Verapaz: Cubilquitz, Tuerckheim II 221 (JDS
7953), fl (holotype, K; isotypes, K, MO, P, US).
Distribution. Endemic to the lowlands of east central Guatemala between
50 and 350 m altitude in northern Alta Verapaz, extreme southern Peten and
northern Izabal Provinces.
GUATEMALA. Alta Verapaz: Finca Yalpemech, 100-150 m, Steyermark 45207 (F); Cerro
Chinaja, 150-700 m, Steyermark 45575 (A, F). Izabal: Rio Chac6n, Johnson 1241 (F, US); be-
tween Bananera and La Presa, 40-300 m, Steyermark 38210 (F), 50-150 m, 39199 (F). Peten: San
Luis, Tun Ortiz 1459 (F, EAP, US).

Amphitecna donnell-smithii is readily distinguished from A. breedlovei

(and from all other species of the genus) by its peculiar short corolla, broadly
hypocrateriform-campanulate from the narrow base rather than variously ex-
panded above a more or less tubular base as in other Amphitecna species. The
fruit of this species is longer than that of A. breedlovei. Vegetatively A.
donnell-smithii is very like the lowland form of A. breedlovei but seems to
have consistently thinner leaves with more prominulous tertiary venation
Williams (1973) separated A. oblanceolata from A. donnell-smithii on
account of its supposedly longer flowers. In fact the type of A. oblanceolata
actually keys out to A. donnell-smithii in the Flora of Guatemala and is mani-
festly conspecific with that species. Calyces of the type of A. oblanceolata are
1.2-1.5 cm long and its corollas are about 2.5 cm long; one of the US isotypes
of A. donnell-smithii (US 1322655) has calyces 1.2-1.5 cm long and corollas
reaching 2.8 cm long, actually exceeding the dimensions of the A. oblanceolata
type. Although A. oblanceolata falls into synonymy under A. donnell-smithii,
most of the material annotated by Williams as A. oblanceolata and included in
his concept of the species is actually referrable to A. breedlovei.

5. Amphitecna isthmica (A. Gentry) A. Gentry, Taxon 25: 108. 1976.

Figs. 8, 10.
Dendrosicus isthmicus A. Gentry, Phytologia 26: 442. 1973.

Small tree to 10 m. Leaves simple, alternate, glabrous, narrowly elliptic,

acute to acuminate, basally cuneate, 7-32 cm long, 2.8-10 cm wide, the midrib
raised beneath, the main veins drying whitish or yellowish beneath, leaf sur-
face gray or olive-gray above and beneath, the veinlets sometimes minutely
appressed beneath; petiole less than 5 mm long or lacking. Inflorescence of 1-2
flowers, usually terminal, the pedicel 2-4 cm long. Flowers with the calyx bil-
abiately split almost to the base, 2.3-3 cm long, each lobe 1.3-1.8 cm wide, ob-
Amphitecna 63


I i C r ;

I I --

FIG. 11. Map 2. Distribution of Amphitecna. A. donnell-smithii (closed stars), A. latifolia (closed
circles), A. molinae (cross), A. sessilifolia (triangles), A. silvicola (squares), A. spathicalyx (cross),
A. steyermarkii (open stars), A. tuxtlensis (star in circle).

tuse, glabrous with a few plate-shaped glands; corolla greenish-white, 3.4-6 cm

long, 1.7-2 cm wide at the mouth, the tube 2.4-3.5 cm long, glabrous, the rim
lepidote-papillate, the lobes fused into a reflexed rim; stamens subexserted,
inserted ca. 1.5 cm from the base of the tube, the anther thecae divergent, each
4-5 mm long and 1-2 mm wide; filaments 2-3 cm long; pistil ca. 4 cm long, the
ovary oblong, 2-3 mm long, ca. 2 mm wide, glandular-lepidote, 2-locular in
the lower 2/3 with the ovules several seriate on 2 indistinct axillary placentae in
each locule, above unilocular with parietal placentation; disc annular-pulvi-
nate. Fruit ellipsoid, apiculate at the ends, to 14 cm long and 7 cm wide; seeds
thick, embedded in the pulp.
Type. Panama. Chiriqui: San Bartolo de Limite, 400 m, Liesner 75, fl
(holotype, MO).
Distribution. Tropical wet forest below 1000 m alt. from Costa Rica to
northwestern Colombia, restricted to the Pacific slope in Costa Rica and
western Panama.

COSTA RICA. Alajuela: cataratas de San Ram6n, Brenes 13498 (F), 13797 (CR, F). San
Jos6: basin of El General, 675-900 m, Skutch 4825 (A, CR, MO, NY, US), 4940 (A, CR, F, MO,
NY, US). Puntarenas: Osa Peninsula, TSC Center, Mathias s.n. (C, F, US).
64 FloraNeotropica

PANAMA.Chiriqui:Progreso,Cooper& Slater277 (F, NY, US); roadto Barriles,Dressier

4901(MO).Cocld:La Mesa,900 m, Gentry7443(MO).Col6n:Rio Guanche,10-20m, Kennedy&
Foster2147(F, MO), 25 m, Moriet al. 5220(MO),20 m, Mori& Witherspoon7980(MO).Pana-
mi: El Llano-CartiRoad, 500m, Gentryet al. 8872(F, MO), 1100-1200ft, Mori& Kallunki4094
COLOMBIA.Antioquia:La Playona, Bernal I (COL);7 km from Turbo, Gentry9232

Amphitecna isthmica is closely related to upland A. sessilifolia from

whichit differsin its generallysmaller,narrowlyellipticratherthan oblanceo-
late to narrowlyobovate leaves, generallynarrowerfruit, and ecology as well
as in the charactersnoted in the key. Althoughdifferencesin calyx thickness
and degreeof splittingare relative,they appearto be constant. The uneven
splittingand thicknessof the calyx of A. sessilifoliausuallyresultin a whitish
edgingalong the split edges of its black-dryingcalyx, especiallynear the base
of the splits;in A. isthmicathe thinnercalyx lacks a whitishedgingalong the
linesof rupture.
I cannot alwayspositivelyseparateA. isthmicafrom A. kennedyion the
basis of sterileor fruitingmaterial.Terminal(noted as terminalor caulineon
one collection) placementof flowers, usually much smallerleaves, and geo-
graphicseparationgenerallydistinguishit from that species.Basedon a single
dissectionfor each species, the ovary of A. isthmicais bilocularto above the
middleratherthanonly at the extremebaseas in A. kennedyi.
For a discussionof the differencesbetweenA. isthmicaand A. parviflora
of intermediatealtitudes,see underthat species.

6. Amphitecnakennedyi(A. Gentry)A. Gentry,Taxon25: 108. 1976.

Figs. 8, 12.
DendrosicuskennedyiA. Gentry,Phytologia26:441. 1973.

Small to medium-sizedtree to 15 m tall. Leaves simple, alternate,glab-

rous or sparselylepidote, oblanceolateto obovate, apically acute to acumi-
nate, basallycuneate,attenuate,the petiolenot differentiated,16-38cm long,
5-14 cm wide, the midribconspicuouslyraisedbeneath,the main veins drying
whitish beneath, leaf surface gray above and grayish beneath, the veinlets
sometimesminutelyimpressedbeneath.Inflorescence1-2 flowersfrom a con-
spicuouslybracteateabbreviatedshort-shooton older branchesor trunk, the
pedicels1.5-3 cm long. Flowerswith the calyx bilabiatelysplit almostto base,
1.5-3 cm long, each lobe ca. 1 cm wide, conspicuouslyapiculate; corolla
greenish-white,4.3-4.8 cm long, 1.3-1.9cm wideat the mouth, the tube 2.7-3.5
cm long, the lobes fused into a reflexedrim; stamenssubexserted,the anther
thecaesomewhatdivergent,4-5 mm long, 1.5-2mm wide, the filaments2.4-2.8
cm long, inserted1.3-1.5cm fromthe base of the tube;pistil ca. 4 cm long, the
ovaryrounded-conical,2-3 mm long, 2 mm wide, denselylepidote,the ovules
severalseriateon 2 parietalplacentae;disc annularpulvinate.Fruitellipsoid,
apiculateat the ends, 11.5-13cm long, 5.5-7 cm wide; seeds thick, wingless,
1.3-1.5cm long, embeddedin the pulp.
Type. Panama. Bocas del Toro: Fila de Almirante,Kennedy& Dressier
1258, fl (holotype,MO).
Amphitecna 65

FIG. 12. Amphitecna kennedyi and A. spathicalyx. A-B, A. kennedyi; A, habit x 3/10; B, inflor-
escence x 3/5 (Kennedy & Dressier 1258). C-G, A. spathicalyx, C, flower x 3/5 (Gentry 5769);
D, fruit x 1/2 (Dressler3943); E, corolla x 3/5; F, ovaryx.s. x 5/6; G, ovaryand disc x 1-1/5
(Gentry5769).(FromFloraof Panama).
66 FloraNeotropica

Distribution.Tropicaland premontanewet forest below 1000m alt. from

Hondurasto centralPanama,exclusivelyon the Caribbeanslope.
HONDURAS. Atlantida: Mt. Cangrejal, 800 ft, Yunckeret al. 8724 (F, GH, MO, NY, US).
COSTA RICA. Lim6n: between BriBri & Rio Sixaola, 50-100 m, Burger et al. 10479 (F); 3
km E of El Carmen, 10 m, Lent 2427 (F).
PANAMA. Bocas del Toro: Cooper & Slater 36 (US); Fila de Almirante, Kennedy &
Dressier 1261 (MO, PMA); Quebrada Huron, Kirkbride & Duke 615 (MO); Fila Almirante, 100-
200 m, Nee & Hansen 14095 (MO), von Wedel 2258 (GH, MO). Panami: Cerro Jefe, 800-900 m,
Gentry & Mori 13415 (MO); Altos de Pacora, 750 m, Mori & Kallunki 4967 (MO).

Localname. Panama:Tapitode monte.

Threeof the collectionscited above-the Honduranone from Mt. Can-

grejal,AtlantidaDept. and two from CerroJefe, PanamaProvince,Panama-
differ from other materialof A. kennedyiin muchthickermore nearlyelliptic
leaves with the venation beneath more stronglywhite-margined.Since these
collectionsare at the two opposite extremesof the geographicdistributionof
the speciesit seemsunwiseto accordthem taxonomicrecognitiondespitetheir

7. Amphitecnalatifolia(Miller)A. Gentry,Taxon25: 108. 1976.

Figs. 7, 11. 13.
Crescentia latifolia Miller, Gard. Dict., ed. 8. 306. 1768.
C. cucurbitina Linnaeus, Mant. P1. 2: 250. 1771. Type: Based on C. latifolia Miller.
C. lethifera Tussac, Fl. Antill. 4: 50.1827. Type illustration: Fl. Antill. 4, pl. 17 (as C. toxi-
C. toxicaria Tussac, Fl. Antill. 4: 50, tab. 17. 1827. (alternate name for C. lethifera).
Dendrosicus saxatilis Rafinesque, Sylva Tell. 80. 1838. Type: Based on C. cucurbitina Lin-
Crescentia obovata Bentham, Bot. Voy. Sulphur 130. 1844. Lectotype: Colombia, Isle of
Gorgona, Barclay 935 (K; isolectotype, BM).
C. palustris Forsyth ex Seemann, Bot. Voy. Herald 183. 1854, pro syn.
C. coriacea Miers, Trans. Linn. Soc. London 26: 177. 1870. Syntypes: Jamaica? P. Browne
s.n. (BM); Jamaica, Prior s.n. (K).
C. cuspidata Miers, Trans. Linn. Soc. London 26: 178. 1870. Lectotype: Venezuela, Fendler
780 (K; isolectotype, GH).
C. elongata Miers, Trans. Linn. Soc. London 76: 178. 1870. Type: Cuba, Wright 361 (holo-
type, K; isotypes, GH, MO).
C. cucurbitina var. heterophylla Kuntze, Rev. Gen. PI. 2: 479. 1891. Type: St. Thomas,
Kuntze s.n. (not seen).
Enallagma cucurbitina (Linnaeus) Baillon ex Schumann, In: Engl. & Prantl, Nat. Pflanzenf.
4(3b): 247. 1894.
E. obovata (Bentham) Baillon ex Schumann, In: Engl & Prantl, Nat. Pflanzenf. 4(3b): 247.
E. latifolia (Miller) Small, Fl. Miami 171. 1913.
Amphitecna obovata (Bentham) L. O. Williams, Fieldiana, Bot. 36: 25. 1973.
Dendrosicus latifolius (Miller) A. Gentry, Taxon 22:638. 1973.

Small scragglytree to 10 m tall and 18 cm dbh., the branchesusually

crooked, the crown open; twigs subtereteor 3- or 4-angled with decurrent
Amphitecna 67


~~~~~~~~~~~~!.. '

.. ..:t


t ~~~~~~~~~~~~~~~~~~~~~
~ ~ ~
? t

?? .
:. ,.......~..-~~~f.
FIG. 13. Amphitecna hab.i.B,
latifolia.A, t F.orer C.
Afte..frit;etry, seed D,. ?79 clyx. .

co? after . i~~~~~~~~~~~~~{

. :':'
': ?E.'
-. ....... . . . ..~~~~~~~~~~~~~~~~~~~I
.. ... :.

~'' ...~~~I? ' ?:'

:: ?";~ ~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~



:" /~ '..t'?
r'..:, '~
? . ?


" . . . ..r I : ,..

i ..,~,,~?~ ~ ~,,.~ ~ ~ ~ i/ ~ ~ ~ ~ ~ ~ ~~~~~~~'" :?'
? ,? ?

? ~~ ~ . . .e~
'.. -? '? ~~~pr??f:~~~~~~~
. ,


FIG. 13 Amphilcna latfolia. , 11QU?? ?r, B fruit C, see D,"...........

.. . ? . '..'.79
corolla after Dwyer 1~~~~~~68.
Al /. ''m lrao Eu r."
68 FloraNeotropica

ridgesbelow the nodes. Leavessimple, alternateto subopposite,obovate, the

tip roundedto almost acute, usuallyapiculate,the base cuneate-attenuate,7-
19cm long, 3.3-10.6cm wide, rigid-chartaceous to coriaceous,secondaryveins
6-15 on a side, the midveindepressedor plane above, raisedbelow, glabrous
beneath,with plate-shapedglands especiallyat the base, dryinggrayish-olive
to brownish,the petiole poorlydefined, more or less winged,0.2-1.1 cm long,
glabrous.Inflorescencea single flower or fascicleof 2-3 flowersborne on the
largertwigs, pedicelsglabrous, 2.2-3.6 cm long. Flowerswith the calyx split
into subequal anterior and posteriorhalves, 2.5-3.7 cm long and 1-2.5 cm
wide, essentiallyglabrous;corolla greenish-white,tubular with a transverse
fold midwayacross the lower side of throat, fleshy, 3.8-6.2 cm long, 1.2-2.4
cm wide at mouth, the tube 2.0-4.0 cm long, the lobes 0.8-1.5 cm long, upper2
and lower 3 lobes respectivelyfused, mostly glabrous,stalked-lepidoteon the
lobes outsideand at the level of stameninsertioninside;stamenssubexserted,
the antherthecae thick, only partiallydivergent,5-6 mm long, the filaments
2.4-2.6 cm long, inserted1.5-2cm frombase of tube, the staminodeca. 10 mm
long, insertedca. 10 mm from base of tube; pistil ca. 5 cm long, the ovary
rounded-conical,4-5 mm long, 3 mm wide, lepidote,the ovules ca. 4-seriatein
each locule of an incompletelybilocularovary, placentationparietalonly at
extremeapex; disc pulvinate, 2 mm long and 5 mm wide. Fruit a pepo or
calabash, more or less sphericalwhen mature, 6-9 cm in diam., glabrous,
minutely papillate with scatteredsunken plate-shapedglands; seeds thick,
corky, 1.3-1.6cm long and 1.6-1.8cm wide, withoutwings.
Type. Plumier,Plant. American.,pi. 109.
Distribution.Circum-Caribbean from the south tip of Florida,Veracruz,
Mexico and the GreaterAntillesto Venezuela,also on the Pacific coast from
Costa Rica to northernEcuador.Restrictedto coastal beachesand mangrove
fringesexceptin Venezuelaand on the GreaterAntilles.
UNITEDSTATES.Florida(DadeCo.): Miamivic., Britton451 (F), Caldwell8785(A, US),
Churchills.n. of 1970(MSC),Curtiss.n. of 1881(A, K),s.n. of 1895(US), 5838(A, F, GH, K, P,
US), Garber59 (GH, MO, US), Hitchcock2624 (F), Mell s.n. of 1911(US), Moldenke721 (K),
Porters.n. (F), Rehder776(A), Rolfs & Quaintance189(F), Simpson540 (GH, US), Sargents.n.
of 1886 (A), Small et al. 6767 (US), 7872 (GH), Thurbers.n. of 1859 (GH), Stearns42 (US),
Young413 (US).
MEXICO.Tabasco:cercaChiltepec,Rovirosa728(US). Veracruz:Playade La Barra,Son-
tecomapan,Calzada624 (F, GH); La Palma, acahual,Calzada794(F); lado sur de la lagunade
Sontecomapan,manglar,Lot 1283(GH);La Palma,Sontecomapa,0 m, Sousa3102(MEXU).
GUATEMALA.Izabal:Jocolo, Johnson 1106 (F, US); Lago Izabalopposite San Felipe,
Steyermark 39689 (F).
BELIZE.Belize:Sibun River, Gentle1601 (A, K, MICH, MO, US); GracieRock, Gentle
1785(A, BM, F, GH, K, MO).Toledo:PuntaGorda,Dwyeret al. 606(MO);TemashRiver,Gen-
tle 4651(MICH);Kinloch39 (F), 40 (F), Stevenson156(F).
HONDURAS.Atlantida:HighlandCreek,nearPuertoSierra,P. Wilson369 (US).
COSTARICA.Lim6n:betweenCauhita& Lim6n,0-10m, Baker&Burger7 (MO,U); Cau-
hita, Gentry1283(MO),1291(MO);10kmNW of Lim6n,Godfrey66377(MO);Portete,Jimdnez
2879 (F); Puerto Lim6n, Pittier 16004(GH); entre BarraParisminay Rio Chiquero,Shank &
Molina4305(F, GH), La Barra,Solfs555 (CR, UCR). Puntarenas:Delta of Rio Esquinas,Alien
5639 (F); Cabo Blanco,0-200m, Burger& Liesner6579(BM, F), 6694(F); nearRinc6n,Gentry
1244 (MO); Punta Guapinal,A. Jimenez 1679 (CR, NY); Curfi, O. Jimenez 1192 (CR); Osa
Peninsula,vic. of Rinc6n,Lent 469 (NY), Raven21526(F), SalasS-375(UCR), S-595(UCR), S-
Amphitecna 69

994 (CRU); Playa Blanca Golfo Dulce, Valerio 463 (F), 464 (CR); Lagartos Bay, Barclay 2768
PANAMA. Bocas del Toro: Isla Col6n, von Wedel 561 (GH, MO, U); Water Valley, von
Wedel 988 (GH, MO); Old Bank Island, von Wedel 2101 (GH, MO, US); Johns Creek, von Wedel
2769 (GH, MO, US). Canal Zone; summit Garden (cult.), Croat 13827 (MO, SCZ); Chagres,
Fendler 210 (K); 1 mi NE of Pifla, Lewis et al. 1842 (MO); along Rio Indio de Gatfin, Pittier 2782
(US); vic. of Fort Sherman, Standley 30919 (US), Tyson & Blum 3797 (MO, SCZ). Col6n: vic. of
Camp Pifla, Allen 3632 (MO); 3 mi SW of Col6n, Croat 14168 (MO); W of Portobelo, Gentry
1742 (MO); Aspinwall, Hayes 862 (NY); Punta Pedro, Icacal between Salud and mouth of Rio
Indio, Howell 40 (MO); Salud, Lao & Holdridge 250 (MO); Rio Indio de Fat6, Pittier 4272 (US);
Plaza Col6n, Rose 22028 (US), 22078 (NY). Darien: on Rio Tuira ca. 3 mi NW of El Real, Duke
4820 (SCZ); S of Jaque, Gentry 4121 (BM, MO); Pto. St. Dorotea, Dwyer 2272 (MO); Rio Tuira
below El Real and Piriaque Island, Stern et al. 891 (GH, K, MO). Los Santos: Azuero Peninsula
near Raja Point, Ostenfeld 20 (C, US); Punta Mala, Tyson 2738 (MO, SCZ). Panama: San Jose
Island, Erlanson 4 (GH, NY, US), Harlow 97 (P, US), Johnston 10, 1179 (both GH), Stimson
5333 (NY, PMA, SCZ), Tyson & Loftin 5074 (SCZ); seaside, Panama, Hayes 826 (NY). San Bias:
Mulatupo, Duke 8552 (MO); near Puerto Obaldia, Pittier 4362 (US). Veraguas: Isla de Coiba,
Dwyer 1568 (MO); Foster 1617 (F, PMA), Kelsall 467 (K). Sin. loc.: Barclay s.n. (BM, K); Hayes
44 (GH).
CUBA. Habana: W of Bataban6, Bro. Le6n 13408 (GH, US). Isla de Piftos: Ensanada de
Siguanea, Britton & Wilson 14858 (US); Nueva Gerona, Killip 44827 (US), 45550 (US). Las Villas:
Hanabanilla Falls, Britton et al. 4869 (F); San Bias, Hunnewell 11580 (GH); Belmonte, Soledad,
Jack 5014 (A, US); San Bias 600-800 ft, Jack 7340 (A, US); Soledad, Rehder 1204 (A, P); vic. of
Sancti Spiritus, Shafer 12135 (MO, U, US). Oriente: near Miranda, Bro. Clemente 2503 (GH); El
Gato, 1000 m, Bro. Clemente 3220 (US); Chivirico, Bro. Clemente 3445 (GH); Cayo Rey, Cle-
mente 4868 (GH); Manacal, Bro. Hioram 7635 (GH, US); S of Holguin, Shafer 1249 (GH, US),
1746 (F); Lebisa Bay to El Purio, Shafer 3425 (F, US); prope Monte Verde, Wright361 (BM, GH,
K, MO, P), 361a (GH). Pifiar del Rio: Rangel, 500 m, Bro. Alain 2701 (GH, US); Sierra de Anafe,
Wilson & Leon 11536 (US); vic. of Punta de Afuera, Wilson 9265 (US). Prov. not known: Santa
Catalina, Caldwell & Baker 7059 (F); de la Sagra s.n. (P); Otto 357 (K, P); Seemann 644 (BM).
JAMAICA. Cornwall: Black River Morass, Proctor 24516 (BM); Great River, 0 m, Procter
32661 (BM, F). Middlesex: 2.5 mi W of Runaway Bay, Proctor 36493 (BM); White River, Alexan-
der Prior s.n. (K); Galina Point, 10 ft., Stearn 772 (A, BM); Roaring River 5-10 ft, Steam 738 (A,
BM). Surrey: near Port Antonio, 0 m, Fredholm 3312 (US); Rio Grande, 600 ft, Harris 5983 (F);
Swift River, Harris 5986 (F). Sin. loc.: W. Wrights.n. (BM); Justin s.n. (K).
HAITI. vic. of Jean Rabel, Leonard & Leonard 12635 (US); Tortue Island, vic. of Basse
Terre, Leonard & Leonard 14027 (K, MO); vic. of St. Louis du Nord, Leonard & Leonard 14264
(US), 14411 (GH, MICH, US). Sin. loc.: Jaeger 207 (K).
REPUBLICA DOMINICANA. Sin. loc.: ex herb. Poiteau s.n. (P); vic. of San Francisco de
Macoris, 400-1000 m, Abbott 2202 (US); Gaspar Hernandez, J. Jimenez 1991 (US); Wrightet al. 7
PUERTO RICO. Rio Grande to Fajardo, Britton et al. 7029 (US); Moca, Sargent A67 (US);
Vieques Island, Shafer 2528 (US); near Dorado, Miller 1621 (US); Bayam6n, Sintensis 920 (BM,
C, GH, K, P, US); Aguada, Sintensis 5586 (BM, C, GH, K, US).
VIRGIN ISLANDS. St. Croix: Borgsen s.n. of 1906 (C), Eggers 657 (C), s.n. of 1870 (C),
Howard 15664 (A, U), Little 16402 (BM, GH, US), Raunkiaer s.n. (C), West s.n. ex herb. Vahl
(C). St. Jan: Raunkiaer s.n. of 1906 (C, P). St. Thomas: Oersted s.n. (C, P, U, US), 1131 (C),
7961 (C); ex herb. Ricole (P).
WINDWARD ISLANDS. Martinique: Belanger 606 (P), 2359 (P); Basse Pointe, Hahn 1394
(BM, GH, K, P, US), 1436 (P, in part), Plee s.n. cult. (P), 660 (P). St. Vincent: Anderson s.n. ex
herb. Forsythe (K), H. Smith & G. Smith s.n. (K).
COLOMBIA. Antioquia: Rio Le6n, 0 m, Feddema 2058 (MICH). Choc6: Titumate, 0 m,
Dawe 866 (K); 3-7 km S of Curiche, 0 m, Duke 9649 (COL); Rio Baud6, Fuchs et al. 21702 (COL);
entre Nuqui y Pangui, Ferndndez 246 (COL); mouth of Rio El Valle, 0 m, Gentry & Forero 7379
(COL, MO), Gentry & Fallen 17494 (COL, MO); Quebrada Togoroma, 0 m, Killip & Cuatrecasas
39134 (COL, F); Jurad6, Romero-Castafleda 470 (COL). El Valle: Buenaventura Bay, Killip
33012 (A, COL, MO). Narifio: Tabacal, Dryander 2607 (K); Gorgona Island, 0 m, Killip & Garcia
70 Flora Neotropica

33122 (COL); Barclay 926 (BM), s.n. (K), Hinds s.n. of 1841 (K), Kelsall 547 (K); Rio Mira, Bar-
clay 852 (BM, US).
VENEZUELA. Sin. loc.: Cono d'Tigre, Moritz 1953 (BM). Aragua: Colonia Tovir, Fendler
780 (GH, K); Ingomar, Pittier 14251 (VEN); Playa de Turiamo, Trujillo 4129 (MV). Carabobo:
La Cumaca, Alston 5896 (BM); alrededores de Mor6n, Schnee 1166 (MV). Distrito Federal: Chi-
chiriviche, Aristeguieta 5444 (VEN); Los Caracas-Carnao, Aristeguieta 7118 (VEN); Los Caobos,
Hoyos 2214 (LaSalle); near Caracas, Bro. Elias 141 (A), Linden 113 (P); E de NaiguatS, 5 m, Little
16102 (VEN); Naiguata, P. Rodriguez 7 (VEN); Rio Los Caracas, Steyermark & Rabe 95961 (NY).
Miranda: Los Chorros, Delgado 288 (F, US, VEN); around Dos Caminos, 800-1300 m, Pittier
5922 (C, P, US); 5.5 km 0 de Aricagua, Steyermark & Carrefo 106890 (F, US, VEN); Los
Chorros, LI. Williams 12243 (F).
TRINIDAD. Baker s.n., cult. (K); Grisebach 1325, cult. (K); Sieber 277 (MO, P).
GUYANA. Georgetown, cult., Mckenzie & Watkins49 G-76 (K).
ECUADOR. Esmeraldas: San Lorenzo, Little 6339 (F, K, US).

Local name. Florida: Black calabash. Cuba: magiiira, giiira de olor.

Puerto Rico: higiierillo. Jamaica: savana calabash. Belize: Morito de rio, wild
calabash, river calabash. Guatemala: mora de costa. Costa Rica: jicaro,
jicarita, calabacillo de playa, jicaro de estero. Panama: wild calabash,
tutunuto, calabash tree, mariquita de Maria, totumito, totumillo. Colombia:
totumillo, matecillo, matesillo,matecito, totumito, calabasillo. Ecuador: cala-
Uses. The fruit is an important food for the paca and is often used medi-
cinally. The seeds are used as a purgative (Duke 4820) and the pulp as a laxa-
tive (Leon & Alain, 1957) and remedy for chest infection (Leon & Alain, 1957).
The wood is occasionally used for plows and similar construction in Cuba.

8. Amphitecna macrophylla (Seemann) Miers ex Baillon, Rev. Hort. 1882:

465. 1882. Figs. 8, 10.
Crescentia macrophylla Seemann, J. Bot. (Hooker) 6: 274. 1854.
C. regia hort. ex. Seemann, Trans. Linn. Soc. London 23: 20. 1862. (nom. nud., pro syn.)
C. nigripes Linden, Cat. 18: 8. 1863. Type: Cultivated, from Mexico (not seen).
Ferdinandia superba hort. ex Seemann, Trans. Linn. Soc. London 23: 20. 1862. (nom. nud.,
pro syn.)
Amphitecna nigripes (Linden) Baillon, Rev. Hort. 1882: 465. 1882.
Enallagma macrophylla (Seemann) Lundell, Wrightia 4: 170. 1971.
Dendrosicus macrophyllus (Seemann) A. Gentry, Taxon 22: 639. 1973.

Small pachycaul tree 2-7 m tall, the leaves in a terminal cluster; branchlets
thick, rather angular. Leaves alternate-verticillate, congested near branch
apex, oblanceolate, obtuse to acuminate at apex, cuneate-attenuate at base,
very large, 28-53 cm long, 5-13 cm wide, subcoriaceous, glabrous, midrib con-
spicuously raised below, somewhat flattened above, secondary veins plane
above, prominent beneath, inconspicuously white-margined, drying olive, the
tan main veins contrastingly lighter below; petiole poorly demarcated, to 1.5
cm long in smallest leaves. Inflorescence a few cauliflorous flowers from a
thick sessile densely bracteate short-shoot, the pedicels 1.5-2(-3.7 fide
Seemann) cm long, glabrous. Flowers with the calyx bilabiately split to near
base or split only 1.5-1.7 cm on one side, 2.2-3.5 cm long, each lobe 0.8-1 cm
wide, obtuse to acutish, glabrous with a very few plate-shaped glands; corolla
Amphitecna 71

white, tubular-campanulate, more or less radially symmetrical, 3.7-5 cm long,

1-1.5 cm wide at mouth of tube, the lobes fused into a frilly-margined rim,
stalked-lepidote on outside of lobes, otherwise glabrous; stamens inserted 1-
1.7 cm from base of tube, the anthers subexserted, the thecae slightly diver-
gent, 4-5 mm long, filaments 1.3-2 cm long; pistil 2.5-3 cm long, the ovary
ovoid, ca. 3 mm long, to 2 mm wide at base, slightly glandular-lepidote, 2-
locular at base, unilocular at apex, disc annular-pulvinate, 1 mm long, ca. 4
mm wide. Fruit (after Graebener, 1899) narrowly ellipsoid, acuminate, 14-16
cm long, 4-4.5 cm wide, the seeds thick, 12 mm long, 14 mm wide.
Type. Specimen without data from plant of unknown origin cultivated at
Kew (K).
Distribution. Middle altitudes around 700-1300 m in the coffee zone of
central Veracruz,Mexico. Collected only twice since 1908 and perhaps nearly ex-
tinct; once rather widely cultivated under glass in Europe.
MEXICO. Veracruz: Cordova, Orizaba, Muller 2201 (NY); Zacuapan, Purpus 4319 (BM, F,
GH, MO, US); Cerro de Chiquihuite, 4000 ft, Conzatti 8 (GH); Cerro de Cuhuatepetl, Tehuacan,
1500-2000 ft, Santos 3644 (MICH); El Mirador, mpio. Totutla, 1000 m, Ventura8416 (ENCB).
CULTIVATED: Paris "cult. serre" 1859 (P); "serres du Museum" 1881 (P); "jardin de
l'Ecole du Medicine" 1895 (P).

9. Amphitecna megalophylla (Donnell Smith) A. Gentry, comb. nov.

Figs. 6, 7, 8.
Neotuerckheimia megalophylla Donnell Smith, Bot. Gaz. (Crawfordsville) 47: 258, f.l. 1909.

Small pachycaul tree 3-10 m tall, unbranched or few-branched, the leaves

in a terminal cluster; branchlets thick, somewhat angular, often ridged, rather
corky-barked. Leaves alternate, mostly congested near branch apex, oblanceo-
late to narrowly obovate, acute to short-acuminate at apex, cuneate-attenuate
at base, gigantic, (25-)50-100 cm long, (5.5-)8-25 cm wide, most leaves about 75
cm long and 15 cm wide, chartaceous to subcoriaceous, glabrous, midrib
slightly raised above, very strongly raised below, secondary veins more or less
plane above, prominent beneath, conspicuously white-margined, drying olive
(blackish when young) with the main veins lighter below; petiole poorly de-
marcated, to 3 cm long, to 1 cm thick. Inflorescence a cauliflorous fascicle of
1-several flowers from a bracteate short-shoot, usually rather low on the
trunk, the pedicels 2.5-5.5 cm long, glabrous. Flowers with the calyx campanu-
late, 1.3-1.8 cm long, 1.2-1.5 cm wide, bilabiately split less than halfway to
base, the lobes rounded, glabrous; corolla pale green or greenish-white,
abruptly hypocrateriform-campanulate above a 1.5-2.5 cm long tubular base,
strongly arcuate especially at top of basal tube, 2-4 cm long, 0.8-1.6 cm wide at
mouth of tube, the lobes more or less fused, "wrinkled and ruffled," with
stalked-lepidote glands on lobes, otherwise glabrous; stamens inserted 1-1.5
cm from base of corolla tube; anthers exserted, the thecae somewhat diver-
gent, ca. 5 mm long; pistil curved, ca. 1.8 cm long, the ovary ovoid, conspicu-
ously ridged, 2 mm long, 1.5-2 mm wide, glabrous or with a few minute lepi-
dote scales, unilocular with 2 protruding parietal placentae from apex to very
near base, bilocular with axile placentation at extreme base; disc annular-
72 Flora Neotropica

pulvinate, lobed, 1.5-2 mm long, 4 mm wide. Fruit oblong-ellipsoid, obtusely

but distinctly 5-8-angled, abruptly very short apiculate at apex, obtuse at base,
6-16 cm long, 5-6 cm wide; seeds thick, wingless, ca. 8-11 mm long and 15-16
mm wide.
Type. Guatemala. Alta Verapaz: Coban, 1350 m, von Tuerckheim I 2278
(lectotype, US; isolectotype, M).
Distribution. Wet forests in the coffee zone on the Caribbean-facing
north slope of the Guatemalan Sierra de las Minas between 300 and 1400 m alt.

GUATEMALA. Alta Verapaz: near Finca Sepacuit6, Cook 120 (US), Cook & Griggs 158
(US), 575 (US); Chamal, Rio Coban, 1200 m, Molina & Molina 12181 (F); Saklak River, Secan-
quim, 300 m, Pittier 295 (GH, NY, US); near Coban, 1260-1440 m, Standley 71591 (F); Rio
Carcha between CobAn and San Pedro Carcha, 1360 m, Standley 89986 (F); Rio Carcha between
Coban and San Pedro Carcha, 1360 m, Standley 90114 (F); Chirriacte, 900 m, Standley 91856 (F),
91881 (F); Rio Icvolay to Quebrada Diablo, 300-350 m, Steyermark 44780 (F, NY, US); Cubil-
quitz, 350 m, von Tuerckheim 8723 (US, syntype); Rio Chiacte near Finca Volcan, Wilson 274 (F).
Izabal: between Los Amates and Izabal, 2000 ft, Kellerman 7168 (NY).

Local name. O-hoch.

Amphitecna megalophylla, A. macrophylla, and A. regalis are absolutely

distinctive in the family in their pachycaul habit with a terminal tuft of gigantic
leaves; vegetatively they are thus similar to certain species of Gustavia, Grias,
and Cespedezia. The fruits are strikingly like those of Theobroma cacao as
pointed out by Griggs (1904).
This Guatemalan species has long been treated as conspecific with Mexi-
can A. macrophylla (Seibert, 1940; Standley & Williams, 1974). However, its
small, bent, medially constricted corolla, short broad calyx, and ridged fruit
are completely different from those of A. macrophylla.
One collection which may be referrable to A. megalophylla-C. L. Wil-
son s.n. from Finca Volcan, Alta Verapaz, Guatemala-has not been included
in the description and distribution. This specimen has similarly very large
leaves but their bases are distinctly rounded above a well-demarcated 3-10 mm
petiole. The twig is also smoother and shinier than in other A. megalophylla
collections. The detached flowering material is identical to that of A. megalo-
phylla. Whether this represents an aberrant (juvenile?) growth form of A.
megalophylla or a new taxon is unclear.

10. Amphitecna molinae L. O. Williams, Fieldiana, Bot. 36: 22. 1973.

Figs. 9, 11, 14.
Dendrosicus molinae (L. O. Williams) A. Gentry, Taxon 22: 640. 1973.

Shrub or small tree to 6 m tall; branchlets angulate when young, terete

with age. Leaves alternate, narrowly obovate, acute to short acuminate at
apex, cuneate-attenuate at base, 7-15 cm long, 2.5-4.5(-6.4) cm wide, subcoria-
ceous, very inconspicuously scattered lepidote, otherwise glabrous, midrib
slightly raised above, strongly raised beneath, secondary veins plane above,
very prominent below, not white-edged, (14-)15-18 on a side separated by 0.5-1
Amphitecna 73

Atat,/ tS /'S D

FIG. 14. Amphitecna tuxtlensis, A. molinae and A. sessilifolia. A, A. tuxtlensis (Beaman 5206);
B, A. molinae (Molina 24295); C, A. sessilifolia (Croat 15712). (All x 1/2).
74 Flora Neotropica

cm, drying olive to gray both above and below, the margin entire and hyaline;
petiole slender, not well demarcated from leaf base, about 1 cm long.
Inflorescence one or two terminal flowers, the pedicels 2.5-3 cm long, glab-
rous. Flowers with the calyx thick, campanulate, 2.2-2.5 cm long, ca. 2 cm
wide, glabrous; corolla white, tubular-infundibuliform, the throat slightly
ventricose, 3.5-4.5 cm long, 1-2 cm wide at mouth of tube, the lobes fused into
a frilly-margined rim, stalked glandular on lobes, otherwise glabrous; stamens
and ovary not examined. Fruit ellipsoid, tapering gradually to both ends, 12
cm long, 6 cm in diameter; seeds not seen.
Type. Honduras. Intibuca: km. 9-11 La Esperanza-Marcala road, 1600
m, Molina & Molina 24295 (holotype, F; isotypes, EAP, NY).
Distribution. Known only from the type locality in open lower montane
forest at 1600 m in southern Honduras.
HONDURAS. Intibuca: km. 9-11 La Esperanza-Marcala road, 1600 m, Molina & Molina
24299 (F).

The flowers of A. molinae are not very different from those of closely re-
lated A. breedlovei. It differs from that species chiefly in the more prominent
and closer together secondary veins of its leaves and its more eastern and
higher altitude distribution. The single known fruit of A. molinae is signifi-
cantly longer and has more tapering ends than the fruits of A. breedlovei.

11. Amphitecna montana L. O. Williams, Fieldiana, Bot. 36: 22. 1973.

Figs. 6, 7, 8.
Dendrosicus montanus (L. O. Williams) A. Gentry, Taxon 22: 640. 1973.

Tree 10-20 m tall; branchlets conspicuously angulate, becoming subterete

with age. Leaves alternate, narrowly obovate, acute to short-acuminate at
apex, cuneate to more or less attenuate at base, (8-)10-30(-34) cm long, (2.4-)3-
9(-11) cm wide, subcoriaceous to chartaceous, very inconspicuously and mi-
nutely scattered lepidote, otherwise glabrous, midrib slightly raised above,
prominently raised below, secondary veins plane or prominulous above, prom-
inent below, sometimes more or less white edged, drying olive to blackish, the
main veins either darker or lighter; petiole 1-4 cm long, merging with base of
leaf blade. Inflorescence a single pendent flower, axillary or on small
branches, the pedicel 6-10 cm long, glabrous. Flowers with the calyx thick,
campanulate, 3-4 cm long, 1.7-3 cm wide, glabrous or with a few lepidote
scales near base, irregularly bilabiately or subspathaceously rupturing in upper
third or half, circumscissilely separating from pedicel; corolla white to pale
greenish-yellow, tubular-infundibuliform, more or less radially symmetrical,
(3.5-)4-7 cm long, 1.5-2.5 cm wide at mouth of tube, the lobes fused into an in-
conspicuous frilly-margined rim, stalked-glandular on lobes, otherwise glab-
rous; stamens inserted 1.2-3 cm from base of tube, the anthers subexserted, the
thecae slightly divergent, 6-8 mm long; ovary ovoid, densely minutely lepidote,
3 mm long, 2 mm wide at base, mostly 2-locular with axile placentation, at
extreme tip unilocular with parietal placentation; disc patelliform-pulvinate, 2
mm long, 5 mm wide. Fruit oblong-ovoid or ellipsoid, very inconspicuously
Amphitecna 75

short-apiculate,smooth surfaced,9-15 cm long, 6-8 cm in diam., the pedicel

ca. 8 cm long; seedsthick, 12 mm long, 18mm wide.
Type. Guatemala.San Marcos:Volcin Tajumulco,2300-2800m, Steyer-
Distribution.SouthernChiapasand adjacentuplandsof westernGuate-
mala; apparentlydisjuncton Montafiade Celaque, Honduras;montane for-
ests from (1500-)1900-2600m.
MEXICO.Chiapas:45-50km NE of Huixtla,1900m, Breedlove& Smith22620(DS), Breed-
love & Thorne31011(DS);CerroMozotal,2100m, Breedlove25718(MO, DS), 40461(MO);Mt.
Ovando,nearEscuintla,Matuda1818(MICH);HaciendaSiltepec,Matuda1947(A, F, K, MICH,
MO, NY); Mt. Ovando, 1900m, Matuda4206 (A, F, GH, NY, MICH, US); Letrero,Matuda
17837(F);betweenCanadaHonda& El Triunfo,1300-2100m, Xoloctzi&SharpX334(MEXU).
GUATEMALA.Huehuetenango:17mi N of Barillas,1500m, Steyermark48766(A, F). San
Marcos:betweenTodos Santosand FincaEl Porvenir,1300-3000m, Steyermark36987(F), 37012
(F); San Marcos-SanRafael road, 2500-2700m, Standley86200 (F, US); 8-10 km W of San
Marcos,2300m, Williamset al. 26915(F, GH, NY, US).
HONDURAS.Lempira:Montafiade Celaque,1200-1500m, Hazlett2287(MO).

Localname. Honduras:Morro.

This speciesis very well characterizedby the ratherlarge distinctlypetio-

late leaves, very long pedicels, and large pendentalmost radiallysymmetric
flowerswith a largecircumscissilelycaducouscalyx. Its relativelybroaderand
moreoblong non-apiculatefruit is uniqueamongthe speciesof northernCen-
tral Americabut similarto fruits of A. kennedyiand A. sessilifolia;both of
these specieshave shorterpedicels.The calyx of A. montanasplits irregularly
and incompletelyand may be both irregularlybilabiateand subspathaceouson
the same plant. Both known calyces of the single Hondurancollection are
spathaceousand the corolla is shorter and the anthers more conspicuously
exsertedthan in other specimens.If all these charactersprove constant, the
disjunctHonduranpopulationmightmerittaxonomicrecognition.The single
Huehuetenangocollection-Steyermark 48766 from 17 miles N of Barillasin
the Sierrade los Cuchumatanes-is representedonly by sterile materialand
immaturefruits. The fruits are describedas 6 inches long and 3 inches in
greatestdiameter,dimensionsthat wouldfit A. montana.
12. AmphitecnaparvifloraA. Gentry,sp. nov. Figs. 6, 8.
Frutexscandens.Folia angusteobovata,acuminata,cuneata.Florescauliflori,calyce2(-3)-
1.8-2.4cm longa, basicomparatelata, disco 5 mm
labiato,corollaalba, late tubulo-campanulata,
lato. Fructusignotus.

More or less scandentshrub;branchletsangulate,the surface wrinkled-

striate, tannish. Leaves alternate,clusterednear tips of branchlets,narrowly
obovate,acuminate,cuneateat base, (6-)12-20cm long, 2-5.5 cm wide, charta-
ceous, glabrous, midvein raised above and below, secondary veins plane
above, prominulousbelow, tertiaryveins plane above and below, not white-
margined, drying blackish-oliveabove and below, the main veins darker
below; petiole usually poorly differentiated,to 1 cm long. Flowerscauliflor-
ous, bornesinglyon bracteateshort-shoots,the slenderpedicels2-2.5 cm long,
76 FloraNeotropica

the calyxirregularly2(-3)-labiate,split to nearbase, 1.4-1.9cm long, the lobes

obtuse to inconspicuouslyapiculate, sparsely minutely lepidote, otherwise
glabrous,tardilycaducous;corollawhite, broadlytubular-campanulate with a
broad base, somewhatexpandedabove and below middle, 1.8-2.4 cm long,
1.7-2cm wideat mouth, the lobes completelyfusedinto an inconspicuousrim;
stamensincluded,the antherthecae divergent,very broad, 2.5-3 mm long, 2
mm wide; ovary conical, 3 mm long, 1.3 mm wide at base, denselyminutely
lepidote;disc annular-pulvinate,1.5 mm long, 5 mm wide. Fruitunknown.
Type. Panama. Chiriqui: La Fortuna hydroelectricproject; in cloud
foreston ridgebehindcamp, 1300-1400m, scandentshrub,cauliflorous,flow-
ers white, 23 March1978,B. Hammel2236 (holotype,MO 2627723;isotypes,
MO 2627724,PMA).
The most distinctivefeature of A. parviflora is its very small corolla,
smallerthanthat of any otherspeciesof AmphitecnaexceptA. donnell-smithii
of Guatemala.Amphitecnaparvifloraprobablyis most closely relatedto A.
donnell-smithiifrom which it differs in the more subcoriaceousleaves, non-
prominulous tertiary leaf venation, cauliflorous flowers, different-shaped
broader-basedcorolla, and muchbroaderdisc. Among Panamanianspeciesof
Amphitecna,A. parviflorais closest to A. isthmicawhich occurs at a lower
altitude and differs in terminal inflorescences,much larger corollas, sub-
exserted anthers, and the main veins drying whitish or yellowish beneath.
Amphitecnasessilifoliausuallyoccursat higheraltitudesand differs in much
largercorollas,subexsertedstamens,longerantherthecae, largerovary, wider
disc, mostly terminalflowers, and especiallya largermuch thickercalyx. The
scandenthabitof A. parviflora,if correctlyreportedand constant,is uniquein
the genusand in tribeCrescentieae.

13. Amphitecnaregalis(Linden)A. Gentry,comb. nov. Figs. 8, 9, 10.

CrescentiaregalisLinden,PI. Nouv. 7. 1859.

Small pachycaultree 4-12 m tall, to 15 cm dbh, the leaves in a terminal

cluster, branchletsthick, rather angular. Leaves alternate-verticillate,con-
gestednearbranchapex, oblanceolate,obtuseto acutish,cuneate-attenuateat
base, gigantic,69-100cm long, 18-35cm wide, subcoriaceous,glabrous,mid-
ribconspicuouslyraisedabove and below, secondaryveinsplaneabove, barely
prominulous below, whitish-margined,drying olive below, blackish-olive
above, petiolenot differentiated.Inflorescencecauliflorous,the pedicelsglab-
rous, ca. 1 cm long. Flowerswith the calyx bilabiate, circumscissilelycadu-
cous, bilabiatelysplit to base, 2.5-2.8 cm long, each lobe ca. 1 cm wide, ob-
tuse, glabrous;corolla white, tubular-infundibuliform, more or less radially
symmetrical, 5.2-6.5 cm long, 2.3-3 cm wide at mouth of tube, the lobes fused
into a frilly-marginedrim, sparselystalked-lepidoteon outsideof lobes, other-
wise glabrous;stamens inserted2-2.5 cm from base of tube, anther thecae
slightlydivergent,6-7 mm long, the filaments1.5-1.7 cm long; ovaryconical,
ca. 3 mm long, to 2 mm wide at base, denselyminutelylepidote;disc annular-
pulvinate,1 mm long, 5 mm wide. Fruitunknown.
Amphitecna 77

Type. Mexico. Chiapas: Ghiesbrecht s.n. (BM, perhaps from cultivated

Distribution. Evergreen tropical forest below 200 m altitude from central
Veracruz to extreme northwestern Chiapas.
MEXICO. Chiapas: versunt septentrionale de la cordillera, Ghiesbrecht s.n. (BM). Oaxaca:
Ubero, 30-90 m, LI. Williams 9443 (F). Tabasco: Chiapas border, km. 59 carreterra Huiman-
guillo-Mal Paso (cuad. 50A), Gonzalez et al. s.n. (MEXU). Veracruz: La Laguna, Hidalgotitlan,
200 m, Brigada Vazquez 105 (F).
CULTIVATED EUROPEAN: without collector, 17 Aug 1878 and Jul 1887, cultivated in
Munich Botanical Garden (M).

Local name. Morro-cimmaron.

Vegetatively the most obvious difference between A. regalis and its two
pachycaul relatives A. macrophylla and A. megalophylla is the barely
prominulous secondary venation of the leaf undersurface; its gigantic leaves
are also apparently larger than in either of its two closest relatives. The corolla
is broader and more infundibuliform and the calyx lobes shorter than in the
similar flower of A. macrophylla; the small flower of A. megalophylla is com-
pletely different. Ecologically A. regalis is separated from the two related
species by its lowland habitat.
I have previously placed C. regalis in the synonymy of A. macrophylla
(Gentry, 1973b). To my knowledge, C. regalis otherwise has been universally
ignored except for its listing in Index Kewensis. Linden's obscure "sub-
nudum" description of sterile material-"Arbuste majestueux a feuilles
sessiles entieres, dures et coriaces, spatulees, longement attenuees, mesurant 3
pieds de longueur sur 10 pouces de largeur. C'est une plante a effet grandiose,
et la digne rivale du Curatella (Theophrasta) imperialis. "-seems hardly ade-
quate for identification. Nevertheless the very large leaves described for C.
regalis fit only this species and Guatemalan A. megalophylla. There are two
sterile collections of this species at the British Museum each consisting of a
single leaf, identified as Crescentia regalis Linden and labelled as originating in
Chiapas. One leaf is 82 cm long and 19 cm wide, the other 63 cm long and 16
cm wide. The larger leaf was collected by Ghiesbrecht from whom Linden ob-
tained much Mexican material, and undoubtedly represents the plant brought
into cultivation by Linden. I have designated it as the type of Crescentiaregalis.
The existence of 1878 and 1887 flowering collections labelled as C. regalis ob-
tained from a cultivated plant at the Munich Botanical Garden and pre-
sumably deriving from Linden's original distribution, further prove that
Linden's plant is indeed this rare Mexican species.

14. Amphitecna sessilifolia (Donnell Smith) L. O. Williams, Fieldiana, Bot.

36:25. 1973. Figs. 11, 14.
Tabebuia sessilifolia Donnell Smith, Bot. Gaz. (Crawfordsville) 25: 156. 1898.
Neotuerckheimia gonoclada Donnell Smith, Bot. Gaz. (Crawfordsville) 47: 259. 1909. Type.
Tonduz 7384, Costa Rica, San Jos6, fl (holotype, US; isotypes, BM, GH, K, M, MO,
78 Flora Neotropica

Enallagma sessilifolia (Donnell Smith) Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 18:
1120. 1938.
Dendrosicus sessilifolius (Donnell Smith) A. Gentry, Taxon 22: 640. 1973.

Shrub or tree to 18 m tall and 25 cm dbh.; twigs subterete, glabrate, usu-

ally moss-covered. Leaves simple, alternate, oblanceolate to narrowly obo-
vate, acute, attenuate with the petiole undifferentiated and essentially lacking,
13-29 cm long, 4.0-8.5 cm wide, chartaceous, secondary veins 11-16 on a side,
the midvein raised above, lepidote above and beneath, a few plate-shaped
glands at base beneath, drying dark brown. Inflorescence a terminal fascicle,
the pedicels glabrous to inconspicuously lepidote, 2.2-7.9 cm long, commonly
reduced to a single flower. Flower with the calyx thick, fleshy, irregularly split
into subequal anterior and posterior halves or merely ruptured apically by the
expanding corolla, one of the lobes sometimes itself split, 2.5-2.8 cm long, 1.9-
2.1 cm wide, glabrous; corolla greenish-white, tubular, usually with an incon-
spicuous transverse fold in the throat, fleshy, 4.1-5.2 cm long, 1.2-1.6 cm wide
at the mouth, the tube 2.5-3.3 cm long, the lobes fused, 0.5-1.2 cm long, glab-
rous; stamens subexserted, inserted 13-18 mm from base of the tube, the
anther thecae thick, somewhat divergent, 5 mm long, 2-3 mm wide, the anterior
filaments 2.3-2.4 cm long, the posterior filaments 2.5-2.7 cm long; pistil 3.8-3.9
cm long, the ovary rounded-conical, 8-9 mm long, 4-5 mm wide, incompletely 2-
locular with 2 protrudingparietal placentae at the top, these fused above middle,
the ovules then irregularly4-seriatein each locule; disc pulvinate, ca. 3 mm long,
9-10 mm wide. Fruit ellipsoidalwith the apex elongated, 12-15cm long, 6.6-7.2 cm
in diam., evenly rough-surfacedunder a lens, inconspicuously scattered-lepidote;
seeds thick, 1.5-2 cm long, 1.5-2.4 cm wide, without wings.
Type. Costa Rica. San Jose: Tablazo, 1800 m, Tonduz 7937 (holotype,
US; isotypes, BR, CR).
Distribution. Costa Rica and western Panama in lower montane rain for-
est between 1300 and 2000 m alt.
COSTA RICA. Prov. unknown: Paso Ancho, Lankester 295 (K). Alajuela: Alto de La
Palma, 1300 m, Brenes 3830 (F); La Palma de San Ram6n, Brenes 168 (CR); Palmira, 5700 ft, A.
Smith A 134 (MO), 1600 m, 576 (A, NY); Zarcero, A. Smith 1570 (A, NY); La Ventolero, 1700 m,
Standley 34733 (US); Palmira, 7200 ft, Stork 4135 (MICH). Cartago: Tapanti, Gentry 1001 (MO),
2044 (CR, MO); 8 km S of Tapanti, 1550 m, Lent 1231 (CR, F); Paraiso, 1600 m, Madriz AMV-37
(CR, UCR). Heredia: Volcan Barba, 1750 m, Wilbur & Stone 9867 (GH, NY). San Jose: El Tab-
lazo, 1700 m, J. Le6n 808 (CR), Tonduz 7384 (BM, CR, GH, K, M, MO, NY, US); La Palma,
1500 m, Werckle 17419 (GH).
PANAMA. Chiriqui: New Switzerland, 1800-2000 m, Allen 1362 (GH, MICH, MO, NY,
US); Finca Collins, Blum & Dwyer 2570 (MO, SCZ); Bajo Chorro, 6000 ft, Davidson 410 (MO);
Cerro Horqueta, 5000-5800 ft, Dwyer et al. 444 (GH, MO, US); Dwyer & Hayden 7694 (K, MO);
Nueva Suissa, Gentry 5982 (MO); 2 mi E of El Hato del Volcin, King 5305 (K, US); El Boquete,
1000-1300 m, Pittier3001 (NY, US); near El Volcan, White227(GH, MO, US).

Local names. Costa Rica: Jicarilla, Guacalillo, Jicaro, Calabacero.

Panama: Calabash.

15. Amphitecna silvicola L. O. Williams, Fieldiana, Bot. 36: 25. 1973.

Figs. 6, 7, 11.
Amphitecna 79

Dendrosicussilvicola(L. O. Williams)A. Gentry,Taxon22:640. 1973.

Shrub or small tree 2-12 m tall; branchlets angled when young, terete with
age. Leaves alternate, narrowly obovate, acute to short-acuminate at apex,
cuneate to attenuate at base, 5-20 cm long, 1.5-5 cm wide, subcoriaceous to
coriaceous, glabrous, midrib slightly raised above and below, other venation
plane above and plane or slightly impressed below, the venation below intri-
cately reticulate and conspicuously whitish-margined, drying blackish or dark
olive above, light olive below; petiole poorly demarcated, to 1 cm long. Inflor-
escence a single terminal or axillary flower or a fascicle of 1 or 2 flowers from
the smaller branches, the pedicel 1.5-3(-4) cm long, glabrous. Flowers with the
calyx campanulate, irregularly bilabiate or trilabiate, split to below middle,
1.4-2.5 cm long, the lobes obtuse to acute, glabrous or with a few lepidote
scales near base, usually a few lepidote glands near apex; corolla white,
tubular-infundibuliform, only slightly bilabiate, 2.5-5 cm long, 1.5-2.5 cm
wide at mouth of tube, the lobes fused into a frilly-margined rim, conspicu-
ously lepidote and stalked-glandular on lobes, otherwise glabrous; anthers
subexserted, the thecae somewhat divergent, 3-4 mm long; pistil 3.5 cm long,
the ovary oblong-conical, 3-5 mm long, 1.5-2 mm wide, slightly lepidote; disc
truncate-conical, 1.5-2 mm long, 4-5 mm wide. Fruit ellipsoid to narrowly
oblong-ellipsoid, 8-14 cm long, 4-5.5 cm wide, gradually tapering to an acumi-
nate or subtruncate apex, tapering basally but more or less truncate at extreme
base; seeds thick, 1.3-1.5 cm long, 1.5-1.7 cm wide.
Type. Mexico. Chiapas: 30 mi E of La Trinitaria, 1500 ft, Breedlove
10064 (holotype, F; isotypes, MEXU,MO).
Distribution. Middle elevation forests between 900 and 1500 m on north
slope of the northern cordilleras from eastern Chiapas to Alta Verapaz,
MEXICO.Chiapas:E of LagunaTzikaw, 1300 m, Breedlove29655 (DS, MICH), 35106
(DS, MO); 4 km E of LagunaTsiskaw, 1300m, Breedlove38800 (DS, MO); 42 km NE of La
Trinitaria,1300m, Breedlove& Thorne21175(DS, MO).
GUATEMALA.Alta Verapaz:H. Johnson867 (US);43 kmNE of Cobin, 1200m, Standley
70148(F, US); regionof Cocola, 1200m, Standley70325(F, US); Rio Coban, 5 km SE of Tactic,
1300-1500m, Williamset al. 40695(F, NY); vic. of San Juan Chamelco,1500m, Wilson40836

Local name. Guatemala: (Quecchi) S-xom-qawa'?ul (= our Lord Moun-

tain's Gourd).

Vegetatively A. silvicola is easily distinguished by the small leaves with a

characteristic shagreened appearance beneath from the intricately reticulate
white-edged impressed venation. The material from the Coban area of Alta
Verapaz, Guatemala, has the leaf undersurfaces with prominent or sub-promi-
nent secondary veins and less shagreened; the pedicels are longer and more
slender (3.8-4 cm vs. 1.5-3 in Chiapas) and the fruits smaller (8 cm vs. 14 cm
long). Presumably these are intraspecific variations, especially in view of the
limited sample size.
80 FloraNeotropica

16. Amphitecnaspathicalyx(A. Gentry)A. Gentry,Taxon25: 108. 1976.

Figs. 11, 12.
Dendrosicus spathicalyx A. Gentry, Phytologia 26: 439. 1973.

Small,leaningtree 5-7 m tall, 15 cm dbh., the barkthicklymoss-encrusted;

the twigs decurrentbelow the nodes. Leavesalternate,narrowlyobovate-ellip-
tic, acute to long acuminate,basallycuneateor attenuate,essentiallysessile,
the extremebase swollen and woody, conspicuouslyjointed at nodes, 7.5-16
cm long, 2.1-4.3 cm wide, chartaceous,the secondarynerves6-12 on a side,
slightlyraisedabove, conspicuouslyso beneath,the veinletsminutelyand in-
tricatelyimpressedabove and beneath, the surfacethus roughenedand sha-
greened.Inflorescencea single flower borne from a bracteateshort-shooton
an older twig, the pedicel glabrous, red when fresh. Flowers with the calyx
membranaceous,4-4.1 cm long and ca. 2 cm wide, spathaceouslysplit adaxi-
ally to 0.8-1 cm from the base with a tendencyto tardilycircumscissiledehis-
cence, glabrous,light greenwhen fresh;corolla creamto whitishbasally, be-
cominggreenish-whitetowardthe mouth, campanulatewith a transversefold
in the throat, 5.5-5.8 cm long, 1.8-1.9 cm wide at the mouth, the tube 3.7-3.8
cm long, the lobes fused into a reflexedrim, 1.7 cm wide, glabrouswith a
glandularepidermisinside and out, becomingstalkedglandular-lepidotedis-
tally outside;stamenssubexserted,inserted1.7-2.1 cm from base of the tube,
the antherthecaeslightlydivergent,7-8 mm long and 1.5-2 mm wide, the fila-
ments 2.3-2.6 cm long, the staminode1.1-1.2 cm long, inserted 1.0 cm from
the base of the tube, bent straightout from the corolla; pistil (partiallyde-
stroyedin all flowers examined)with the ovary roundedconical, 4 mm long
and 3-4 mm wide, lepidote, the ovules widely spaced, irregularly8-seriateon
each of 2 projectingparietalplacentae,the placentaealmostmeetingbut fused
only at the extremebase;disc annular-pulvinate, 2 mm long, 9 mm wide. Fruit
Type. Panama. Panama: Cerro Campana, Gentry 5769, fl (holotype,
MO;isotype, SCZ).
Distribution.Endemicto the top of CerroCampanain centralPanama,
at about900 m alt.; tropicalwet forest.
PANAMA. PanamA:Cerro Campana, Croat 14765 (MO, SCZ); Dressier 3943 (MO).

The unique,thin, dorsallysplit, apiculate,spathaceouscalyxof A. spathi-

calyx is distinctive.Except for the calyx this speciesappearsrathersimilarto
A. silvicola. The only other specieswith an apiculatespathaceouscalyx is A.
steyermarkiiwhichhas very differentoblanceolateleaves and a narrowergib-

17. Amphitecnasteyermarkii(A. Gentry)A. Gentry, Taxon 25: 108. 1976.

Figs. 6, 9, 10, 11.
Dendrosicus steyermarkii A. Gentry, Phytologia 26: 445. 1973.
Amphitecna 81

Tree8-10m tall. Leavessessile,alternate,glabrous,oblanceolate,acuteto

shortly acuminate, the base cuneate-attenuate,11-25 cm long, 2.8-6.5 cm
wide, subcoriaceous,the main nervesraisedbelow, dryinggray above, brown-
ish-graywithtan mainveinsbelow. Inflorescenceone to severalterminalflow-
ers, the pedicel4-5 cm long. Calyxspathaceouslysplit to base, apiculateat tip,
3.2 cm long, glabrous;corollaca. 4.5 cm long, 1.4 cm wide at mouth of tube,
the tube 2.8 cm long, the lobes ? fused into a reflexedrim, tube mostlyglab-
rous, slightlylepidote-papillateat top and on lobes; stamenssubexserted,the
antherthecaedivergent,each 4 mm long, filamentsca. 2.5 cm long, the stami-
node ca. 1 cm long, insertion 1-1.5 cm from base of tube; pistil ca. 3.5 cm
long, ovary oblong-conical, 3 mm long, 2 mm wide, minutely lepidote-
papillate, bilocular in lower half with ovules irregularly2-3-seriate on 2
axillaryplacentaein each locule, unilocularwith ovules severalseriateon each
of 2 protrudingparietalplacentaeat top; disk pulvinate,1.5 mm long, 3-4 mm
wide. Fruitoblong-ellipsoid,10-11cm long, 4-5 cm wide, widestnearmiddle,
the base gibbous, rathersharp-angled,taperingto a blunt apex; seeds strongly
2-lobed,thick, wingless,12-14mm long, 1.5-1.8mm wide.
Type. Guatemala.Huehuetenango:betweenDemocraciaand canyon of
Chamushu,1000-1100m, Steyermark51252(holotype,F; isotype, NY; photo-
copy, MO).
Distribution.Oak forests between 1000 and 1200 m altitude in Chiapas
and westernGuatemala.
MEXICO.Chiapas:6-8 km E of FronteraComalapa,1000m, Breedlove39067 (DS); Cor-
cega, PuebloNuevo,Matuda17649(F).

Localname. Guatemala:Morro.

Amphitecnasteyermarkiiis remarkablein the genus in its spathaceously

split Parmentiera-likecalyx with a distinctlyapiculatetip. The only other spe-
cies of Amphitecnawith a similarcalyx is PanamanianA. spathicalyxwhich
has verydifferentintricatelyimpressed-reticulate moreor less ellipticleaves.
Identificationof the fruitingcollection from western Chiapas (Matuda
17649) with this species is somewhat tentative. The leaves (probably new
growth)are membranaceousbut sharethe characteristicoblanceolateshapeof
this species. If this peculiaroblongoid fruit with its gibbous almost square-
corneredbase is not A. steyermarkiithen it clearlyrepresentsan undescribed
species.The fruitingmaterialpreviouslydoubtfullyreferredto A. steyermarkii
(Gentry,1977b)apparentlybelongsto A. apiculata.The strikinglybullatefruit
surfaceof thesecollectionsprovesof no taxonomicvalue, as I have now seen a
singlegatheringwith fruitsrangingfrom quite smooth to bullate;the unusual
surfacetextureis a resultof abnormalgrowthas I previouslysuggested.
18. Amphitecna tuxtlensis A. Gentry, sp. nov. Figs. 11, 14.
Frutex5-15m altus. Folia oblanceolatavel perangusteelliptica,acuminata.Floresramiflori
82 FloraNeotropica

(vel cauliflori?),pedicello1.5-2cm longo, calycecampanulato,2-3-labiato,caduco,corollaalba,

tubulo-infudibuliformi, basitubulari.Fructusignotus.

Tree 5-15 m tall; branchletsangulatewhen young, becomingterete with

age. Leaves alternate, oblanceolate to very narrowly elliptic, acuminate,
graduallytaperingto an attenuate-cuneatebase, 8-18 cm long, 1.5-5 cm wide,
chartaceous,inconspicuouslyscattered-lepidote,otherwiseglabrous, midrib
very slightlyraisedabove, prominentbelow, the secondaryveins planeabove,
prominentbelow, not prominentlywhite-edged,dryingdarkgrayabove, olive
withtan mainveins below;petioleca. 1-2cm long, mergingwith attenuateleaf
base. Inflorescencea singleflowerborneon a smallbranchletbelow the leaves
(or cauliflorous?),the pedicel 1.5-2 cm long. Flowers with the calyx rather
thinly coriaceous, campanulate, 1.6-2.5 cm long, 1-1.8 cm wide, glabrous
exceptfor a few scatteredlepidotescalesnearbase or apex, evenly2-3-labiate-
ly splitto below middle,circumscissilely separatingfrompedicel;corollawhite,
tubular-infundibuliform above a 2-2.5 cm long pronouncedbasal tube, more
or less radiallysymmetrical,4-5.5 cm long, 2-3 cm wide at mouth of tube, the
lobes subacute,more or less fused into a marginalrim, conspicuouslylepidote
at top of tube outside, sparselystalked-lepidoteon lobes, otherwiseglabrous;
antherssubexserted,the thecaedivergent,5 mm long; ovaryand disc not seen.
Fruitnot seen.
Type. Mexico. Veracruz:lado NE de Lago Catemacoen Cerrosal E de
Cogame,450 m, Mapa21.5-57.0;selvaalta perennifolia,vegetaci6nprimaria,
arbol de 15 m, flor blanca, 29 Oct 1971,J. Beaman5206 (holotype, MO; iso-
type, F, MEXU, MSC, XAL).
Distribution.Endemic to evergreenrainforestat intermediatealtitudes
(ca. 450 m) in the Sierrade los Tuxtlas,Veracruz,Mexico.
MEXICO.Veracruz:Estaci6nBiologicade los Tuxtlas,Mapa 24.0, 56.5, vegetaci6npri-
maria,suelo arcilloso,escasaarbol 10 m, frutoverde,JuanI. Calzada529 (MEXU);El Bastonal
Station,Martha,18?23' N, 94? 57' W, selvaalta, arbolabundante,5 m, plantacauliflora,flor

The small acuminaterather thin leaves are reminiscentof A. donnell-

smithiibut the floweris much largerthan in that species.The corolla shapeis
very like that of A. montanaand the circumscissilecalyx also agreeswith A.
montanawhichis no doubt the closest relativeof A. tuxtlensis.The new spe-
cies differsfrom A. montanaespeciallyin its shorterpediceland smaller,thin-
ner, evenly splittingcalyx as well as in significantlysmallermostly oblanceo-
late leavesand relativelylowlandhabitat.The isolatedSierrade los Tuxtlasis
over400 km NW of the westernmoststationfor A. montanain Chiapas.
One collection describes A. tuxtlensis as a small cauliflorous tree while the
type, from a largertree, includesattachedflowerson small branchlets;I sus-
pect that the "cauliflorous"collectionwas also ramiflorous,especiallyin view
of the smallsize of the tree fromwhichit was taken.


2. Crescentia Linnaeus, Sp. P1. 2: 626. 1753; Gen. PI., ed. 5. 274. 1754.
Crescentia 83

PteromischusPich6n, Bull. Soc. Bot. France92: 227. 1946. Type: P. alatus (Humboldt,
Bonpland& Kunth)Pich6n = CrescentiaalataHumboldt,Bonpland& Kunth.

Small to medium-sized trees, branching rachitic, crown open. Leaves sim-

ple or 3-foliolate, borne on thick twigs in alternate fascicles resulting from the
condensation of short-shoots. Inflorescence of 1-2 cauliflorous flowers arising
from nodes on the trunk and older branches. Flowers with the calyx large,
usually bilabiately split; corolla off-tan, usually with maroon penciling, espe-
cially on lobes and on the tube inside, the lobes deltoid, acuminate; stamens
subexserted, the anthers glabrous, the thecae thick, somewhat divergent; pol-
len grains single, 3-colpate or 6-colpate with the three primary colpi transverse-
ly divided at middle, the exine microreticulate; ovary ovoid-elliptic, lepidote,
1-locular, the ovules multi-seriate on 4 parietal placentae. Fruit a pepo or cala-
bash, large, more or less spherical, indehiscent with a hard woody shell, pulpy
inside; seeds small, less than 8 mm long and 9 mm wide, flat, not winged,
embedded in the pulp.
Type species. C. cujete L.
Six species in tropical America ranging from Mexico and the West Indies
to Amazonian Brazil, also cultivated through most of the tropics.

Key to Species of Crescentia

1. Leaves3-foliolate (rarely5-foliolatein part)and simplein each fascicle;Mexicoto Costa
Rica,rarelycultivated. 1. C. alata.
1. Leavesall simple;widespread,especiallyin cultivation.
2. Leavesoblanceolateto obovate,largerleavesmorethan4 cm long and 1.5 cm wide;fruit
morethan5.5 cm long;widespread,especiallyin cultivation.
3. Fruitoblong witha truncatebase;leavesnoticeablyshiny;corollalobes obtuse;south-
westPuertoRico. 6. C.portoricensis.
3. Fruitgloboseor ellipsoid;leavesdull;corollalobestriangularand moreor less acumi-
nate;widespread,especiallyin cultivation.
4. UpperOrinocoandupperAmazon;fruitlessthan8 cm long. 2. C. amazonica.
4. Caribbeancoast of Mexico and northernCentralAmerica,also widelycultivated
andnaturalized;fruitmorethan8 cm long. 3. C. cujete.
2. Leaveslinearor microphyllous,largestleavesless than 1.2(-2.5)cm wide or less than 2.5
cm long; fruitless than 5 cm long; drycoastalthicketsof Greaterand LesserAntillesand
5. Leaveslinear;PuertoRico, Hispaniola,LesserAntilles,andBelize. 4. C. linearifolia.
5. Leavesmicrophyllous,obovate;Cuba. 5. C. mirabilis.

Species of Crescentia are defined entirely by vegetative characters and dif-

ferences in fruit size. Even the most distantly related species (e.g. C. alata and
C. cujete) are interfertile. Nevertheless these plants tend to form dense aggre-
gations of homogeneous morphotypes where they occur, and taxonomic
recognition of different-appearing ecologically differentiated dominants seems
in order despite the tenuous nature of their reproductive barriers and dis-
tinguishing characters.

1. Crescentia alata Humboldt, Bonpland & Kunth, Nov. Gen. Sp. PI. 3: 158.
1819. Figs. 15, 16.
CrescentiatrifoliaBlanco,Fl. Filip., ed. 1. 489. 1837.Type:not seen.
84 FloraNeotropica

Otophora?paradoxa Blume, Rumphia 3: 146. 1847. Type: not seen.

Parmentiera alata (Humboldt, Bonpland & Kunth) Miers, Trans. Linn. Soc. London 26:
166. 1868.
C. ternata Sesse & Mociflo, La Naturaleza, ser. 2, 1 (append.): 94. 1889. Type: Mexico, Sesse
& Mociflo 2227 (F).
Pteromischus alatus (Humboldt, Bonpland & Kunth) Pich6n, Bull. Soc. Bot. France 92: 227.

Small tree to 25 cm dbh and 8 m tall, the branchescrooked, the crown

open; branchletsessentiallylacking, the smaller branchesthick, subterete,
with alternatecondensed, short-shootprojectionseach bearinga fascicle of
leavesfrom the center. Leavesboth compoundand simplefrom each fascicle,
the compoundleaves3-foliolate(rarely5-foliolatein part)with sessileoblance-
olate leaflets 1-4.5 cm long and 0.3-1.2 cm wide, round-tippedor slightly
emarginate,lepidote but otherwise glabrous, rigid-chartaceous,the petiole
winged,2.7-11.5 cm long, 3-15 mm wide, lepidote,oblanceolate,rigid-charta-
ceous as the leaflets, the simpleleavesmuchsmaller,1.9-3.2cm long and 0.5-1
cm wide, similarto single leaflets or petioles of compoundleaves. Inflores-
cenceone or 2 cauliflorousflowersborneon largerbranchesor trunk,pedicels
lepidote,0.5-0.8 cm long. Flowerswith a muskyodor, calyx split nearlyto the
base into two subequal lobes, each 1.4-1.9 cm long and 0.7-1.4 cm wide,
mostly glabrous, slightly lepidote at base, usually with sunken plate-shaped
glands;corolla off-tannish, tubular-campanulate with a horizontalpleat mid-
way acrossthe lowerside of throat, fleshy, 4.0-6.5 cm long, 1.8-2.5cm wide at
mouthof tube, the tube 2.8-4.2 cm long, the lobes 1-1.5cm long, more or less
triangularand fused at the bases, mostlyglabrous,sparselyglandular-lepidote
outside;stamenssubexserted,the antherthecaeca. 2 mm thick, only partially
divergent,5-7 mm long, the anteriorfilaments2.1-3.4 cm long, inserted0.8-
1.7 cm from the base of the tube, the posteriorfilaments 2.3-3.8 cm long,
inserted 1.1-1.5 cm from the base of tube; pistil 4.3-5.8 cm long, the ovary
rounded-conical,lepidote, 5-6 mm long, 4-5 mm wide, the ovules multi-seriate
on 4 parietalplacentae;disc pulvinate,2-3 mm long, 0.9-1.4 mm wide. Fruita
pepo or calabash,more or less spherical,7-10 cm in diam.; seeds small, thin,
wingless,6-7 mm long, 7-9 mm wide, scatteredthroughthe pulp of the fruit.
Type. Mexico. Guerrero:Acapulco,Humboldt&Bonpland3858 (P).
Distribution.Mexico to Costa Rica, mostly along the Pacific Coast. A
characteristicand often dominantelementof the drysavannas.
MEXICO. Sin. loc.: Linden 228 (MICH); Sesse & Mocifo 2227 (F); Stange s.n. of 1962
(MICH); Laguna Verde, Liebmann 11349 (C). Baja California: San Jose del Cabo, Jones 24011
(A, MO). Chiapas: 20 mi S of La Trinitaria, 2800 ft, Breedlove 11788 (DS, F, MICH, US); 3-5 mi
N of Cintalapa, 900 m, Breedlove & Thorne 30469 (DS, MO); Arista, Matuda 17365 (F, K,
MEXU). Chihuahua: Batopilas, Knobloch 548 (MICH, MSC, US); Urique, Knobloch 887 (BM,
MSC). Colima: 27 mi NE of Colima, Clarke et al. 2137-1 (MICH); Tecoman, Emrick 141 (F); 9 mi
E of Colima, 480 m, McVaugh 14997 (MICH); Colima, Palmer 150 (C, MICH, US); 10 mi E of
Colima, 550 m, Reeder & Reeder 2374 (MEXU); between Pihuamo & Colima, Templeton 8932
(F); 18 km S of Manzanillo, West 3511 (GH, MO). Guerrero: Zacacoyuca, Becerra s.n. (MEXU);
Temisco, Bruff 1320 (MEXU); carretera a las Grutas de Cacahuamilpa, German 176 (MEXU);
Cerro de Jumil, Germtn 464 (MEXU); Coyuca, Hinton 5840 (BM, K, US); Pungarabato, Hinton
5941 (A, K, MO); Coyuca, Hinton 6315 (BM, F, K), 711 (NY); 2 mi W of Coletlita, 2700 ft, Kubi-
cek 3746 (MICH), vic. Acapulco, Palmer 630 (A, BM, F, GH, K, MICH, MO, NY, P, U, US);
Santo Tomas, 270 m, Mexia 8934 (F, GH, K, MO, NY). Jalisco: Sin. loc.: Diguet s.n. (P); between
Crescentia 85

F 1 ~ ~ ?.

FI. 5.Cesenic "aa n"C

!:i'te '-,Ccjt~A ai /,B ri / Gnr
32 oa xs ' D lw I dra i 25 crl n etl o e anm)/; - C
"%, ~~
hbtx25(ra ==ro
7~G ?': ri / Gnr 0 .
'ai aia aMnail,1020m c/lg 10 MC) - mEo hml,3
m, c~ugh 2516 MIH);Isa Ccias,Peez551(MXU) Cscaot, Redws'""64

FIG. 15. Crescentia alata and C cujete. A-E, C. cujete; A, habit x 2/5, B, fruit x 2/5 (Gentry
352); C, ovary x.s. x 4; D, flower in dorsal view x 2/5; E, corolla in ventral view x 2/5; F-G, C.
alata; F, habit x 2/5 (Croat 273); G, fruit x 2/5 (Gentry 5043). (From Flora of Panama).

Bahia Navidad & La Manzanilla, 150-200 m, McVaugh 21002 (MICH); 7-8 km E of Chamela, 30
m, McVaugh 25216 (MICH); Isla Cocinas, Perez 551 (MEXU); Cascalote, Rzedowski 16647
(MICH). Mexico: Temascaltepec, 1230 m, Hinton 755 (A, K, NY); Ixtapan, 1000 m, Hinton 1165
(US); Tonatico, 1200 m, Matuda 27908 (MEXU); 10 km W de Nanchititla, Medrano et al. s.n.
(MEXU). Michoacan: Coahuayula, Emrick 156 (F); Pihuato, Jones 34 (US); Apatzingan, Leaven-
worth & Hoogstraal 1595 (F, GH, MO), 1709 (F, MO); La Salada, Nelson 6913 (GH, NY, US);
Las Mesitas, 1000 m, Langlasse 197 (GH, P, US). Morelos: near Amacuzac, Feddema 1643
(MICH); 23 km S of Cuernavaca, 5500 ft, Hitchcock & Stanford 7048 (F, GH); Morelos, Paray
460 (MEXU); Yautepec, Rose et al. 8590 (US); 25 mi N of Taxco, Rowell et al. 17M931 (BM, F,
NY, US); Tojutla de Juarez, Salazar s.n. (MEXU). Nayarit: Jesus Maria, 600-700 m, Feddema
86 FloraNeotropica

1309(MICH);2 mi NE of Mazatlan,675 m, McVaugh15339(MICH);Acaponetato Concepcion,

Rose 1529(US); Acaponeta,Rose et al. 14466(NY, US). Oaxaca:Tehuantepec,50 m, Conzatti
3690(MEXU,US); 2 kmE of SalinaCruz,King210 (MICH,US); 10-15km NW of Tehuantepec,
50 m, King767(MEXU,MICH,US); 2-4 km E of Tehuantepec,King1166(MICH,NY, US); 16
km NNE of Tehuantepec,King1465(US);Hilotepec,100ft, Nelson2572 (GH, US); SalinaCruz,
Orcutt3474(MO);SalinaCruz,3-100m, LI. Williams9677(F, MICH).San LuisPotosi: 17mi N
of Valles, 150 m, Fosberg29445 (US); Rasc6n,Pringle5293 (GH); 15 km N de Taumin,50 m,
Rzedowski10869(MICH). Sinaloa:Rinc6nde Santo Conitaca,Conzatti794 (MEXU);vic. of
Labradas,Ferris& Mexia 5129 (A); Cofradia,H. Gentry5043 (MO, NY); Las Flechasto La
Rastra,Goldman315 (GH, NY, US); Las Palmas,Gonzalez732(MEXU);nearMazatlAn,Gregg
1208 (MO);Mazatlan,Jones 22354 (MO); Villa Union, Lamb 426 (GH); Los Labrados,5 m,
Mexia 948 (A); Villa de San Ignacio,Narvaez& Salazar420 (US); Rinc6nde Santo Coniteca,
Narvdez& Salazar794(US);San Ignacio,J. Ortega420 (F, K);Mazatlan,Ortega6527(GH, US),
sin. loc. Ortega7302(F); Villa Union, Rose et al. 13898(NY, US); Mazatlan,Rose et al. 14021
(NY, US);betweenMazatln & VillaUnion, Shreve6398(F);betweenEscuinapaand Acaponeta,
Templeton7492(MICH);El Salto Grande,Vela1452(MEXU).Sonora:Hermosillo,Gallego539
(US);Guaymas(cult.), Palmer85 (BM, GH, C, K, MICH,NY, U, US); vic. of Alamos, Rose et
al. 12910(GH, NY, US). Veracruz:Zacuapan,Coulter1007(K);Mozomboa,60 m, Doranteset
al. 843 (F); nearPlan del Rio, Gillyet al. 69 (MICH);2 km S of Tampico,Palmer434 (F, GH, K,
GUATEMALA.Alta Verapaz:Cahabon1904, Cook s.n. (US); Santa Mariade Cahabon,
250 m, Pittier212 (BM, US). Guatemala:Fiscal, 3700 ft, Deam 6065 (F, GH, MICH, MO, NY,
US); nearFiscal, Standley59607(F), 80375(F). Jalapa:El Rancho, 1000ft, Kellerman7050(F,
US). Jutiapa:near Jutiapa, 900 m, Standley60505 (F), 850 m, 75020 (F). Santa Rosa: near
Cuilapa,895 m, Standley77959(F); SE of Chiquimulilla,150m, Standley78775(F);vic. Chiqui-
mulilla,325m, Standley79285(F). Zacapa:PuertoBarriosHwy, 3 mi NE of Zacapaturnoff,200
m, Croat41890(MO);Gualan, 122m, Kellerman5710 (MO, US); vic. Zacapa,200 m, Standley
74208(F);nearLa Fragua,200-500m, Standley74386(F);nearSantaRosalia,250-300m, Steyer-
EL SALVADOR.La Libertad:near Comasagua,900 m, Carlson261 (F). San Salvador:
Tonacatepeque,Calder6n241 (GH, US); vic. Tonacatepeque,Standley19425(GH, US). San Vi-
cente:San Vicente,350-500m, Standley21685(GH, NY, US). Sonsonate:vic. of SanAntoniodel
Monte,250m, Standley22145(GH, NY, US).
HONDURAS.Choluteca:10 mi S of Choluteca,Croat273 (MO);4 km E of Choluteca,
Davidse& Pohl 2411 (MO); 11 km E of Choluteca,Harmon& Fuentes5955 (NY); vic. Cholu-
teca, Standley24490(F), L. Williams&Molina 10856(F, GH, MEXU).Comayagua:Las Limas,
3000ft, EdwardsP-127 (A, F), P-376 (A, F, K, US); vic. Comayagua,600 m, Standley& Chac6n
5836 (F). MorazAn:SantaClara, Glassman1573(F, NY); El Zamorano,Molina25791 (BM, F,
MO, NY); vic. El Zamorano,Standley131(F); Yeguare,Valerio358 (F). Olancho:vic. Juticalpa
NICARAGUA.Sin. loc.: Levy 1383 (P); Wrights.n. (GH, US). Chinandega:vic. Chichi-
galpa, 90 m, Standley 11243 (F), 11317 (F), 11525 (F). Chontales:between Boaco cutoff &
Acoyapa,100-250m, Bunting&Light692(F, NY);vic. Juigalpa,160m, Standley9364(F).
COSTARICA.Guanacaste:nearCailas,Brenes12719(CR);nearLiberia,Gentry356 (MO,
WIS);nearCaflas,Gentry814 (MO),872 (MO),1352(MO);nearLiberia,Lems100(F); 16km SE
of Liberia,Lems5136(NY);nearLiberia,Lent2548(CRU);nearCaflas,Little&Budowski20132
(CR);SE of Bagaces,Little& Budowski20293(CR);betweenSantaRosa & La Cruz,Hatheway
1447 (CR); Hac. Sta. Rosa, Oersteds.n. (BM); near Caflas, Salas S-549 (CRU); near Liberia,
Taylor4414(NY); 15km SE of La Cruz, Wilbur&Stone9768(F).
PANAMA.Chiriqui:David(cult.), Seemann1130(K).

Local names.Mexico:Sinaloa:tecomate,ayale, tecomata.Jalisco:sirian.

Michoacan/Guerrero:cirian, sirian, syrial, tecomate. Oaxaca: jicara, jica-
rita, morro. Guatemala:morro, jicaro. El Salvador:morro, morrito. Hon-
duras:jicaro, morro,guacal, morrito.Nicaragua:jicaro, jicarito. Philippines:
cruzcruzan,hoja cruz.
Crescentia 87

[4/~~~~~~~ '

,I. ,6. M.a. Do..... (e, 11.

, ne........J(stars_,

FIG. 16. Map 3. Distribution of Crescentia. C. alata (closed circles), C. linearifolia (stars), C.
mirabilis (stars in circles), C. portoricensis (open circles).

Occasional hybrids with C. cujete are encountered. These have mostly

simple leaves with some incompletely 3-foliolate leaves and the small fruits of
C. alata. Collections which are apparently hybrids are Hinton 5804 and 7111
(K) from Coyuca, Guerrero, Mexico, Standley 22146 (US) from Sonsonate, El
Salvador, Steyermark 37816 (F) from San Marcos, Guatemala, and Gentry
813 (MO) from Guanacaste Province, Costa Rica.
This species is rarely utilized or cultivated by the campesinos, who prefer
its larger-fruited relative, C. cujete. The fruits are sometimes used for cups,
ladles, and rattles. The three leaflets and winged petiole together form a kind
of cross and the tree has acquired some religious significance from this resem-
blance. According to Seemann (1854), the chief use of the fruit in Mexico is to
boil the pulp with sugar as a remedy for complaints of the chest (consumption?).

2. Crescentia amazonica Ducke, Arq. Inst. Biol. Veg. 4: 61. 1938. Figs. 17, 18.

Small tree to 10 m; branchlets mostly lacking, smaller branches thick,

subterete, with alternate short-shoot projections each bearing a fascicle of
leaves from its center; leaves of various sizes in each fascicle, simple, oblance-
olate to narrowly obovate, acute, attenuate at base, 5-22 cm long, 1-5.3 cm
wide, petiole merging gradually into leaf base, scattered lepidote, otherwise
glabrous or with a few scattered trichomes along midvein. Inflorescence of one
88 Flora Neotropica

FIG. 17. Crescentia amazonica, C. linearifolia and C. mirabilis. A-B, C. linearifolia; A, habit
(Liogier 16871); B, fruit (Ekman 8822). C, C. mirabilis (Ekman 15535). D-F, C. amazonica; D,
pre-anthesis flower (Gentry et al. 25811); E, fruit (Gentry et al. 25811); F, habit (Ducke s.n., RB
34696). (All x 1/2 except C. amazonica leaves x 1/3).
Crescentia 89

FIG. 18. Map 4. Distribution of Crescentia amazonica.

or two flowers, cauliflorous. Flowers indistinguishablefrom those of C.

cujete. Fruit a calabash, subspherical to oblong-ellipsoid, 4-4.5 cm diam., 5.5-
7(-8 cm fide Aristeguieta5808) long, the shell smooth, thin, hard;seeds small,
thin, wingless,scatteredthroughpulpof the fruit.
Type. Brazil. Amazonas: Borba, Rio Madeira, Ducke s.n. (RB 34697)
(holotype,RB; isotypes, K, MO, US).
Distribution.Along the upperand middleAmazonand Orinocoand their
majortributariesin seasonallyinundatedvarzea.
COLOMBIA. Matacormena, Cuatrecasas 4292 (F, US).
VENEZUELA. Amazonas: cerca de Tamatama, 121 m, Williams 15274 (F, VEN); Alto
Orinoco, Croizat 856A (VEN); Brazo Casiquiare, Holt & Blake 677 (NY, US, VEN). Apure: Isla
Orupe, Trujillo 10839 (MY); entre el Rio Meta y el Caflo, Velez 2596 (VEN). Barinas: Reserva
Forestal Caparo, Marcano-Berti 2806 (MER). Bolivar: San Felix-Pto. Ordaz, rebalses del Caroni,
Aristeguieta 5808 (VEN); alrededores de Moitaco, Trujillo 5870 (MY); La Paragua, 80 m, Wil-
liams 12757 (F, US, VEN). Guarico: selva de galeria del Rio Tiznados, Aristeguieta & Zabala 7087
(VEN); estero de Camaguan, Aristeguieta 7791 (VEN).
BRAZIL. Amazonas: Sao Paulo de Olivenca, Ducke 552 (F, IAN, MO, NY, US), s.n. (RB
34696) (MO, P, RB, U, US); Codajas, Laguinho, Ferreira 58-252 (INPA), 6455 (IAN, INPA).
PERU. Loreto: mouth of Rio Nanay, 120 m, Gentry & Revilla 21162 (F, MO, USM); Rio
Atacuari, vic. of Isla Tigre, Mori et al. 9214 (MO); Yarinacocha, Rio Ucayali, Tessmann 3478

Local names. Venezuela: tapara montanera, totumo, taparo de agua,

tapara. Brazil: cuia pequena do igapo, cuia maraca, cuiupi, cuia pequena.
90 FloraNeotropica

Thisspeciesof the seasonallyinundatedvarzeaand tahuampaforestsmay

prove no more than a small-fruitedwild form of widely cultivatedC. cujete.
Its fruitsare constantlysignificantlysmallerthan those of cultivatedC. cujete
but such a changemightbe expectedin naturalizedpopulationsof C. cujeteif
artificialselectionfor the largefruitssuitablefor containerswererelaxed.Bio-
geographicallyC. amazonicais a bit improbable-all other Crescentieaeare
found in CentralAmericaand the West Indies (threeof these also reaching
northwest South America). Crescentia amazonica is mentioned by the earliest
Europeanexplorersso the introductionof its ancestors,if not natural,was at

3. CrescentiacujeteLinnaeus,Sp. PI., ed 1, 2: 626. 1753. Fig. 15.

C. acuminataHumboldt,Bonpland& Kunth,Nov. Gen. Sp. PI. 3: 157. 1819.Type:Hum-
C. arboreaRafinesque,SylvaTell. 81. 1838.Type:"Antillesand SouthAmerica",not des-
C. cuneifoliaGardner,J. Bot. (Hooker)2: 422. 1840.Type: Gardner3342, Brazil,Goyaz
C. angustifoliaWilldenowex Seemann,Trans.Linn.Soc. London,ser. 1, 23: 20. 1862.pro
C.fasciculataMiers,Trans.Linn.Soc. London,ser. 1, 26: 171. 1868.Type:Oersteds.n.,St.
Thomas(holotype,P; isotype,fragm.BM).
C. plectanthaMiers,Trans. Linn. Soc. London, ser. 1, 26: 170. 1868. Type: Mrs. A. Fry
s.n., Brazil,Rio de Janeiro(P).
C. spathulataMiers,Trans. Linn. Soc. London, ser. 1, 26: 173. 1868. Type: Sinclairs.n.,
C. cujete var. puberulaBureau& K. Schumann.In: Martius,Fl. Bras. 8(2): 403. 1897.
Lectotype:Dombeys.n., Peru,Lima(P).

Treeto 10m tall and 30 cm dbh., the branchesusuallycrooked,the crown

open; branchletsmostly lacking,smallerbranchesthick, subterete,with alter-
nate short-shootprojectionseach bearinga fascicle of leaves from its center.
Leavesof varioussizes withineach fascicle, simple, obovate, the tip obtuse to
acute, the base attenuate,petiole lacking, (1.5-)4-26cm long, (0.7-)1-7.6 cm
wide, chartaceousto rigid-chartaceous,the midvein raised above; lepidote
above and beneath,otherwiseglabrousabove, beneathglabrousor pubescent
along midveinwith simpleand forked trichomes,plate-shapedglandsat base
of blade beneath, drying grayish-olive.Inflorescenceone or 2 cauliflorous
flowersborneon largerbranchesor trunk,the pedicellepidote, 1.5-3cm long.
Flowerswith calyx bilabiatelysplit to the base, each lobe 1.8-2.6 cm long and
1.3-2.4 cm wide, mostly glabrouswith plate-shapedglands on the upperhalf
of lobes, slightlylepidoteat the base;corollayellowish-tanwith purplishvena-
tion on the lobes and purplishlines on the tube outside, tubular-campanulate
with a transversefold midwayacrossthe lower side of the throat, fleshy, 4.1-
7.4 cm long and 3.1-4.5 cm wide at the mouth of the tube, the tube 2.8-4.5 cm
long, the lobes triangularwiththe apexextendedas a narrowpoint, 2.5-3.1 cm
long, sparselylepidote or stalked-lepidoteon the tube outside and near the
mouth of tube inside, more denselyso at level of stameninsertion,papillate-
glandularon the lobes outsideand sparselyso inside;stamenssubexserted,the
Crescentia 91

antherthecaethick, partiallydivergent,5-8 mm long, ca. 3 mm wide, the fila-

ments2.8-3.3 cm long, inserted7-15 mm from the base of the tube, the stami-
node 1-3 mm long, inserted5-11 mm from the base of the tube; pistil 4-5 cm
long, the ovary roundedconical, 5-7 mm long, 4-7 mm wide, 3-6 mm thick,
lepidote, the ovules multi-seriateon 4 placentae;disc annular-pulvinate,3-4
mm long, 8-11 mm wide. Fruita pepo or calabash,sphericalto ovoid-elliptic,
(8-)13-20cm in diam., to 30 cm long, the thin hard shell smooth, lepidote-
punctate;seeds small, thin, wingless, 7-8 mm long, 4-6 mm wide, scattered
throughthe pulp of the fruit.
Type. Specimenwithoutinformation(LINN779.1).
Distribution.Extensivelycultivatedthrough most of tropical America,
the naturalrangeobscure.Certainlynativeat least to northernCentralAmeri-
ca and Mexico. Apparentlywild-growingtrees elsewhereare probably de-
scendedfrom cultivatedtrees.
MEXICO.Sin. loc.: 1857, Schotts.n. (F). Campeche:Champolan,Goldman542 (US); 48
km sobre el Rio Candelaria,20 m, Menendez532 (F); 26 km E of Escarcega,Sallee E 575
(MEXU);vic. of Xpujil, 153km E of Escarcega,300 m, Shepherd62 (F, MICH,WIS). Chiapas:
TanateRiverat Habenal,3300ft, Breedlove12745(DS, MICH);Rio Grijalva,munic.Acala, 1600
ft, Breedlove2825 (DS); Chicla, Collins& Doyle 109 (US); road to BalacAn,40 m, Davidse &
Davidse9403(MO);3 km N of PuebloNuevoSolistahuacan,Lathrop5129(DS);Escuintla,cult.,
Matuda16643(F, MICH);Acagoyagua,Matuda16818(F, MICH);Tanat6River,munic.Tene-
japa, 3200 ft, Ton 394 (DS). Guerrero:Pungarabato,cult., Hinton et al. 5804 (A, BM, K, US),
7112(A, K, NY); Acapulco,Palmer149(GH, K, US), Thiebaud41 (P). Nayarit:vic. of San Bias,
Ferris6850(A, US). Oaxaca:Palapa,distr.Tuchilan,200 m, Conzatti3719 (MEXU,US); Tolo-
sita, 70 m, cult., Ll. Williams(F). Sinaloa:Sinaloa,Ortega4533 (US). Tabasco:BalancAn,cult.,
Calzada2376 (F); carretera10 jct. with carretera20, 35 m, Novelo et al. 92 (MO); cruce de
caminos Villahermosa-Teapa y Teapa-Pichucalco,50 m, Pennington& Sarukhan9149 (NY);
sabanasentrelos Rios Chabley ChumpAn,50 m, Pennington& Sarukhan9192 (NY). Veracruz:
Tlacotalpan,Nelson 488 (US);Tenochtitlan,22-75m, Wing66 (GH). Yucatan:sin. loc., Gaumer
437(A, BM, C, F, GH, MO, NY, US), Gaumer& sons 23773(F, GH, US); Izamal,cult., Gaumer
&sons 23804(F, US); Progreso,Goldman672 (US);Merida,Schott521A (F); Piste, Steggerda56
GUATEMALA.Alta Verapaz:Cubilquitz,350 m, von Tuerckheim8724(US). El Progreso:
4-8 km S of El Progreso,570m, Iltis G-65(WIS).Escuintla:Masagua,400 ft, DonnellSmith2096
(GH, K, US); betweenMasagua& San Jose, 75 m, Standley64142(F); near San Jose, Standley
64002 (F). Peten: La Libertad,cult., Lundell2325 (A, MICH), 3279 (A, MICH). Quiche:El
Rabinal,Cook 7456(US). Retalheu:Champerico,0 m, Standley66605(F); betweenNuevaLinda
andChamperico,120m, Standley87726(F). SantaRosa:vic. Guazacapan,220 m, Standley78670
(F). Suchitepequez:Mazatenango,1905,Kellermans.n. (US); Santo Tomas, Marshall28 (GH).
BELIZE.Belize:19.5mi NW of Belize,Croat23295(MO);TurneffeIsland,Egler42-23(F);
BelizeForestDept. s.n. (K); 11 mi N of Belize, Gentry7850(MO), 7851 (MO);road to Burrell
Boom,Jansen&Ives 1122(MO);5 mi W of Belize, Wiley212 (MO).El Cayo:Pine Ridge,Bartlett
11212(US); LittleMountainPine Ridge,Bartlett13103(MICH,US). OrangeWalk:nearHoney
Camp,Meyer23 (F). Toledo:SanAntonio,Barlett13048(MICH,US).
HONDURAS.Atlantida:LancetillaValley,introduced,Standley54402(A, F, US). Col6n:
WisperniniCamp,50-75ft, von Hagen& vonHagen1400(F, NY). Morazan:Montailade la Flor,
von Hagen& von Hagen1295(NY). Olancho:betweenJuticalpaandEl Asib, 380-400m, moreor
EL SALVADOR.La Libertad:nearComasagua,900 m, Carlson(F). San Salvador:Tonate-
peque, Calderon239 (NY, US), 240 (US), 241 (NY), Standley19477(GH, US), Standley19497
(GH, MO, US), Standley19498(US).
92 Flora Neotropica

COSTA RICA. Alajuela: QuebradaMaquina& Azul near San Carlos, Fournier 1580
(CRU);nearCapulinon Rio Grandede Tarcoles,Standley40138(US). Guanacaste:nearCaflas,
Daubenmire46 (CRU),514 (CRU), Gentry403, 813, 868, 869 (all MO);Playasdel Coco, Gentry
352 (MO, WIS);Nicoya, Tonduz13644(K, NY, P). Heredia:PuertoViejo, Gentry1182(MO).
Lim6n:Cauhita,Gentry1280, 1292, 1293(all MO);NW of PuertoLim6n,Godfrey66416(MO).
Puntarenas:Puerto Jimenezde Osa, Brenes12125(CRU);Cabo Blanco, Burger& Leisner6705
(F, NY); below Rio Barrancabridge, Gentry1368 (MO); Mata de Lim6n, Maja & Salas 4-22
(CRU), Ortizs.n. (HNCR6767)(CRU);Terraba,Pittiers.n. (US); Matade Lim6n, Ugalde6358
(CRU).San Jose: Villa Col6n, Echeverria62 (HNCR34462)(CRU, F); Rio Torres, Tonduzs.n.
(HNCR1310)(BM, CR, US); San Jose, 1100m, Tonduz7098(HNCR1310)(BM, F, GH, K, NY,
PANAMA.Bocasdel Toro:nearAlmirante,Gentry2807 (MO);MarianoCreek,von Wedel
2894 (GH, MO). CanalZone: nearLos Rios, Gentry3690 (MO);GatunStation,Hayes 94 (BM,
GH); BalboaHeights,Sharps.n. (SCZ);ChivaChivaTrail, Tyson&Blum3565 (SCZ).Chiriqui:
Sin. loc.: Wagoners.n. (M);Progreso,Cooper&Slater288 (US). Darien:villageof Paya, Gentry
4397(MO);El Real, Gentry4505(MO).Los Santos:S of Pedasi,Stimson5293(MO,SCZ).Pana-
ma: Caflitas,Gentry1405(MO);Rio Maestro,Gentry2245(MO);Rio Piragua(Tabardi),Gentry
2523 (MO); Bella Vista, Standley25343 (US); Taboga Island, Standley27885 (US); between
MatiasHernandesand JuanDiaz, Standley31974(US). San Blas:mountainsabovePuertoObal-
dia, Gentry1478(MO).
BAHAMAS.Sin. loc.: 1716,Dale s.n. (BM). Acklin'sIsland:Brace4447 (F, NY). Cat Is-
land: Byrne350 (A, WIS). Eleuthera:Correll& Hill 45241 (NY). Great RaggedIsland:Gillis
CUBA. Sin. loc.: de la Sagras.n. (P), 324 (P); Lomadel Jaguey,700 m, Eggers5292 (K).
Camaguey:nearCamaguey,Brittonet al. 13240(NY);La Gloria,Shafer29 (NY, US); La Gloria,
in swamp,Shafer343 (NY, US). Habana:Havanna,Humboldt& Bonpland1300(P-HB);Mad-
suga,Shafer27 (NY). Islade Pinos:nearNuevaGerona,Curtiss311 (BM, F, GH, K, MO, NY, P,
US);NuevaGerona,cult., Killip43846(US). LasVillas:vic. of Soledad,Howard4976(GH, MO,
NY, U, US); Josefa, Soledad, Jack 4044 (A); Limones,Soledad, Jack 5400 (A); Santa Clara,
Rutten-Pekelharing 380 (U); Cieneguita,Combs9 (F, GH, K, MO, NY). Matanzas:nearMatan-
zas, Rugel663(NY). Oriente:GuantanamoBay, Britton2205(NY);El Cobre,Santiago,Bro. Cle-
mente 6055 (GH); Monte Picote near Palmaritodel Cauto, 400 m, Morton 9692 (US); S of
Holguin,Shafer1355(GH, NY, US); sin. loc., Wrights.n. (GH, NY);MonteVerde, Wright361a
(BM, GH). Pinardel Rio: Rio San Juande Zayas,Shafer11839(NY, US); Buenaventurato San
Juande Guacamalla,Wilson9331(NY, U).
CAYMANISLANDS.GrandCayman:W. Bay, Brunt2110 (BM); 1 mi SE Georgetown,
JAMAICA.Sin. loc.: BenzonEE152-4172(C). Middlesex:Moneague,Alexanders.n. (K);
near Widcombe,NE of Liguanea,1000ft, Proctor 10190(BM);FerryPen, Harris6533 (BM);
nearLydfordPO, ca. 1400ft, Howard&Proctor14273(A, BM);ClarendonParish,1-1/2 mi S of
May Pen, Stearn213 (BM);PortlandRidge, ClarendonParish,50 ft, Steam 788 (A, BM);Tre-
lawny, cult., Whitefoord1478 (BM). Surrey:vic. of Kingston,St. AndrewParish, Britton 786
(NY);nearPort Antonio,PortlandParish,Fredholms.n. (US);E of Rockfort,Harris9382(NY);
Kingston,Hitchcocks.n. (MO);CrossRoadsPO, St. AndrewParish,SteamC27(BM).
HAITI. BetweenHinchaand San Rafael,Cook et al. 133 (US);Massifde la Pille, Port-au-
Princeto Nornea Bateau,EkmanH8034 (US); Massifde la Hotte, EkmanH10856(US); Prin-
cipio, Jaeger88 (P, US); La Vallee,TortueIsland,Leonard& Leonards.n. (US); Bayeux,near
PortMargot,Nash 917 (NY);Port-au-Prince,Rose22088(NY, US).
REPUBLICADOMINICANA.SantoDomingo,sin. loc., sin. coll., ex herb.Maire(P), ex.
herb.Poiteau(P); San Pedrode Macoris,Rose et al. 4308(NY, US); sin. loc., 1857,Schomburgk
s.n. (K).
PUERTORICO.DesecheoIsland,Brittonet al. 1603a(NY); Llanosde Coamo, Britton&
Britton9454 (NY); Candelaria,Cook & Collins256A (US); Areciba,Heller & Heller334 (NY,
US);Cambalache,Little 13462(F, US); CaboRojo to Boqueron,Little13622(F, NY, US);Mona
Island,Sardinera,Little & Wadsworth7759(NY); Mona Island,Little & Wadsworth16463(F,
US, WIS);Yuncos,Mt. Santode Le6n, Sintenis2740 (US);propeManati,Sintenis6546(BM, F,
GH, K, MO, NY, P, US);Rio Piedras,Stevenson5566(US).
VIRGINISLANDS.St. Croix:La Grange,Eggers873(C);Bassin,Ricksecker70(F, MO, P,
Crescentia 93

US), 73 (F, MO, NY, US); Anna's Hole, Thompson 467(NY), Thompson 1062 (NY). St. Thomas:
Boboni, Oersted s.n. (BM, P); Foden at Flaghill, Ostenfeld s.n. (P), 478 (C). Tortola: Pasea Hall
Estate, Fishlock 208 (NY).
LEEWARD ISLANDS. Antigua: Confort Hall, Box 1546 (BM, US); Boggy Peak, Rose et
al. 3432 (NY, US). Nevis: Cunu Gut, cult., Beard 468 (K). Saba: Booby Hill, Fra. Arnoldo 787
(U); sin. loc., Boldingh 1930b (U). St. Barthelemy: St. Jean, Questel 201 (P), 231 (NY), 868 (NY).
Guadeloupe: Sin. loc.: Bena s.n. (P), Duchassaing s.n. (P), Questal 1408 (P, US); Trois Rivieres,
Rodriguez 3263 (P); St. Maire, 25 m, Stehle 1080 (A, US). St. Eustatius: Boldingh s.n., 2954 (both
U); Stoffers 3814 (U).
WINDWARD ISLANDS. Dominica: Taylor 118, 119, 120 (all GH); Bataca, Bonbee 6369
(P). Martinique: Fort de France, cult., Duss 1236 (F, NY, US); Basse Terre, Duss 3066 (NY); sin.
loc., Hahn 889 (K, P); Foret de la Calabasser, Hahn 1436 (BM, P); sin. loc., Plee s.n. (P). St.
Lucia: sin. loc.: without collector (very old) (BM); Marquis Estate, Proctor 18122 (BM, US). St.
Vincent: Smith & Smith 217? (K). Grenada: 1905, Broadway s.n. (NY).
SOUTHERN DUTCH ANTILLES. Aruba: Fra. Arnoldo 267 (U); Boldingh s.n. (U); Bol-
dingh 6414, 6416 (both U); Curran & Haman 402 (GH). Bonaire: Punk Vierkant, Stoffers 705 (U);
Kust Sta. Barbara, Stoffers 791 (U); Put. Bronswinkel, Stoffers 817 (U); Montague, Stoffers 879
(U); Hobau, Stoffers 1023 (U); Karpata, Stoffers 1081 (U); sin. loc., Boldingh 7916 (U). Curacao:
Fr. Arnoldo-Broeders 3903 (A); Shalome, Stoffers 19 (U); Sta. Barbara, Stoffers 412 (U).
COLOMBIA. Antioquia: Medellin, Archer 423 (US), Toro 121 (NY); Paso de Caramanta,
600-800 m, probably planted, Pennell 10843 (GH). Atlantico: Puerto Colombia, Elias 1043 (F,
US). Bolivar: San Martin de Loba, cultivated and wild, Curran 184 (US); La Popa near Carta-
gena, 50-175 m, Killip & Smith 14072 (A, GH, F, NY, US); Vilches, Rio Sinu, Pennell 4707 (NY).
Casanare: Orocu6, 140 m, Cuatrecasas 4348 (US). Cundinamarca: Tocaima, 435 m, Andre 1821
(K, P); La Mesa, 950-1370 m, Garcia-Barriga 12174 (US); 1-6 km N of Puerto Bogota, 550 m,
Gentry et al. (MO); Tocaima, Perdz-Arbeldez 2475 (US). Guajira: Serrania La Macuira, Saravia
2355 (US). Magdalena: Santa Marta, Smith 1132 (F, GH, K, NY, P, US). Norte de Santander:
Ocafia to Pamplona, Kalbreyer 1015 (K). Santander: near Baranca, Dugand 149 (F). Valle: Rio
Calima, 5-50 m, Cuatrecasas 6429 (F).
CULTIVATED: Venezuela, Trinidad (also naturalized: Britton et al. 2594 (GH, NY, US),
Couva Bay, border of mangrove swamp), Guyana, Surinam, Guyane Francaise, Ecuador, Peru,
Brazil, Bolivia, Paraguay, Bermuda, Indonesia, Philippines, Society Islands, Ghana, Nigeria,
Liberia, India, Vietnam.

Local names. English (Jamaica, Bahamas, Grand Cayman, Bermuda,

Belize): calabash, wild calabash, tree calabash. French (St. Barthelemy, Marti-
nique, Guyane Francaise): calebassier. Dutch (Curasao, Aruba): kalabash,
calbas, calbas rondo. Dominica: callebasse longue, callebasse coricon, calle-
basse boite. Cuba: giiira, guira del monte, calabasa. Puerto Rico: higuera,
higuero. Mexico: west coast (Sinaloa to Guerrero): tecomate, ayale, cerial,
cirian mazo. Yucatan (Maya): hoco, luch, huaz. Eastern and southern Mexico
to Honduras: jicara, jicaro, morro. Honduras (Wisperini): Kabami. Costa
Rica: jicaro. Panama: totumo. Colombia: totumo, totumo cimarron. Vene-
zuela: totuma, tapara. Ecuador: mate, pilche, pilchimate. Peru: coast:
totuma, totumo. Amazonia: tsapa, huingo, pati, sacha huingo, buhango. Bra-
zil: cueira, cuia. Bolivia: porobamba.

Uses. Extensively cultivated for the hard shells of its fruits which are
widely used as household utensils, bailers for canoes, and as a storage con-
tainer. As already noted, use of the fruit was already widespread in pre-
Colombian times. The pulp is sometimes used medicinally as in the Cuban
"jarabe de giiira, used against chest infection or as a laxative (Leon & Alain),
or for "tortions of the Guts or dry Gripes" (Wafer). The pulp is occasionally
94 FloraNeotropica

reportedas more or less edible as by Waferwho reportsthat the Indiansfre-

quentlysuckedout the juice (spittingout the pulp) on forced marches.In the
early 1800's the fruit was eaten on the Ecuadorianisland of Puna (Barclay
361). Even an infusion of the flowersis used for earache(Leon & Alain) and
for whoopingcough (Marshall28). In Guyanathe seeds are used as a contra-
ceptiveand the leaf boiledwith othersto give a feverremedy(Hardy302). The
fruitshellis often paintedor engravedand usedas a curioor wall-hanging.
Two variantsof this polymorphicspeciesmay deservetaxonomicrecogni-
tion. Both differ from typical C. cujetein smallermore coriaceousleavesand
fruit. One of these, the commonspeciesof the coastal savannasof Belize, is a
dominantmemberof the savannahformationand forms completelyhomo-
geneous populationswith 1.5-9 cm by 0.8-2.5 cm leaves and 8 cm diameter
fruits. It seems more distinct from cultivated C. cujete than, for example,
many of the acceptedCubanspeciesof Tabebuiaare from each other. Addi-
tional field workis neededto evaluateits status.The secondsmall-leavedvari-
ant of C. cujete occurs in the MountainPine Ridge region of westernBelize
and adjacent Peten-Belize. Cayo: Mountain Pine Ridge, Bartlett 13103,
11212. Guatemala.Peten: La Libertad,Lundell3279. It has 0.7-1.7 cm wide
leaves less than 8 cm long and is thus very similarto the coastal savannah
form, but can easilybe distinguishedby discolorousleaveswith conspicuously
light tannish undersurfaces.If additional collections prove that naturally
occurringpopulationsof C. cujetefrom coastalMexicoare uniformlylike cul-
tivatedC. cujetein largefruitsand largethin leaves,thesetwo allopatricsmall-
leavedpopulationswould probablyhave to be considereddistinctsubspecies.
However,the apparentlynaturaloccurrenceof a similarsmall-leavedvariant
on Curacao-said to develop large thin leaves when cultivatedand irrigated
(Stoffers,pers. comm.)-suggests that much of this ecotypicvariationis only
4. CrescentialinearifoliaMiers,Trans.Linn. Soc. London26:172. 1868.
Figs. 16, 17.
CrescentiamicrocarpaBello, Anales Soc. Esp. Hist. Nat. 10: 294. 1881. Type: Bello s.n.,
PuertoRico(not seen).

Smalltreeor shrubto 7 m tall and 20 cm dbh, with open crownof slender

spreadingwand-likebranches. Leaves in fascicles of 2-6 from the enlarged
nodes, usually more or less linear, often almost needle-like,0.5-8.8(-15) cm
long, 0.1-1.8(-2.5)cm wide, roundedto acuminateat apex, cuneate-attenuate
to base, petiole lacking, chartaceous,secondaryvenationstronglyascending,
very indistinct, hardly differentiatedfrom similarlyindistincttertiaryvena-
tion, conspicuously scattered lepidote below, otherwise glabrous, drying
grayish-olive.Inflorescenceusually a single axillary flower, scatteredalong
branchtips, the pedicels0.5-1 cm long, glabrousor sparselysubpuberulous.
Flowerswith calyxbilabiatelysplitto base, eachlobe 1.1-1.5cm long and 0.8-1
cm wide, glabrous except for some scatteredplate-shapedglands; corolla
greenishwith brown striations, tubular-campanulate with a transversefold
midwayacrosslower side of throat, fleshy, 3-3.5 cm long, 1.5-2.5 cm wide at
Crescentia 95

mouthof tube, the tube 3-3.5 cm long, the lobes irregularlytriangularwiththe

apex extendedas a narrowpoint, glandular-lepidoteon lobes and sparselyon
outside of tube; stamens exserted or subexserted,the anther thecae thick,
divergent,4-5 mm long; ovary ovoid, the ovules multiseriate;disc annular-
pulvinate.Fruita pepo or calabash,globose to ellipsoid, 3-5 cm long, 3-4 cm
wide, the thin hard shell smooth, lepidote-punctate;seeds small, thin, wing-
less, scatteredthroughthe pulpof the fruit.
Type. Virgin Islands. St. Thomas:Oersteds.n. (holotype, P; isotype, C
(as 7963),MO; fragm., BM).
Distribution. Dry coastal forests below 100 m from Hispaniola and
Puerto Rico to the northernmostLesser Antilles; apparently disjunct in
REPUBLICADOMINICANA.IslaSaona,Alain&Alain28167(NY);E of JuanLopez,Bara-
hona Peninsula,EkmanH6933 (A, F, GH, S); Coueve-en-hautto Trujin, BarahonaPeninsula,
Ekman H7039 (K, MO, NY, S, US); near Cabo Rojo, Liogier 16871 (F, NY); Castillo de la
Isabela,J. Jimenez2323(US).
HAITI.GonaveIsland,Ekman8822(S); GonaveIsland,Eyerdam227 (GH, NY, US); Tor-
tue Island,Leonard&Leonard15324(NY, US).
PUERTORICO. Bahia Puerco, Britton& Britton8853 (NY); Guanica,Britton& Shafer
1906(NY, US); propeCabezade San Juan, Sintensis1898(MO, NY, P); propeCoamo, Sintensis
2980(BM,GH, K);Montalva,Brittonet al.4818(F, MO, NY), 8297(NY, US); IslaPifteros,Little
21669(BM,GH, NY, US).
VIRGINISLANDS.St. Thomas:Oersted7963(C);Tortola:ex. herb.Richard(P). St. Jan:
Britton&Shafer601(NY);Raunkiaers.n.(C). St. Martin:Boldingh2554B (NY).
LEEWARDISLANDS.SaintBarthelemy:LeGallo960 (U), 967 (A, NY, U).

Local name. Puerto Rico: Higiierita, higiierito. Dominican Republic:


Some Puerto Ricanand VirginIslandmaterial,notablyLittle21669 from

PifterosIsland and Raunkiaers.n. from St. Jan, has ratherdifferent larger
oblanceolateto verynarrowlyellipticleavesto 15 cm by 2.5 cm and mightrep-
resentintrogressionfrom C. cujete or possibly "foundereffect" variationin
an isolatedsmallislandpopulation.
The single mainlandcollection from northernBelize cited above is re-
ferredherewith muchhesitation.Morphologicallyit falls well withinthe range
of variationof West IndianC. linearifoliabut it is not very differentfrom the
small-leavedBelize populationof C. cujete of which it could be an extreme
form. Its leaves are 2-8 by 0.4-1.2 cm, and the clearlylinearlargestleaves are
20 timesas long as wide;a few microphyllousleavesalso occurat some nodes.
If this collectionwereincludedunderC. cujete, as geographicalconsiderations
mightsuggest,then C. linearifoliaitself would probablyhave to be reducedto
the synonymyof C. cujete. It is unfortunatethat no fruitswerepreservedwith
the Belizecollection.

5. CrescentiamirabilisEkmanex Urban,FeddesRepert.22: 88. 1925.

Figs. 16, 17.
96 FloraNeotropica

Smalltree or shrub. Branchletsthick, the nodes enlarged.Leavesfascicu-

late at the nodes, tiny, obovate to elliptic, roundedat apex, attenuateat base,
5-25mm long, 4-10 mm wide, glabrous,coriaceous,dryingdarkabove, olive-
graybelow. Flowersas in C. linearifolia.Fruitsunknown.
Type. Cuba. Camaguey:insula Cayo Sabinal prope El Fuerte, Ekman
15535(holotype,S; isotype,NY).
Distribution. Coastal marshes of northeasternCuba; endemic to the
Puerto Padre region of extreme northern Oriente Province and adjacent
CUBA. Oriente:El Cupey, Puerto Padre, CurbeloX53 (Roig 5364) (NY); La Resbalosa,
PuertoPadre,CurbeloX81 (Roig5662)(NY).

The statusof this species,separatedonly by the smallerleavesfrom close-

ly relatedC. linearifolia,is distinctlytentative.Neverthelessthe tiny obovate
leaves of C. mirabilisgive it such a remarkablydifferent fascies from other
Crescentiaspeciesthat I am compelledto retainit, at least provisionally,pend-
ing accumulationof sufficientadditionalmaterialfor evaluationof intraspe-
cific variability.
The Roig collectionsfrom PuertoPadre,Oriente,have largerleaves(1.4-5
x 0.5-2 cm) than the type and suggestthat the differencein leaf size between
C. mirabilisand C. cujetemay proveinsufficientfor specificrecognition.

6. CrescentiaportoricensisBritton,Bull. TorreyBot. Club43: 457. 1917.

Fig. 16.
Vinelikeshrubwith long slenderbranches.Leavesmostlyin fasciclesof 2
or 3 (in partalternate,fide Little)from the slightlyenlargednodes, obovateto
elliptic-obovate,obtusely acute to abruptlyacuminateat apex, cuneate at
base, 5-15 cm long, 2-8 cm wide, notablyshiny, secondaryand tertiaryvena-
tion prominulousabove and below, tertiaryvenation tending to parallelthe
secondaryveins, dryingolive-grayabove, olive below, sparselylepidoteabove
and below, otherwiseglabrous,petiole slender,0.6-1.5 cm long. Inflorescence
a singleaxillaryflower, the pedicel 1-2 cm long. Flowerswith the calyxbilabi-
ately split to base, the lobes 2 cm long; corolla whitish, reticulate-veined,
tubular-campanulate, 4 cm long, the lobes ca. 1 cm long. Fruit(only immature
seen; largermeasurementsafter Little) a pepo or calabash, oblong-ellipsoid,
terete, acute at apex, distinctlyangulateat base, 7-10 cm long, 3-3.5 cm in
diam.;seedsthin, flat, ca. 5 by 5 mm.
Type. Puerto Rico. Maricao:Rio de Maricao, 500-600m, Britton, Stev-
ens &Hess 2455 (holotype,NY; isotype,NY).
Distribution.Rarein the foothills of southwesternPuerto Rico, between
300 and 800m.
PUERTORICO.IndieraFria, nearMaricao,430-800m, Brittonet al. 4523 (F, MO, NY);
Parmentiera 7


3. Parmentiera de Candolle, Rev. Bign. (Bibl. Univ. Geneve) 19. 1838, non

Shrubs to quite large trees; often armed with a spine below each node.
Leaves opposite or subopposite, mostly palmately 3-5-foliolate, often with a
slightly winged petiole. Inflorescence a single flower or fascicle of 2-3 flowers,
usually borne cauliflorously on the old wood or trunk, sometimes terminally.
Flowers with the calyx spathaceous, split ventrally to near its base, inconspicu-
ously lepidote; corolla white or greenish-white, large, broadly campanulate to
infundibuliform, usually with a transverse fold across the lower side of the
throat (tubular and much smaller in P. parviflora); stamens subexserted,
anthers glabrous, the thecae thick, straight, somewhat divergent; pollen grains
3-colpate, the exine finely reticulate; ovary cylindrical, the epidermis glandular
and appearing more or less lepidote when dry, the ovules multiseriate; disc
annular-pulvinate. Fruit elongate-linear to oblong, indehiscent, often longitu-
dinally ridged, with a firm, fleshy outer cortex and a fibrous-fleshy core; seeds
between cortex and core, small, flattened, encircled by a narrow vestigial wing.
Type species. P. edulis DC. = P. aculeata (HBK.) Seemann.
Nine species of Central America ranging from central Mexico to north-
west Colombia.

Key to Species of Parmentiera

1. Leaves palmately 4-7-foliolate. 9. P. valerii.
1. Leaves 3-foliolate (sometimes simple in part, rarely incompletely 4-foliolate in part).
2. Flowers tubular, less than 2 cm long; fruits terete and unridged, less than 5.5 cm long and
1.3 cm wide. 6. P. parviflora.
2. Flowers broadly campanulate, more than 3.5 cm long; fruits more than 11 cm long or con-
spicuously ridged and more than 2 cm wide.
3. Branchlets usually with 2 short incurved spines at most nodes; Mexico to Honduras.
4. Fruits deeply ridged, thick, more than 3 cm wide when fresh, less than 17 cm long;
leaflets entire, more than 1.5 cm long. 1. P. aculeata.
4. Fruits terete, slender, less than 1 cm wide, more than 16 cm long; leaflets often ob-
tusely dentate near apex, frequently as small as 0.3 cm long. 4. P. millspaughiana.
3. Branchlets unarmed (the nodes sometimes accentuated but not spiny); Nicaragua to
northwest Colombia.
5. Fruits slender, less than 2.4 cm wide even when fresh, terete or somewhat 8-angled,
not ribbed; flowers terminal or cauliflorous; central Panama to Colombia.
6. Petioles winged; calyx acute, not apiculate; more or less cauliflorous; central
Panama. 2. P. cereifera.
6. Petioles unwinged; calyx apiculate or acuminate; flowers terminal; eastern Pana-
ma or Colombia.
7. Calyx apiculate with an acumen 5 mm long; fruit terete, not ribbed when dry;
northwest Colombia. 7. P. stenocarpa.
7. Calyx acuminate, the acumen less than 3 mm long; fruit distinctly 8-angled;
east central Panama. 5. P. morii.
5. Fruits (where known) thick, more than 4 cm wide when fresh, octagonal and con-
98 Flora Neotropica

spicuously ridged in cross-section; strictly cauliflorous; Nicaragua to central

8. Flowers on conspicuously elongated short shoots; buds more or less blunt-tipped;
Nicaragua, above 1000 m. 8. P. trunciflora.
8. Flowers one or two per node on abbreviated short shoots; buds and calyces more
or less acuminate; Costa Rica and Panama, below 1000 m. 3. P. macrophylla.

1. Parmentiera aculeata (Humboldt, Bonpland & Kunth) Seemann, Bot. Voy.

Herald 183. 1854; Trans. Linn. Soc. London 23: 18. 1860. Fig. 19
Crescentia edulis Desvaux, J. Bot. (Desvaux) 4: 112. 1814, non P. edulis de Candolle. Type:
not indicated, Mexico, Oaxaca (n.v.).
C. aculeata Humboldt, Bonpland & Kunth, Nov. Gen. Sp. PI. 3:158. 1819.
C. edulis Mocihio ex de Candolle, Prodr. 9:244. 1845, pro syn., non Desvaux.
Parmentiera edulis de Candolle, Prodr. 9: 244. 1845. Type: Sesse & Mocifio, drawing, Mexi-
co, not seen.
Crescentia musaecarpa Zaldivar ex Heller, Reisen in Mexico 414. 1853. Type: not indicated,
Mexico, nom. nud. (n.v.)
Parmentierafoliolosa Miers, Trans. Linn. Soc. London 26: 166. 1870. Type: Schiede 1207,
Mexico, Veracruz (holotype, K; isotypes, BM, MO, US).
P. lanceolata Miers, Trans. Linn. Soc. London 26: 167. 1870. Type: Based on Crescentia
edulis Desvaux.

Small tree to 10 m tall, branchlets terete, glabrous or slightly puberulous,

the nodes armed with a thick thorn subtending each leaf. Leaves opposite or
subopposite, rarely alternate (some branches of Gaumer 23440), mostly 3-
foliolate, a few simple, rarely palmately 4-foliolate (some leaves of Hinton
5805 are pinnately 5-foliolate!), the leaflets elliptic-obovate, acute to obtuse,
entire, the base cuneate or attenuate, terminal leaflet 1.5-6 cm long, 0.6-3 cm
wide, lateral leaflets 1.5-5 cm long, 0.6-2.5 cm wide, with barbate chambers in
the nerve axils beneath; petiolules lacking or to 1 cm long, petiole 1.3-3.5 cm
long, strongly grooved on top, narrowly winged. Inflorescence a few-flowered
fascicle or reduced to a single flower, terminal or axillary near the branch tips,
or from nodes of older wood. Flowers with the calyx basically spathaceous,
sometimes additionally split, acute to short acuminate in bud, 2.5-4 cm long;
corolla white, campanulate, 5-7 cm long, 2-2.5 cm wide at the mouth; stamens
subexserted, the anther thecae somewhat divergent, thick, 5-6 mm long; ovary
oblong-linear, lepidote, the ovules multiseriate in each locule. Fruit to 17 cm
long, more than 3 cm in diam. when fresh, curved, costate with thick ribs;
seeds small, thin, 3-4 mm long, 3 mm wide.
Type. Mexico. Campeche: Humboldt & Bonpland s.n. (P).
Distribution. Mexico to Honduras, frequently in cultivation or in dis-
turbed areas along roads or near habitations; naturalized in Queensland,
MEXICO. Sin. loc.: Liebmann 8022 (C), 11348 (US). Campeche: Apazoli, Goldman 575
(US). Chiapas: Ocozocautla, 850 m, Breedlove & McClintock 23526A (DS); Rosario, 2500 ft,
Laughlin 862 (DS); Chicomuselo, 800 m, Matuda 4491 (A, F, MO, NY); Rosario, 2500 ft, Pittier
1860 (US). Guerrero: Copacabana, Boege 456 (MEXU); Coyuca, Hinton 5805 (A, BM, F, K),
7128 (BM, K, US). Mexico: Rinc6n del Carmen, 1460 m, Hinton 1016 (K, MO, P). Michoacan:
Huetamo, Hinton 7126 (A, K, NY), 7276 (K, NY). Morales: near Cuernavaca, Rose & Hough
4372 (US). Nayarit: near Jesus Maria, Feddema 1327 (MICH); Santiago, Lamb 587 (GH, NY).
Oaxaca: Monte San Felipe, Andrieux 175 (P); Canotlan, 1700 m, Bravo s.n. (MEXU); Hac. de
Parmentiera 99

i- ~----- _^^ t^ ......... ..

FIG. 19. Map 5. Distribution of Parmentiera aculeata.

Guadeloupe, 1650 m, Conzatti 5251 (MICH,NY); cerca Tuxtepec, 30 m, Hernandez & Ramos 501
(MEXU); Chiltepec, 25 m, Hernandez 576 (F); 1-5 mi E of Temascal, Janzen 10476 (MO); 21.4 mi
S of Tuxtepec, LeDoux & Dunn 2315 (MO); Chiltepec, 20 m, Martinez-Calder6n 149 (A, US); vic.
of Mitla, 1600-1700 m, Messer 190 (MICH); vic. of Cuicatlan, 1800-2500 ft, Nelson 1683 (GH,
US); valley of Oaxaca, 5000 ft, Pringle 7524 (F, MO, US); San Mateo del Mar, 5 m, Zizumbo &
Colunga 138 (MO). Puebla: Huaytamalco, 300 m, Mdrquez 614 (F); Mesa de San Diego, 400 m,
Pennington & Sarukhan 9286 (NY); Caltepec, 1908, Purpus s.n. (BM, F, GH, MO, NY, US); 3950
(BM, F, GH, MO, NY, US); between Macozcalco & San Antonio Canada 1000-1800 m, Smith et
al. 4070 (F, US). San Luis Potosi: 1 mi S of Tamazunchale, Barkley 17M302 (F); Tamazunchale,
Edwards 663 (F, MO); Valles, Kenoyer A347 (F); 12 mi NE of Xilitla, King 4379 (F, MICH, US);
San Dieguito, Palmer 136 (GH, US); Rasc6n, Pringle 5297 (GH); Quinta Chilla, Vaughn et al.
1143 (MO). Sinaloa: Mocorito (cult.), Collins & Kempton 68 (US); Culiacan, Palmer 1532 (F, GH,
NY, US); Rosario, Rose 1583 (US). Tabasco: 21 km W of Cardenas, 5-15 m, Conrad et al. 2914
(MO, US); San Juan Bautista, Rovirosa 388 (K, NY). Tamaulipas: Tampico, Fisher 46116 (F); 7
mi N of El Lim6n, Lloyd & Mueller 3650 (GH); vic. of Tampico, 15 m, Palmer 80 (GH, MO, US);
vic. of Gomez Farias, 350 m, Palmer 269 (F, GH, K, MO, NY, US). Veracruz: Santa Cruz Hidal-
go, 160 m, Avendano et al. 104 (F, MO); 2 km de P. Vicente, Chavelas & Zamora s.n. (MEXU);
San Pablo, 100 m, Gutierrez 92 (MSC, WIS); Los Mangos, Hernandez 1344 (F); Tierra Blanca, 30
m, Hernandez & Orozco (MEXU); Nautla, Jarquin & Lot 41 (MEXU, US); Papantla, Liebmann
8023 (F); Pital, Liebmann 8024 (C, F); Poza Rica, 250 m, Lot 810 (GH); Acahual, Martinez-Cal-
der6n 1218 (BM, F, MICH,MO, U); Cazones, Mell s.n. (NY); Orizaba, Muller 2201 (NY);
Papantla, Schiede 1207 (BM, K, MO, US) El Triunfo, Papantla, Vela 1461 (MEXU); 2 km S de
Vigueta, Ochoa 1935 (MEXU); 28 mi N of Tantoyuca, Ozment et al. 390 (WIS); Laguna de Sonte-
compan, 60 m, Pennington & Sarukhan 9133 (NY); Zongolica, Vera Santos 2915 (MICH); Cerca
del Cerro Chico, Ventura 3597 (MICH). Yucatan: Izamal, Gaumer & Sons 23440 (F, GH, MO,
NY, US); sin. loc., Millspaugh 30 (F); Merida, Schott 242 (F).
100 Flora Neotropica

GUATEMALA.Alta Verapaz:Cajab6nto ChurnjigaOxec, Cook& Griggs681 (US);Caha-

b6n, Goll 148(US); Panzos,Pittier353 (NY, US);cultivatedfromseedsfrom Rio Salinas,Steyer-
mark45716(F). Guatemala:Chinautla,4000ft, DonnellSmith2691(GH, K, NY, P, US); Pinula,
Hayess.n. (GH);EstanciaGrande,600 m, Standley59295(F). Peten:N of El Cruce,500 ft, A.
Gentry8324(F, MO);Yaxha-Remate Rd, Lundell2079(F, MICH);LagunaPacay,200 m, Molina
15737(F, NY); Retalhuleu:9 km N of Champerico,40 m, Harmon2293 (MO);vic. Los Delicias,
200 m, Standley88041(F);vic. Retelhuleu,240 m, Standley88538(F). San Marcos:betweenOcos
& Ayutla, 5-20 m, Steyermark37927 (F). Santa Rosa: Santa Rosa, 3000 ft, Heyde & Lux 3109
(GH, K, NY, P, US).
BELIZE.Sin. loc.: Winzerling1119(US). Belize:2 mi W of BelizeRiveron NorthernHwy,
Liesner& Dwyer1443(MO);RoaringCreek,Lundell340 (F). Cayo:Cocquericot,Bartlett12037
(MICH,NY, F); nearVaca, Gentle2433(A, F, MICH);5 mi S of Georgeville,A. Gentry7762(C,
F, MO, NY); WesternHighwaybetweenBananaBankand WarMemorial,Proctor30354(BM);
2.5 mi S of jct. of Western& HummingbirdHwys, 75 m, Spellman& Newey 1872(MO).Orange
Walk:IndianChurch,Arnason& Lambert17045(MO);Toledo: Rio GrandeRiver,Sampson25
(K),Schipp1146(A, BM, F, GH, K, MICH,MO, NY).
EL SALVADOR.La Union:vic. of La Union, 150m, Standley21056(GH, NY, US). San
Salvador:San Salvador,Calder6n353 (GH, NY, US); Renson317 (NY, US). SantaAna: Santa
Ana (cult.), Allen & van Severen7137(US). Sonsonate:vic. of Izalco, 400-600m, Pittier 1935
(US);vic. of Nahulingo,220m, Standley22010(US).
HONDURAS.Cortes:La Lima, Johansen30 (F); vic. of La Lima, 30 m, Standley& Cal-
der6n7551(F). Valle:Amapala,10m, 1945, Valerios.n. (F).
CULTIVATED:Australia(Queensland:Brass33502 (K); White12847(A). Escapedfrom
cultivation),Bogor,EastAfrica,GoldCoast,Cuba,PuertoRico, Tobago.

Local name. Mexico: San Luis Potosi: chayote, chote. Puebla: chote,
cuajilote. Guerrero: cuajilote, cuajiote. Oaxaca: cuajilote. Veracruz: chote,
chotes, guachilote, cuajilote. Guatemala: guajilote, cuajilote. Belize: cow
okra, cat, kat. El Salvador: Cuajilote.

Williams (1973) has argued that Seemann did not formally propose a
combination Parmentiera aculeata based on Crescentia aculeata HBK., since
he cited the name as based on a Schiede collection from Hooker's herbarium
"distributed under the name of Crescentia aculeata. " Even though Seemann
did not specifically state in 1854 that his Parmentiera aculeata was based on C.
aculeata HBK., he clearly intended this as indicated in his 1860 revision of
Crescentieae where C. aculeata HBK., and P. aculeata Seem. are listed to-
gether under P. edulis. The combination has long been, and should continue to
be, attributed to Seemann.

2. Parmentiera cereifera Seemann in J. Bot. (Hooker) 3: 302. 1841.

Figs. 20, 21.

Small tree to 20 cm dbh. and 7 m tall, usually several-branched near the

base, the main branches strongly ascending, the crown open; twigs terete, sub-
puberulous, the nodes often with a pair of small leaves in the axil of a larger
pair, foliaceous pseudostipules sometimes present, 6-10 mm long and 3-4 mm
wide. Leaves opposite, 3-foliolate; leaflets elliptic to elliptic-rhomboid, acute
to acuminate, basally attenuate, the terminal leaflet 3.6-9.5 cm long and 1.4-4
cm wide, the lateral leaflets 3.0-6.5 cm long and 1.4-3.6 cm wide, membrana-
ceous, secondary veins in 4-6 pairs, somewhat lepidote at least beneath,
simple-puberulous along the midvein above and beneath, especially in axils of
Parmentiera 101

"' i: >/ cjjH

FErf a mC

FIG. 20. Parmentiera cereifera and P. macrophylla. A-E, P. cereifera; A, inflorescence x 1/2; B,
ovary x.s. x 9; C, fruit x 3/5; D, fruit x.s. x 1/2; E, leaves x 1/2 (Gentry 5610). F-J, P.
macrophylla; F, leaves x 1/2; G, corolla x 1/2; H, ovary and disc x 3/4; I, calyx x 1/2; J, fruit
x.s. x 1/2 (Gentry 754). (From Flora of Panama).
102 Flora Neotropica

lateral nerves, usually with a few plate-shaped glands along the mid-vein be-
neath, drying olive to blackish; petiolules not differentiated, petiole conspicu-
ously winged, 2.4-6.2 cm long, the wing to 0.4 cm wide, widest apically,
basally attenuate, the midrib slightly puberulous. Inflorescence a single cauli-
florous flower or a fasicle of 2-3 flowers borne on abbreviated short-shoots on
larger branches or trunk, the pedicels lepidote, 0.9-1.6 cm long. Flowers with
the calyx spathaceously split on the ventral side, 1.9-4.9 cm long, 0.9-2.9 cm
wide, inconspicuously lepidote, at least at the base; corolla pure white, tubular
with a horizontal pleat midway across the lower side of throat, slightly fleshy,
3.7-6.4 cm long and 1.8-2.9 cm wide at the mouth, the tube 2.5-4.0 cm long,
the lobes basally fused, 1-1.1 cm long, mostly glabrous throughout, the epi-
dermal cells apparently glandular, a few stalked-lepidote trichomes at level of
stamen insertion inside and on lobes outside; stamens subexserted, the anther
thecae thick, partially divergent, 5-8 mm long, the anterior filaments 2.6-3.5
cm long, inserted 1.1-1.8 cm from base of corolla tube, the posterior filaments
2.7-3.2 cm long, inserted 1.2-1.7 cm from base of the corolla tube, the stami-
node 1-1.9 cm long, inserted 9-23 mm from base of corolla tube; pistil 3.1-4.8
cm long, the ovary linear-cylindrical, 7-8 mm long, 1.5-2 mm wide, glabrous
with glandular epidermis, the ovules irregularly 8-10-seriate; disc annular-
pulvinate, 1.5-2 mm long, 6 mm wide. Fruit elongate-linear, subterete in cross
section, 39-54 cm long, 1-2.4 cm wide, glabrous to sparsely lepidote, waxy yel-
low, divided into a fleshy part surrounding the seeds and a fibrous central core
surrounded by the seeds; seeds small, thin, 3-4 mm long, 3-4 mm wide, with a
narrow but distinct mucilaginous wing.
Type. Panama. PanamA: Seemann s.n. (holotype, K).
Distribution. Apparently restricted to limestone outcrops on the Atlantic
slope of the Canal Zone and adjacent central Panama, where it is locally com-
mon. It is widely cultivated as a curiosity throughout the tropics.

PANAMA.CanalZone:SummitGarden(cult.), Correaet al. 1689(PMA);nearSalamanca

Hydrographic Station,Dodgeet al. 16963(MO);Boy ScoutroadnearMaddenLake,Gentry2053,
2488, 5060(all MO);Barbacoa,Hayess.n. (BM);Matachin,Hayes 141 (GH), 604 (NY);Madden
Dam,Kirkbride&Elias267 (NY, SCZ);vic. of MaddenDam, Lewiset al. 21 (GH, MO, US); vic.
of Gatuncillo,Piper5611,5641, 6009(allUS);betweenGorgonaandMamei,Pittier2237(BM, C,
NY, US); Rio Chagres,Seemanns.n. (K);ChilibreRiverbelowChilibre,Seibert1503(MO, US);
Rio PedroMiguel,Standley29979(US);vic. of Fort Sherman,Standley31132(MO, US); Mamei
and Gorgona, Wagoner35 (M); MaddenDam, Woodsonet al. 1554(A, MO, NY); vic. of Sala-
mancaHydrographicStation, Woodsonet al. 1607(A, MO, NY); sin. loc., Gaillards.n. (US),
Gaillards.n. (NY, US);Johansen16 (US).
CULTIVATED:Dominica, Puerto Rico, Cuba, Costa Rica, Florida, Camerouns,and

Local name. Palo de velas, candle tree.

3. Parmentiera macrophylla Standley, Publ. Field Mus. Nat. Hist., Bot. Ser.
4: 263. 1929. Fig. 20, 21.
P. macrophyllaStandley,Trop.Woods16: 12. 1928,nomennudum.

Small to medium-sized tree to 30 cm dbh. and 25 m tall, the trunk usually

Parmentiera 103

erect and unbranchedfor much of its length, the crown open; twigs terete,
glabrousto slightlyand inconspicuouslypuberulous.Leavesmostly opposite,
sometimessubalternate,often ? fasciculate,3-foliolate (very rarelywith an
extrabasal leaflet), the leaflets rhomboid-elliptic,acuminateto acute, basally
attenuate,the terminalleaflet 6.8-20 cm long, 2.6-5.9 cm wide, the lateralleaf-
lets 4.7-17 cm long, 1.8-5.5 cm wide, membranaceous,secondaryveins 5-7 on
a side, somewhatlepidoteor deciduous-lepidote,especiallybeneath,otherwise
glabrousexceptfor a ciliatefringeof shortsimpletrichomesalong the margins
of the well developed axillary domatia beneath, usually with a few plate-
shapedglands along the midveinbeneath,dryingdark olive above, pale olive
beneath;petioluleswingedbut more or less differentiatedfrom the decurrent
leaflet bases, the terminalpetioluleto 3.5 cm long, the lateralpetiolulesto 1.1
cm long, the petiole unwingedexcept on immatureplants, furrowedabove,
glabrous, 3.6-7.2 cm long. Inflorescenceof one or 2 flowers borne along the
trunk,the pedicelslepidote.Flowerswith the calyx spathaceouslysplit, 2.6-5.0
cm long, 0.8-1.2 cm wide, basallylepidote, otherwisemostly glabrouswith a
conspicuouslyglandularepidermiswhen fresh;corolla purewhite with a faint
greenish tinge outside, tubular-campanulatewith an inconspicuous saccate
bulgein the floor of the throat, fleshy, 4.9-6.3 cm long, 1.5-2.5 cm wide at the


FIG. 21. Map 6. Distribution of Parmentiera. P. cereifera (open squares), P. macrophylla (tri-
angles), P. millspaughiana (closed stars), P. morii (star in circle), P. parviflora (closed circles), P.
stenocarpa (crosses), P. trunciflora (open circles), P. valerii (open stars).
104 Flora Neotropica

mouth, the tube 3.6-4.5 cm long, the 3 lower lobes 0.8-1 cm long, the upper 2
fused half their length, glabrous throughout except for a few glandular-lepi-
dote enations at the level of stamen insertion, the epidermis conspicuously
glandular; stamens subexserted, the anther thecae thick, 1.5-2 mm wide, par-
tially divergent, 5-6 mm long, the anterior filaments 3.3-3.8 cm long, inserted
1.5-1.7 cm from the base of the tube, the posterior filaments 3.3-3.8 cm long,
inserted 1.7-2.1 cm from the base of the tube, the staminode 4-16 mm long,
sometimes totally fused to the corolla tube; pistil 5.7-5.9 cm long, the ovary
narrowly cylindric, octagonal in cross section, 6-7 mm long, 2 mm wide, glab-
rous with a glandular epidermis, the ovules ca. 6-seriate in each locule; disc
annular-pulvinate, 1.5 mm long, 5 mm wide. Fruit cylindric with conspicuous
longitudinal ribs, suboctagonal in cross section, 44-50 cm long, ca. 6 cm
wide when fresh and 2.8 cm wide when dried, waxy yellow when fresh, drying
black, divided into a fleshy part outside the seeds and a fibrous core inside the
seeds; seeds small, thin, 3-6 cm long, 8-9 cm wide, with a distinct, narrow
mucilaginous wing when fresh.
Type. Panama. Bocas del Toro: region of Almirante, Cooper 402 (holo-
type, F; isotypes, K, US).
Distribution. Tropical wet forest and premontane rain forest, mostly
above 500 m, in Costa Rica and the western half of Panama.
COSTARICA.Alajuela:VillaQuesada,A. Smith1798(F, MO, US). Lim6n:Talamanca,in
forestsof Shirores,Tonduz&Pittier997 (US);Monteverde,Stork894 (F).
PANAMA. Bocasdel Toro: ChanguinolaValley,Dunlap421 (F); aboveAlmirante,Gentry
2802, 2815 (both MO);vic. of ChiriquiLagoon, von Wedel1101(GH, MO);WaterValley, von
Wedel1421 (GH, MO, US); Darkland,von Wedel2619 (GH, MO, NY, US). Cocl6:vic. of El
Valle,800-1000m, Allen 1782(F, GH, MO, NY, US), Croat14312(F), Duke 12174(MO),Dwyer
et al. 4550 (MO),Folsom3898 (MO),Gentry754(F), 757(F), Gentry& Dwyer3663 (MO),3658
(MO),Hunter&Allen307 (MO),Kirkbride1089(MO),Lewiset al. 1780(MO),Luteyn1214(GH,

One problematical collection which may belong to this species is not in-
cluded in the above description. This is Dressier 3679 (F, MO) from Cerro
Jefe, Panama Province, Panama. It is described as cauliflorous and the leaves
are identical with those of P. macrophylla but the single preserved flower is
much smaller (calyx 1.7 cm long, pedicel 0.9 cm long, corolla 2.5 cm long, 2
cm wide at mouth of tube) than in other material of P. macrophylla.

4. Parmentiera millspaughiana L. O. Williams, Fieldiana, Bot. 36: 28. 1973.

Fig. 21.
Shrub 3-5 m tall, branchlets terete, the nodes armed with a conspicuous
thorn to 4 mm long subtending each leaf. Leaves opposite to subopposite or
with 1 or 2 smaller simple leaves (= pseudostipules?) from the same axil as a 3-
foliolate one, subrotund to obovate, rounded at apex, terminal leaflet 0.5-3.5
cm long (without the usually long-attenuate base), 0.4-2.5 cm wide, lateral
leaflets 0.3-3 cm long, 0.2-2 cm wide, scattered lepidote, nerve axils below con-
spicuously barbate and more or less domatiate, usually slightly puberulous at
Parmentiera 105

base of midvein above, sometimesvery sparselypuberulousover upper sur-

face; petiolulespoorly differentiatedfrom leafletbases, petiole 0.5-4 cm long,
narrowlywinged. Inflorescencea single terminalflower, the pedicelthickish,
less than 1 cm long. Flowerswith the calyx spathaceous,ratherobtuse, acute
in bud, 1.8-2 cm long, corolla greenish,campanulate,3.5-4 cm long, 1.3-1.5
cm wide at mouth of tube, the lobes more or less fused, stamensexserted,the
anther thecae thickish, somewhat divergent, 4-5 mm long; ovary not
examined.Fruit linear, 16-24cm long, 0.5-1 cm wide, the surfacesmooth or
finelystriate,not ridged;seedsminute,unlobed,3 mm long, ca. 2 mm wide.
Type. Mexico. Yucatan:sin. loc., Gaumer338 (holotype, F; isotypes, A,
BM, C, K, MO, NY, UPS, US).
Distribution.Dry thorn scrubforests of the YucatanPeninsula,disjunct
in ColimaStateon the Pacific coast of Mexico.
MEXICO.Colima:km. 293, 15mi SE of Manzanillo,ca. 0 m, McVaughet al. 1626(MICH,
MO);oeste Rio Armeria,Miranda9118(MEXU);Manzanillo,Palmer1347(GH, U, US); 12 km
E of Manzanillo,10-50m, West3508 (GH). Campeche:Chan Laguna,Lundell1021 (F, MO,
MICH, NY, US). Yucatan:Ticul, Enriquez309 (MEXU);Kinchil,Enriquez821 (MEXU, US);
San Anselmo, Gaumer1613 (F, GH); Suitun, Gaumer23289 (BM, F, GH, MO, US), 23439(F,
GH, MO, NY);km. 27 Meridaroad,Lundell&Lundell8198(MICH);San Ignacio,Miranda8226
(MEXU,US);Merida,Schott382 (BM,F, MO,US), 382a(F).

Localname. (Maya)kat-puc,catpxiu,catcunc,xcatcunk,xcatecuk.

5. ParmentieramoriiA. Gentry,sp. nov. Figs. 21, 22.

Arborparva,ramulislenticellisverrucatis.Foliolaangusteelliptica,acuta;petiolusnon ala-
tus. Floresterminales,calyceacuminato,corollatubulari-campanulata, tubo non conspicuecur-
vato. Fructuslinearisvel lineari-oblongus,8-angulatus.

Smalltree ca. 5 m tall; branchletsterete, unarmed,conspicuouslywarty-

lenticellate.Leavesopposite, 3-foliolate,the leafletsnarrowlyelliptic,acuteat
base and apex, entire,terminalleaflet 3.5-12 cm long, 1.3-4.3 cm wide, lateral
leaflets 1.6-8 cm long, 0.8-3.3 cm wide, lepidote, especiallybelow, nerveaxils
below usuallydomatiatewith a fringeof trichomesarounddomatia,otherwise
glabrous;petiolulespoorlydifferentiated,less than 1 cm long, petiole0.5-5 cm
long, unwingedbut the top grooved with slightly thickened lateral ridges.
Inflorescence1 or 2 terminalflowers,the pedicels1-1.5cm long, sparselylepi-
dote. Flowerswith the calyx spathaceous,mostly glabrous, inconspicuously
lepidote near base, acuminate,ca. 3.5 cm long, includingthe ca. 3 mm long
acumen; corolla greenish-whiteto creamy-yellow,evenly tubular-campanu-
late, the tube without a noticeablebulge or "S"-shapedbend, 5-5.5 cm long,
ca. 2.5 cm wide at mouth of tube, the broadlobes ca. 1 cm long, mostlyglab-
rous, very sparsely glandular-lepidoteon lobes and upper half of tube;
stamenssubexserted,the antherthecae divergent,ca. 4 mm long; ovary not
examined;disc patelliform, 1 mm long, 5 mm wide. Fruit linear to linear-
oblong, octagonal,taperingto base and apex, 11-30cm long, 1.7-2.2 cm wide,
with 8 conspicuouslongitudinalridges when dried, seeds 4-5 mm long, 6-7
mm wide, with a subhyalinemarginalfringe representinga distinctvestigial
106 Flora Neotropica

FIG. 22. Parmentiera morii and P. stenocarpa. a-B, P. morii; A, habit (Mori 7727); B, fruit (Mori
& Kallunki 2259). C-E, P. stenocarpa (Gentry 9435); C, corolla, ventral view; D, corolla, side
view; E, calyx. (All x 1/2).
Parmentiera 107

Type. Panama. Panami: 8-11 km from Pan Am. Hwy., 300-400 m, Mori
7727(holotype, MO; isotype, US).
Distribution. Known only from the El Llano-Carti Road wet tropical for-
est in central Panama.
PANAMA. Panama: El Llano-Carti Road, 10 km from Pan Am. Hwy., Mori & Kallunki
2259 (MO), 9.4 km from Pan Am. Hwy., 350 m, Mori & Kallunki 6399 (MO).

This species is very closely related to P. stenocarpa of northwestern

Colombia. It differs from P. stenocarpa in a distinctly 8-angled longitudinally
striate-ridged fruit, more prominent leaf venation beneath, a longer less con-
spicuously apiculate calyx, a more gradually broadening corolla with the tube
not at all "S"-shaped, and more suberous warty-lenticellate twigs. It is possi-
ble that these differences would break down if Parmentiera is discovered in the
geographically intermediate Darien region, but the available evidence supports
recognition of two species.

6. Parmentiera parviflora Lundell, Lloydia 3: 211. 1940. Fig. 21.

Shrub to 4 m tall and 8 cm dbh., branchlets terete, the nodes armed with a
thick thorn subtending each leaf on young branches, these usually reduced and
often not evident on older branches. Leaves opposite to subopposite, mostly 3-
foliolate, a few simple, the leaflets obovate to elliptic, acute to obtuse, the base
acute to attenuate, usually in part obtusely few-dentate, terminal leaflet and
simple leaves 2-7 cm long, 1.3-3 cm wide, lateral leaflets 1-5 cm long, 0.7-3 cm
wide, scattered lepidote, the nerve axils beneath barbate and domatiate, other-
wise glabrous; petiolules not differentiated, the petiole 1-5 cm long, narrowly
winged. Inflorescence a single terminal flower, the pedicel slender, 0.6-1 cm
long. Flowers with the calyx subspathaceous, usually 2-parted at apex, the
lobes rather obtuse, 0.8-1.1 cm long, corolla greenish, more or less tubular,
1.7-2.0 cm long, 0.4-0.5 cm wide at mouth of tube, the lobes more or less
fused, 3-4 mm long, scattered glandular lepidote outside and on lobes inside;
stamens subexserted, the anther thecae slightly divergent, 2 mm long; ovary
linear-oblong, 2 mm long, 1 mm wide, essentially glabrous; disc pulvinate, 1
mm long, 2 mm wide. Fruit fusiform-oblong, acute at base and apex, (2.3-)4-
5.5 cm long, (0.8-)1-1.3 cm wide, not ridged, the surface smooth or with
several prominulous longitudinal lines; seeds thin, wingless, conspicuously
bilobed, ca. 1 cm wide from lobe tip to lobe tip, each lobe 5-6 mm long.
Type. Mexico. Tabasco: Reforma, Balancan, Matuda 3210 (holotype,
MICH; isotypes, A, F, US).
Distribution. Endemic to the Peten region of northern Guatemala and
adjacent Mexico, below 100 m alt.
MEXICO. Tabasco: Reforma, Balanc,n, Matuda 3209 (A, F, MICH); Laguna de la
P61vora, Rovirosa 171 (K, US).
GUATEMALA. Pet6n: Rio Pasi6n below Sayaxche, mouth of Rio Pucte, Lundell 17709
(US); Rio Pasi6n above Altar de Sacrificios, Lundell 17896 (US); Rio Cancuen between El Cambio
and mouth of Rio Machaquila, 75 m, Steyermark 45894 (A, F, MICH, NY); Rio Santa Monica
108 FloraNeotropica

between Cedral and Ceibal, 50-60 m, Steyermark46027 (A, F); Santa Elena, camino para
Sayaxch6km. 70, Tun-Ortiz835(F, MO, NY).

Localname. crucetillo.

7. ParmentierastenocarpaDugand& L. B. Smith, Caldasia7(31): 17. 1955.

Figs. 21, 22.
Small shrubbytree 4-6 m tall, to 8 cm dbh.; branchletsterete, unarmed,
slightlylenticellate.Leaves opposite, 3-foliolate, the leaflets ovate-ellipticto
obovate, usually conspicuouslyacuminate,cuneate at base, entire, terminal
leaflet 5-12 cm long, 1.7-5 cm wide, lateralleaflets 2.5-8.5 cm long, 1.3-4 cm
wide, lepidote,especiallybelow, nerveaxilsbelow usuallydomatiateand with
a fringeof trichomesaroundthe domatia;petiolulesnot clearlydifferentiated,
petiole 1.2-4 cm long, unwingedbut the top flattenedand with slightlythick-
ened lateralridges.Inflorescenceone or two terminalflowers, the pedicels1-2
cm long, lepidote, long acuminate(occasionallybilabiateat apex with each
lobe long acuminate),2.7-3 cm long, includingthe 5 mm long acumen,in bud
with the acumenalmostas long as the pedicel;corollawhite, tubular-campan-
ulate, somewhat "S"-shaped in side view; 4-5 cm long, 2-2.5 cm wide at
mouthof tube, the broadlobes ca. 1.5 cm long, mostlyglabrous,verysparsely
glandularlepidoteon lobes; stamenssubexserted,the antherthecaedivergent,
5 mm long, thickish;pistil 4-5 cm long, ovary linear, lepidote, 5 mm long, 1
mm wide, disc patelliform, 1 mm long, 5 mm wide. Fruit linear, cylindrical,
taperingto base and apex, 30-40cm long, 1-2cm wide, not ridged,the surface
smooth or with barely prominulouslongitudinallines when dried; seeds 4-5
mm long, 6-7 mm wide, with a subhyalinemarginalfringerepresentinga dis-
Type. Colombia.Antioquia:Nicocli, 5 m, Haught4919 (holotype,COL;
Distribution.Endemicto the lowlandwet forest of northernChoc6 and
COLOMBIA.Antioquia:Rio Le6n, 15km W of Chigorado,100m, Feddema1857(MICH,
NY, US). Choc6: Rio Truand6,Gentry9435 (COL,F, GH, MO);Rio Truand6,6 km arribade
confluenciadel Rio Chintad6,Romero-Castarleda
6100(COL,MO, NY); confluenciade los Rios
Chintad6y Truand6,Romero-Castafleda 4682 (COL);45 km from Atrato Riverin "cativales",

Localnames.Palo de vela, plantanillo.

Uses. Fruitsaidto be edible.
8. ParmentieratruncifloraStandley& L. Williams,Ceiba3: 219. 1953.
Fig. 21.
Tree 6-9 m tall, to 20 cm dbh, branchletsterete, the nodes thickenedbe-
low the leaves, almost subspinescent.Leaves(in juvenilecondition)opposite,
3-foliolate,the leaflets obovate-elliptic,acutishat apex, cuneateat base, to 5
x 2 cm, membranaceous,lepidote,with trichome-lineddomatiain nerveaxils
Parmentiera 109

beneath,petiolulesnot well-differentiated,petioleca. 3 cm long. Inflorescence

cauliflorousnear base of trunk, with 2-5 buds clusteredat apex of a bracteate
woody short-shoot1-1.5 cm long, the pedicels(in bud) 1-1.5 cm long, densely
lepidote. Flowersnot known. Calyx in bud to 2 cm long, obtuse to acutish,
Type. Nicaragua.Jinotega:SierraW of Jinotega, 1050-1400m, Standley
Distribution.Knownonly fromthe type localityand environs.
NICARAGUA. Jinotega: Sierra W of Jinotega, 1050-1400 m, Standley 9732 (F), 10227 (F),
10239 (F).

The speciesis veryincompletelyknownand not verydifferentfrom Costa

Rican P. valerii except in its 3-foliolate ratherthan palmately 5-7-foliolate

9. ParmentieravaleriiStandley,J. Wash.Acad. Sci. 17: 16. 1927. Fig. 21.

Tree 12-13 m tall, 40-45 cm dbh.; twigs terete, noticeably lenticellate.
Leaves3 or 4 togetherin suboppositefascicles,usuallyon 2-5 mm long short
shoots, palmately4-5(-7)-foliolate,the leafletsobovate, acute, cuneateat base,
the terminalto 8 cm long and 3.2 cm wide, others smaller,the smallest2 cm
long and 1.3 cm wide, slightlylepidote, pubescentin axils beneath;petiolules
0.5-1.2 cm long, not well differentiatedfrom leaf base, the petiole 1.5-7 cm
long, petioleand petiolulesindistinctlylepidote. Inflorescenceof 1-2cauliflor-
ous flowers borne on lower part of trunk, the pedicels 1.9-2.5 cm long, scat-
tered-lepidote.Flowerswith the calyx spathaceous,blunt-tipped,3.9-4.1 cm
long, 1.5-1.8 cm wide, scattered-lepidote;corolla greenish-white,6.5-7.5 cm
long, 1.8-2.9 cm wide at mouth of tube, infundibuliformabove a 2.5-3 cm
long and 0.8-1.2 cm wide tubularbase, mostly glabrous,somewhatscattered-
lepidotewithout, at least towardapex;stamensabout equal, the antherthecae
divergentto subdivaricate,4.5-5 mm long, 1.5 mm wide, the filaments2.5-2.8
cm long, inserted2.8-2.9 cm from base of tube, the staminode12-13mm long,
inserted very high on the corolla tube; pistil 4.8-4.4 cm long, the ovary
tapered-cylindrical,5-6 mm long, 2-3 mm wide, apparentlyglabrous but
rough-surfaced,the ovulesca. 4-seriatein each locule; disc annular-pulvinate,
2-3 mm long, 8-10 mm wide. Fruitcylindrical,sometimescurved,smooth, 20-
31 cm long, ca. 4 cm wide, whendryonly 2.7-2.8 cm wide, somewhatlepidote,
blackwith brownishspotting;seedsthin, essentiallywingless,4-6 mm long, 7-
8 mm wide.
Type. Costa Rica. Guanacaste:NaranjosAgrios, near Tilaran,Standley
& Valerio 46411 (holotype, US).
Distribution.Endemicto the middle slopes of the northernCosta Rican
volcanosaround700 m alt.
COSTA RICA. Alajuela: Zapote de Upala, Volcan Miravalles, Poveda & Campos s.n. (CR).
Guanacaste: Volcan Orosi, 17 km S of Nicaragua border, premontane wet forest, 675 m (reported
but not collected by Holdridge et al. (1972)).
110 Flora Neotropica

Local name. Pepino de danta.

II. Bignoniaceae tribe Tourrettieae K. Schumann in Engler & Prantl, Nat.

Pflanzenf. 4(3b): 250. 1894.

Herbaceous vines; branchlets angled, nodes without interpetiolar glandu-

lar fields; pseudostipules lacking. Leaves opposite, 2-3-ternate, often with a
trifid, dichotomously branching tendril, the leaflets serrate and often more or
less subdivided. Inflorescence a terminal spike-like raceme, the upper flowers
sterile and caducous. Flowers with the calyx red, bilobed, caducous; corolla
tubular, bilabiate, shorter than the calyx, pubescent at least toward the tip;
stamens 4, staminode absent, the anthers glabrous; pollen grains single, 3-
colpate, the exine almost psilate; ovary ovoid, densely short-echinate; disc
annular. Fruit an uncompressed, ovoid, septicidally dehiscing capsule, usually
not splitting to the base, densely echinate with uncate spines; seeds small, thin,
with a narrow, hyaline-membranaceous wing.
Type genus. Tourrettia Fougeroux.
One species of upland areas of continental tropical America from south-
ern Mexico to Argentina; also in the Peruvian lomas.


4. Tourrettia Fougeroux, Mem. Acad. Paris 1784: 205. 1787; corr. Schreber,
Gen. 2: 406. 1791, orth. et nom. cons.
Dombeya L'Htritier, Stirp. Nov. 33. 1786, nom. rej. Type: D. lappacea L'Heritier = T.
lappacea (L'HWritier)Willdenow.
Medica Cothenius, Disp. Veg. 7. 1790; non Miller, 1754. nom. nov. for Dombeya L'H6ritier.

Description as for the tribe and species.

Type species. T. lappacea (L'Her.) Willd.

1. Tourrettia lappacea (L'Heritier) Willdenow in Linnaeus, Sp. PI., ed. 4. 3:

263. 1801. Figs. 23, 24.
Dombeya lappacea L'H6ritier, Stirp. Nov. 33, pl. 17. 1785. Type: Dombey 391, Peru (holo-
type, P; isotypes, FI, P).
Tourrettia volubilis J. F. Gmelin, Syst. Nat. 2: 940. 1791. Based on the same as the

Herbaceous vine, the branchlets tetragonal. Leaves opposite, biternate

with a subterminal, finely branched, spirally coiling tendril, the leaflets mem-
branaceous, glabrous, conspicuously and unevenly serrate, 3-8 cm long, 1.5-5
cm wide, the lateral pair of a ternation often divided to base. Inflorescence
spicate but with short pedicels, narrowly bracteate, the bracts scarlet, the ter-
minal flowers mostly sterile with bright red calyces. Flowers with the calyx of
Tourrettia 111



l~. .

23. Tourrettia
lappacea. B, fruits. (All
A, habit; x 3/5. From Flora ofPanama)
112 Flora Neotropica



FIG. 24. Map 7. Distribution of Tourrettia.

fertile flowers 1.3-1.5 cm long, somewhathirsute,deeply 2-partedmore than

half its length, the upperlobe long-acuminate,the lower lobe acute, the limbs
red;corollapurplishto greenish,tubular,1.6-1.8cm long and apically3-5 mm
wide, bilabiate,the tube 1-1.1 cm long, the lip 6-8 mm long, all lobes fused,
forminga protectivehood over the anthers,this open below, pubescentout-
side, mostly glabrousinside, pubescentat the base of the stamens;stamens4,
the antherthecaedivaricate,glabrous, 1-2 mm long, shorterfilaments8-9 mm
long, longerfilaments9-11 mm long, staminodeabsent,inserted7-8 mm from
the base of the tube; pistil 1.4-1.5 cm long, the ovary ovoid, densely short-
tuberculatewith thick, single-celledtrichomes, 2.5-3 mm long and 1.5 mm
wide, 4-celled, the ovules uniseriatein each locule; disc cupular-membrana-
ceous, 1 mm long and 1.5 mm wide. Capsuleovoid, somewhatwoody, ca. 3.5
1.7 cm,
x 1.7 cm, strongly echinatewith
stronglyechinate with long to 11 cm
long spines cm long with recurved
long with recurvedtips
Tourrettia 113

and shorterstraightspines, 2-valved,apicallysepticidallydehiscent,often not

splittingto base; seeds thin, flat, 6-7 mm long, 4-5 mm wide, the darkbrown
body encircledby a narrowbrownalmostmembranaceouswing.
Type. Peru. Sin. loc., Dombey391 (holotype,P; isotypes,FI, P).
Distribution.SouthernMexicoto northernArgentina,at intermediateele-
vations in the Andes and the CentralAmericancordilleras.Mostly between
1500 and 2500 m, also occurringat much lower altitudes in the Peruvian
lomas. AbsentfromeasternSouthAmerica.
MEXICO.Sin. loc.: Hartweg547 (P). Chiapas:BarrancaHonda, Siltepec,Matuda4044
(MICH);Mt. Tacana,Matuda2434 (MICH);FincaPrusia,Munic.AngelAlbinoCorzo, 2400 ft,
Ton3618(DS, MICH).
GUATEMALA.Chimaltenango:1800m, Morales1241(F). Guatemala:nearFincala Au-
rora, 1500m, Aguilar202 (F); slopes of Lago de Amatitlanbelow Moran, 1300m, Steyermark
52159(F). Huehuetenango:3 km SWof SanAntonioHuista, 10km ESEof La Democracia,1500
m, Iltis & Lind G-153(F); Jacaltenango,3500-5400ft, Nelson 3585 (F). Quezaltenango:Finca
Pirineos,below SantaMariade Jesus, 1500-1650m, Standley84585(F); betweenFincaPirineos
and Patzulin,1200-1400m, Standley86700(F);FincaPirineos,betweenSantaMariade Jesis and
Calahuache,1300-1500m, Steyermark33245(F). Retalhuleu:nearChivolandiaalongroadto San
Felipe,650 m, Standley87183(F). Sacatep6quez: BarrancoaboveDuefias,1590-1800m, Standley
63240(F). San Marcos:Rio Ixpil, below Rodeo, 750 m, Standley68749(F); FincaEl Porvenir,
3500 ft, S. White5269 (MICH);betweenFincaEl Porvenirand Loma Corona,9 mi NW of El
Porvenir,1300-2000m, Steyermark37711(F). Suchitepequez: nearPuebloNuevo, 750 m, Stand-
ley 66892(F); lower slopes of VolcanZunil, SE of SantaMariade Jesfis, 1200-1300m, Steyer-
mark 35233 (F); Volcan Santa Clara, 1.5-2 mi W of Finca El Narajo, 1250 m, Steyermark
HONDURAS.El Paraiso:Cafetalesde MontafiaApauhis,cercade Danli,Molina7452(F).
NICARAGUA.Matagalpa:nearXelaju, 1500m, Williamset al. 29227(BM,F, MO).
COSTARICA. Alajuela:San Ram6n,Brenes23140 (NY); La Verbena,Pittier9111 (CR,
US); Guadelupe,SmithP2190(GH);ZarceroRegion,Smith4140 (MO), 1948/23(US). Cartago:
Cartago,Cooper6000 (GK, K, US), Torres129 (US); El MuflecoS of Navarro,Standley33708
(US); La Estrella,Standley39186 (US); Tucurrique,Tonduz12971(BM, M, P, US); Cartago,
Werckles.n. (HNCR 11573)(CR, US). Heredia:Rio Virilla,Biolley 1963(CR, US); entreCari-
blanco and VaraBlanca,Jimenez2047 (NY); VaraBlanca,Kupper291 (M); road to Sarapiqui,
1500m, Lent3247 (MO).San Jose:ParqueBolivar,Brenes65 (CR);Montede la Carpintera,Pit-
tier4395(CR);Rio Torresa SanFranciscode Guadalupe,Pittier9071(CR, US); CiudadUniversi-
taria, Sdenz1076(CRU);vic. of El General,Skutch2420 (K, MO, NY, US); vic. of SantaMaria
de Dota, Standley41854 (US), Standley& Valerio44081(US); La Hondura,Standley& Valerio
51090(US); Rio Torresa San Franciscode Guadelupe,Tonduz7097(HNCR9071) (GH, K, M,
NY, US), 9774(CR, US); foretsdu Copey, Tonduz11738(CR,US).
PANAMA.Chiriqui:trail from Paso Anchoto MonteLirio,Allen 1477(GH, MO);E side
of CerroPando, D'Arcy 6637 (C, K, MO, P, UCWI);CerroPunta, Graham275 (GH); vic. of
Boquete,Lewiset al. 372 (GH, MO, US);Maurice725(US);Pittier7001(US);Rio ChiriquiViejo
valleynearEl Volcan, White23 (GH, MO); Finca Leridato Boquete, Woodsonet al. 1116(A,
MO, NY); Woodson&Schery237(GH, MO,US).
COLOMBIA.Sin. loc.: Tenasuca,Trianas.n. (P). Antioquia:Boquer6n,withoutcollector,
Jan 1928(MEDEL);trail to Parquede los Orchideas,1600-1800m, Gentryet al. 24599 (COL,
MO). Cesar:Sierrade PerijA,San Antonio, 1700 m, Cuatrecasas& Romero-Castarieda 25335
(COL);aboveManaure,800 m, Haught3950 (COL).Magdalena:SierraNevadade SantaMarta,
10?58'N, 73058'W, 1270 m, Rio Frio, Kirkbride1978 (COL);Quebradala Sirena, 10?59'N,
73059'W, 1100m, Kirkbride2176 (COL). Putumayo:Valle de Sibundoy,2200 m, Cuatrecasas
11668(COL).Tolima:QuindiuRoad, 2600m, Alston 7750(BM);Ibague,cuestade Tolima, 1844,
Goudot s.n. (P); Mariquita,Linden 4135 (P), 1800 m, Triana4111 (BM); Monte Quindio,
Anaime,1900m, Uribe-Uribe3260(COL).
VENEZUELA.Barinas:Sto. Domingo-Barinitas, Aristeguieta4924 (VEN). DistritoFede-
ral: entre El Lim6n y El Aguacatal,750 m, Pittier 9239 (US, VEN); Finca Jeremba,carretera
Tovar-Pto.La Cruz, 1800-2000m, Trujillo5310(MO,MY). Merida:entreCrucehaciaEjidoy La
114 Flora Neotropica

Chorrera,1100m, Benitezde Rojas 1495(MY);de Portachuelohacia El Cacique,950-1100m,

Bernardi3176(VEN);entreChiguaray La Azulita, 1300m, Velasco& Ramia451 (MY), 1600m,
522 (MY). Yaracuy:camino de Aroa, 1300m, Trujillo& Ferndndez781 (MY). Zulia:cerca la
fronteraColombiana,Sierrade Periji, 1160-1440m, Steyermarket al. 105738(MO,VEN).
ECUADOR.Chimborazo:vic. of Huigra,Rose & Rose 22416(NY, US). El Oro: between
SantaRosaand La Chorita,0-1000m, Hitchcock21125(NY, US); vic. of La Chonta,Rose et al.
23477(NY, US). Loja:betweenLojaandSanLucas,2100-2600m, Hitchcock21442(US). Pichin-
cha: El Volante,roadChiriboga-Santo Domingode los Colorados,Asplund19048(S); Tandapi,
alt. 1500m, Heilborn748(S, US);Mindo, 1950m, Heinrichs584 (NY);aboveTandapi,2300-2450
m, Holm-Nielsenet al. 7093(MO).Tungurahua:Hda. Rio VerdeGrande,valleyof Rio Pastaza,
Asplund7867(S); valleyof Rio PastazabetweenMachaiand La Victoria,1400m, Asplund8521
(FI, NY, S, US); valley of Rio PastazabetweenBafiosand Cashurco,1300-1800m, Hitchcock
21772(NY, US);vic. of Rio Margaritas,1225m, Penland&Summers154(MO);nearBaflos,1800
m, Rimbach714 (S). Sin. loc.: Andre 2747 (P); Hda. La Merced,Benoist 4176 (P); km. 25,
Benoist4474(P); Spruce5102(BM).
PERU. Cajamarca:Cumbil-Catache,Prov. Santa Cruz, 500 m, Lopez & Sagastegui5171
(TRUJ);El Bosque, Vargas& Novoa 2223 (TRUJ);San Miguel, Weberbauer 5246 (F). La Liber-
tad: CerroCampana,600 m, Angulo 2021 (MO, TRUJ), 675 m, Lopez 216 (MO, USM), 718
(TRUJ).Lima:Lomasde Atocongo,500-600m, Ferreyra1535(MO,USM),250 m, Ferreyra3904
(USM),500 m, Ferreyra4018(USM),Muelleret al. 931b(LZ);Amancaes,Ferreyra4047(USM);
Lachay,600-650m, Ferreyra3881(USM),400m, Ferreyra11847(USM);nearLima,Martinet367
(P); Dist. Lurigancho,Campoy, 330 m, Mexia 4035 (MO);Amancaes,Ridoutt 12171(USM);
12172(MO,USM);Lomasde Asia, Ridoutt13033(MO,USM);8 km E of Pachacamac,Saunders
173 (BM);CerroJeronimo,Soukup1550(USM).Piura:arribade Palo Blanco, 500 m, Lopezet
al. 7691(TRUJ).Sin. loc.: Dombey874 (P); Ruiz&Pavons.n. (BM).
BOLIVIA.Tarija:Emboruzu,rutaBermejo-Tarija,117km S of Tarija,Krapovickaset al.
ARGENTINA.Salta:CerroAstillero,Pierotti1362(LIL);Oran,Tres Palcas, Willink267
(LIL);San Ignacio,Willink298 (LIL).

Local names. Guatemala: espina de cuchumb de montafia, guisquilillo,



This revision is based on the study of herbarium specimens from the

following herbaria: A, AAU, BM, C, CEPEC, COL, CR, F, FI, G, GH,
whose curatorsI thank for making their collections available.
Various phases of this study have been supported both directly and in-
directly by a series of grants from the National Science Foundation.


Aublet,J. B. C. F. 1775.Histoiredes plantesde la Guianefrancaise.2. Paris.

Baillon,H. 1882.Crescentianigripes.Rev. Hort. 54:464-466.
. 1888 (1891). Histoire des plantes 10: 1-58. Paris.
Baker,H. & B. Harris.1959.Bat-pollinationof the silk-cottontree, Ceibapentandra(L.) Gaertn.
s.l. in Ghana.J. W. Afr. Sc. Ass. 5:1-9.
Bentham,G. &J. D. Hooker. 1876.Generaplantarum2: 1026-1053.London.
Bojer,W. 1837.Hortusmauritianus.Mauritius.
Literature Cited 115

Brown, R. 1810. Prodromus florae Novae Hollandiae. London.

Bultman, J. D. & C. R. Southwell. 1976. Natural resistance of tropical American woods to terres-
trial wood-destroying organisms. Biotropica 8: 71-95.
Bureau, E. 1864. Monographie des Bignoniacees. Paris.
. 1865. Description du genre nouveau Schizopsis. Adansonia 5: 369-381.
. 1868a. Revision des genres Tynanthus et Lundia. Adansonia 8:270-294.
. 1868b. Description du genre nouveau Saldanhaea. Adansonia 8: 353-358.
& K. Schumann. 1896-1897. Bignoniaceae. In: Martius, Flora Brasiliensis 8(2).
Buurman, J. 1978. Contribution to the pollen morphology of the Bignoniaceae, with special ref-
erence to the tricolpate type. Pollen et Spores 19:447-519.
Candolle, A. P. de. 1838. Revue sommaire de la famille des Bignoniac6es. Bibl. Univers.
Geneve 1-24.
. 1845. Prodromus systematis naturalis regni vegetabilis. 9:142-248. Paris.
Chamisso, A. de. 1832. De plantis in expeditione Romanzoffiana. Linnaea 7: 653-723.
Correa, P. 1926. Diccionario das plantas uteis do Brasil. Rio de Janeiro.
D'Arcy, W. D. 1979. Scrophulariaceae. In: Flora of Panama. Ann. Missouri Bot. Gard. 66: 173-
Dobbins, D. R. 1971. Studies of the anomalous cambial activity in Doxantha unguis-cati (Bignoni-
aceae) II. A case of differential production of secondary tissues. Amer. J. Bot. 58: 697-705.
Don, G. 1838. A general history of the dichlamydous plants. 4: 214-233. London.
Dop, P. 1925. Contribucion a l'6tude des Bignoniacees. Bull. Soc. Bot. France 72: 887-891.
. 1929. La Classification des Bignoniac6es Asiatiques et Africaines. Bull. 5oc. Hist.
Nat. Toulouse 58: 641-644.
Dugand, A. 1942. Dos nuevas Bignoniaceas del valle del Magdalena. Caldasia 1(5): 29-35.
Endlicher, S. L. 1836-1840. Genera plantarum. Vienna.
Fabris, H. A. 1965. Bignoniaceae. In: Flora Argentina. Revista Mus. La Plata, Secc. Bot., nov.
ser., 9: 273-419.
Gentry, A. H. 1969. Tabebuia (Bignoniaceae): the tortuous history of a generic name. Taxon 18:
. 1973a. Generic delimitations of Central American Bignoniaceae. Brittonia 25: 226-
_ 1973b. Studies in Bignoniaceae VII. Dendrosicus, Enallagma, and Amphitecna.
Taxon 22: 637-640.
. 1973c. Bignoniaceae. In: Flora of Panama. Ann. Missouri Bot. Gard. 60: 781-997.
. 1974a. Coevolutionary patterns in Central American Bignoniaceae. Ann. Missouri
Bot. Gard. 61: 728-769.
_ 1974b. Flowering phenology and diversity in tropical Bignoniaceae. Biotropica 6:
_ 1976a. Bignoniaceae of southern Central America: Distribution and ecological
specificity. Biotropica 8:117-131.
. 1976b. Relationships of the Madagascar Bignoniaceae: A striking case of conver-
gent evolution. Plant Systematics and Evolution 126: 255-266.
. 1976c. Amphitecna-Enallagma-Dendrosicus revisited. Taxon 25:108.
. 1977a. Endangered plant species and habitats of Ecuador and Amazonian Peru.
pp. 136-149. In: G. Prance & T. Elias, eds., Extinction is forever. New York Bot. Gard.
. 1977b. Notes on Middle American Bignoniaceae. Rhodora 79:430-444.
. 1977c. Bignoniaceae. In: Flora of Ecuador 7: 1-172.
. 1978a. Bignoniaceae. In: Botany of the Guayana Highland. Mem. New York Bot.
Gard. 29: 245-283.
. 1978b. Extinction and conservation of plant species in tropical America: a phyto-
geographical perspective. pp. 115-131. In: I. Hedberg, ed., Systematic botany, plant utiliza-
tion, and biosphere conservation.
. 1979a. Distribution patterns of neotropical Bignoniaceae: some phytogeographical
implications, pp. 339-354. In: K. Larsen & L. Holm-Nielsen, eds., Tropical Botany, New
116 Flora Neotropica

1979b. Bignoniaceae. In: Flora de Venezuela. (in press).

& A. S. Tomb. 1979. Taxonomic implications of Bignoniaceae palynology. Ann.
MissouriBot. Gard.66: 756-777.
Goldblatt,P. 1976.New or noteworthychromosomerecordsin the Angiosperms.Ann. Missouri
Bot. Gard.63: 889-895.
& A. Gentry. 1979. Cytology of Bignoniaceae. Bot. Not. 132: 475-482.
Gomes,J. C., Jr. 1955.Contribuicaoa sistematicadas BignoniaceaeBrasileiras.Arq. Serv.Flor-
est. 9: 261-296.
. 1957. Bignoniaceae. Flora do Itatiaia. Rodriguesia 22: 111-127.
Gould,E. 1978.Foragingbehaviorof Malaysiannectar-feeding
bats. Biotropica10:184-193.
Graebener,L. 1899.CrescentiamacrophyllaSeem.Gartenwelt3: 568-569.
Griggs,R. F. 1904.A vegetablemimic.Theplantworld7: 196-198.
Grisebach,A. H. R. 1866.Catalogusplantarumcubensium.Leipzig.
. 1874. Plantae lorentzianae. Abh. Konigl. Ges. Wiss. G6ttingen 19:1-231.
Harborne,J. B. 1967.Comparativebiochemistryof the flavonoidsVI. Flavonoidpatternsin the
Bignoniaceaeand Gesneriaceae.Phytochemistry1966:1643-1651.
Hemsley,W. 1882.Botany.In: Godman& Salvin,Biologiacentrali-americana. 2. London.
Holdridge,L., W. Grenke,W. Hatheway,T. Liang,&J. A. Tosi, Jr. 1971.Forestenvironments
in tropicallife zones,a pilotstudy.Oxford.
Humboldt,A. & A. Bonpland. 1808. Plantae aequinoctiales1: 107-110.Also transcriptionof
Humboldt'snotesat Kew.
____ , ,__ & C. Kunth. 1819. Nova genera et species plantarum 3: 132-159.
Jacquin,N. J. von. 1760.Enumeratiosystematicaplantarum.Leiden.
. 1763. Selectarum stirpium americanarum historia. Vienna.
. 1797. Plantarum rariorum horti caesarei schoenbrunnensis. Vienna.
. 1800-1809. Fragmenta botanico. Vienna.
Janzen,D. H. 1971.Euglossinebees as long-distancepollinatorsof tropicalplants. Science171:
Jussieu,A. L. de. 1789.Generaplantarum.Paris.
Kuhlmann,J. G. 1941.Uma Bignoniaceaepoucoconhecida.Rodriguesia5: 365-366.
Kunth,C. 1818.Revisionde la familledes Bignoniacees.J. Phys. Chim. Hist. Nat. Arts 87: 444-
Leon, Bro. &Bro. Alain. 1957.Bignoniaceae.Florade Cuba4:414-446.
Leinfellner,W. 1973.Das Gynozeumder BignoniaceenII. Die U-formigeplazentavon Schlegelia
Oesterr.Bot. Z. 121:13-22.
Linden,J. J. 1863.Cataloguedes plantesexotiques18:8.
Lindley,J. 1847.Thevegetablekingdom.London.
Linnaeus,C. 1753.Speciesplanatarum,ed. 1, 2. Stockholm.
. 1754. Genera plantarum, ed. 5. Stockholm.
Macbride,J. F. 1961. Bignoniaceae.In: Flora of Peru. Publ. Field Mus. Nat. Hist., Bot. Ser.
Meisner,C. F. 1840,1841.Plantarumvasculariumgenera.Leipzig.
Menninger,E. 1970.Floweringvinesof the world.NewYork.
Miers,J. 1863. Reporton the plantscollectedby Mr. Weir, especiallythe Bignoniaceae.Proc.
Roy. Hort. Soc. London.3:179-202.
Miquel,F. A. G. 1844.Symbolaead floramsurinamensem. Linnaea18:241-271.
. 1850. Stirpes surinamenses selectae. Leiden.
Ogura,M. G., A. Cordell,& N. R. Farnsworth.1977.PotentialanticanceragentsIV. Constitu-
entsof JacarandacaucanaPitt. (Bignoniaceae).Lloydia40:157-168.
Paclt,J. 1952.Synopsisof the genusCatalpa(Bignoniaceae)III. Candollea13:241-285.
Pichon,M. 1945.Notes surles Bignoniacees.Bull.Soc. Bot. France92: 222-227.
. 1946. Sur le centre de dispersion des Bignoniac6es. Bull. Soc. Bot. France 93: 121-
Prance,G. T. 1977.Theethnobotanyof the PaumariIndians.Econ. Bot. 31: 129-139.
Rafinesque,C. S. 1838.Sylvatelluriana.Philadelphia.
Raven, P. 1975. The bases of Angiospermphylogeny:cytology. Ann. MissouriBot. Gard. 62:
Literature Cited 117

& D. Axelrod. 1974. Angiosperm biogeography and past continental movements.

Ann. MissouriBot. Gard.61: 539-673.
Record,S. & R. Hess. 1940.Americantimbersof the familyBignoniaceae.Trop. Woods63: 9-38.
& C. Mell. 1924. Timbers of tropical America. New Haven.
Rohrhofer,J. 1931.Morphologischestudienan den Staminodiender Bignoniaceae.Oester.Bot.
Z. 80: 1-30.
Sandwith,N.Y. 1938.Bignoniaceae.In: Pulle, Floraof Surinam4(2): 1-86.Amsterdam.
. 1954. Bignoniaceae. In: Flora of Trinidad and Tobago 2: 316-354. Port-of-Spain.
. 1968. Contributions to the flora of tropical America LXXVI: Notes on Bignonia-
ceae XXIV: Arrabidaeain Martius's"Flora Brasiliensis"and subsequently.Kew Bull. 22:
_& D. Hunt. 1974. Bignoniaceas. In: Flora ilustrada catarinense. BIGN: 1-172.
Sauer,C. 0. 1969.TheearlySpanishMain. Berkeley.
Schlechtendal,D. F. L. von. 1833.De plantismexicanis.Linnaea8: 513-528.
. 1853. De plantis variis mexicanis. Linnaea 26: 365-384.
& A. de Chamisso. 1830. Plantarum mexicanorum a cel. viris Schiede et Deppe col-
lectarum.Linnaea5: 72-174.
ScottElliot,G. F. 1890.Ornithophilousflowersin SouthAfrica.Ann. Bot. (London)4: 267-280.
Schumann,K. 1894. Bignoniaceae.In: Engler& Prantl,Die natiirlichenPflanzenfamilien4(3b):
Seemann,B. 1851.Sketchof the vegetationof the Isthmusof Panama.J. Bot. (Hooker)3: 300-
. 1854. Botany of the voyage of the H. M. S. Herald. London.
. 1860. Synopsis crescentiacearum: an enumeration of all the Crescentiaceous plants
at presentknown.Trans.Linn.Soc. London23: 1-22.
. 1863. Revision of the natural order Bignoniaceae. J. Bot. (Seem.), 1: 18-23.
Seibert,R. J. 1940.Botanyof the Mayaarea:MiscellaneouspapersXXI: The Bignoniaceaeof the
Mayaarea.Publ. CarnegieInst. Wash.522:375-434.
_. 1948. The use of glands in a taxonomic consideration of the family Bignoniaceae.
Ann. MissouriBot. Gard.35:123-136.
Southwell,C. R. & J. D. Bultman.1971.Marineborerresistanceof untreatedwoods over long
periodsof immersionin tropicalwaters.Biotropica3: 81-107.
Splitgerber,F. L. 1842.Observationesde Bignoniaceissurinamensibus.
Physiol.9: 5-16.
Spruce,R. 1859.On five newplantsfromeasternPeru.J. Linn.Soc., Bot. 3:191-204.
Standley,P. C. 1926.Treesandshrubsof Mexico.Contr.U. S. Natl. Herb.23.
& L. O. Williams. 1974. Bignoniaceae. In: Flora of Guatemala. Fieldiana, Bot.
Steenis,C. G. G. J. van. 1927.MalayanBignoniaceae,theirtaxonomy,origin, and geographical
description.RecueilTrav.Bot. Neerl.24:787-1049.
. 1928. The Bignoniaceae of the Netherland Indies. Bull. Jard. Bot. Buitenzorg, ser.
3, 10: 173-290.
. 1929. A revision of the Queensland Bignoniaceae. Proc. Roy. Soc. Queensland 41:
.1978. Bignoniaceae. In: Flora Malesiana, ser. 1, 8(2): 114-186.
Steudel,E. G. 1831.Nomenclatorbotanicus.Stuttgart.
Tokarnia,C. H., J. Dobereiner,C. Canella& D. J. GuimarAes.1969.Intoxicacaoexperimental
por Pseudocalymmaelegans(Vell.) Kuhlm.em bovinos. PesquisasAgropec. Bras. 4: 195-
Tournefort,P. de. 1719.InstitutionsReiHerbariae,ed. 3, 1. Paris.
Urban,I. 1916.UberRankenundPollenderBignoniaceen.Ber. Deutsch.Bot. Ges. 34: 728-758.
Vahl,M. 1798.Eclogaeamericanae2. Copenhagen.
Wafer,L. 1699.A newvoyageanddescriptionof the Isthmusof America. 1967 reproduction
editedby L. E. ElliottJoyce.Lichtenstein.
Willdenow,C. L. 1802(1801).Speciesplantarum,ed. 4, 3. Berlin.
Williams,L. 0. 1973.Bignoniaceaeof tropicalNorthAmerica.Fieldiana,Bot. 36:21-29.
118 FloraNeotropica

CRESCENTIEAE 1-18.A. tuxtlensisA. Gentry
1. Amphitecna 2. Crescentia
1-1. A. apiculataA. Gentry 2-1. C.alata HBK.
1-2. A. breedloveiA. Gentry 2-2. C. amazonica Ducke
1-3. A. costataA. Gentry 2-3. C. cujeteL.
1-4. A. donnell-smithii(Sprague)L. 2-4. C. linearifoliaMiers
Williams 2-5. C. mirabilisEkmanex Urban
1-5. A. isthmica(A. Gentry)A. Gentry 2-6. C. portoricensisBritton
1-6. A. kennedyi(A. Gentry)A. Gentry 3. Parmentiera
1-7. A. latifolia(Miller)A. Gentry 3-1. P. aculeata(HBK.)Seemann
1-8. A. macrophylla(Seemann)Miers 3-2. P. cereiferaSeemann
1-9. A. megalophylla(DonnellSmith) 3-3. P. macrophylla Standley
A. Gentry 3-4. P. millspaughiana L. Williams
1-10.A. molinaeL. Williams 3-5. P. moriiA. Gentry
1-11.A. montanaL. Williams 3-6. P. parvifloraLundell
1-12.A. parvifloraA. Gentry 3-7. P. stenocarpaDugand& L. B.
1-13.A. regalis(Lindley)A. Gentry Smith
1-14.A. sessilifolia(DonnellSmith) 3-8. P. truncifloraStandley&
L. Williams L. Williams
1-15.A. silvicolaL. Williams 3-9. P. valeriiStandley
1-16.A. spathicalyx(A. Gentry)
1-17.A. steyermarkii(A. Gentry) 4. Tourrettia
A. Gentry 4-1. T. lappacea(L'Heritier)Willdenow
List of Exsiccatae 119

Abbott,W., 2202(1-7). Breedlove, D. E., 2825 (2-3), 10064 (1-15),
Acosta-Solis,M., 6024(2-3),6461(2-3). 11788 (2-1), 12746 (2-3), 21175 (1-15),
Adams,C. D., 12788(1-7). 22620 (1-11), 23526A (3-1), 24946 (1-2),
Aguilar,I., 202 (4-1). 25718 (1-11), 28886 (1-1), 29655 (1-15),
Alain, Bro., (see Liogier). 30469 (2-1), 31011 (1-11), 32819 (1-2),
Alexander,R., s.n. (2-3). 33928 (1-1), 34091 (1-1), 34432 (1-2),
Allen,P. H., 1362(1-14),1477(4-1), 1782(3-3), 34472 (1-1), 35106 (1-15), 38684 (1-2),
3632(1-7),5639(1-7). 38800 (1-15), 39067 (1-17), 40461 (1-11).
Alston,A. H. G., 5896(1-7), 7750(4-1). Brenes, A. M., 65 (4-1), 168 (1-14), 3830 (1-14),
Anderson,A., s.n. (1-7). 12125 (2-3), 12719 (2-1), 13498 (1-5),
Andre,A., 1821(2-3). 13797 (1-5), 23140 (4-1).
Andrieux,G., 175(3-1),225 (3-1). Breteler, F., 4327 (2-3).
Archer,W. A., 423 (2-3). Britton, N. L., 451 (1-7), 601 (2-4), 786 (2-3),
Aristeguieta,L., 4924(4-1),5444(1-7), 1603 (2-3), 1906 (2-4), 2205 (2-3), 2455 (2-6),
7087(2-2),7118(1-7), 7791(2-2). 2594 (2-3), 4523 (2-6), 4818 (2-4), 4869 (1-7),
Arnason,T. & J. Lambert,17023(2-4), 7029 (1-7), 8297 (2-4), 8853 (2-4), 9454 (2-3),
17045(3-1). 13240 (2-3), 14858 (1-7).
Arnoldo,Fra., s.n. (2-3),267 (2-3). Broadway, W., s.n. (2-3), 540 (2-3).
Asplund,E., 7867(4-1), 8521(4-1),9342(2-3), Bruff, J., 1320 (2-1).
9343(2-3), 19048(4-1), 19542(2-3). Brunt, M., 2110(2-3).
Aublet,J. B.C. F., s.n. (2-3). Bunting, G. S. & L. Licht, 692 (2-1).
AvendanoR., S. et al., 104(3-1). Burchell, W. J., 1058 (2-3), 9576 (2-3).
Burger, W., 6579 (1-7), 6694 (1-7), 6705 (2-3),
Baker,C. F., s.n. (1-7). 10479(1-6).
Bang,M., 1161(2-3),2536(2-3). Byrne, R., 350 (2-3).
Barclay,G., s.n. (1-7), 361(2-3), 852(1-7),
926(1-7),2768(1-7). Calderon, S., 239 (2-3), 240 (2-3), 241 (2-1,
Barkley,F. A., 17M302(3-1). 2-3), 353 (3-1).
Bartlett,H. H., 11212(2-3), 11769(1-2), Caldwell, O. H., 7059 (1-7), 8785 (1-7).
12037(3-1), 13048(2-3), 13103(2-3). Calzada, J. I., 158 (1-1), 529 (1-18), 624 (1-7),
Beaman,J., 5206(1-18). 794 (1-7), 2376 (2-3).
Beard,J. S., 468 (2-3). Carlson, M. C., 261 (2-1).
Becerra,A., s.n. (2-1). Cedillo T., R., 66 (1-1), 226 (1-1).
Belanger,C. P., 606(1-7). Chavelas, P. & C. Zamora S., s.n. (3-1).
Bena,P., s.n. (2-3). Churchill, J., s.n. (1-7).
Benitezde Rojas,C. E., 1495(4-1). Clarke, C., 2137-1 (2-1).
Benzen,A., 4172(2-3), Clemente, Bro., 2503 (1-7), 3220 (1-7),
Berlin,B., 1501(2-3), 1544(2-3), 1545(2-3), 3445 (1-7), 4868 (1-7), 6055 (2-3).
1546(2-3). Collins, G. & Doyle, 109 (2-3).
Bernal,P., 1(1-5). Collins, G. & J. Kempton, 68 (3-1).
Bernardi,L., 3176(4-1). Combs, R., 9 (3-1).
Biolley,P., 1963(4-1). Conrad, J. et al., 2914 (3-1).
Blancaneaux,1864(1-2). Conzatti, C., 8 (1-8), 794 (2-1), 3690 (2-1),
Blanchet,J. S., 295 (2-3). 3691 (2-1), 3719 (2-3), 5251 (3-1).
Blum,K. E., 2570(1-14). Cook, O. F., s.n. (2-1), 120 (1-8), 133 (2-3),
Boege,W., 456 (3-1), 1412(2-1). 158 (1-8), 256A (2-3), 575 (1-8), 681 (3-1),
Boldingh,I., s.n. (2-3), 1930b(2-3),2554b(2-4), 1683 (2-3), 7456 (2-3).
2954(2-3), 3903(2-3),6414(2-3),6416(2-3), Cooper, G. P. & G. M. Slater, 36 (1-6),
7916(2-3). 277 (1-5), 288 (2-3), 402 (3-3).
Bonbee,6369(2-3). Cooper, G. T., 6000 (4-1).
B0rgsen,F., s.n. (1-7). Correa, M. et al., 1689 (3-2).
Box, H. E., 1546(2-3). Correll, D., 45241 (2-3).
Brace,L. J. K., 4447(2-3). Croat, T. B., 273 (2-1), 13827 (1-7), 14168 (1-7),
Brass,L. J., 33502(3-1). 14312 (3-3), 14765 (1-16), 23295 (2-3),
Bravo,H. H., s.n. (3-1). 41890 (2-1).
120 Flora Neotropica

Croizat, L., 856A (2-2). Focke, H. C., 767 (2-3).

Cuatrecasas, J., 4292 (2-2), 4348 (2-3), Folsom, J., 3898 (3-3).
6429 (2-3), 11668 (4-1), 25335 (4-1). Fosberg, R., 1617 (1-7).
Curran, H.M. & Haman, 184 (2-3), 402 (2-3). Fournier, O., 1580 (2-3).
Curtis, A. H., s.n. (1-7), s.n. (1-7), 311 (2-3), Fredholm, A., s.n. (2-3), 3312 (1-7).
5838 (1-7). Freeland, J. & L. Spetzman, 23 (2-1).
Fry, Mrs. A., s.n. (2-3).
Dale, S., s.n. (2-3). Fuchs, H. P., 21702 (1-7).
D'Arcy, W. G., 6637 (4-1).
Daubenmire, R., 46 (2-3), 514 (2-3). Gaillard, Mrs. D. D., s.n. (3-2).
Davidse, G., 2411 (2-1), 9403 (2-3). Gallego, J., 539 (2-1).
Davidson, M. E., 410 (1-14). Gallegos, J. M., s.n. (2-1).
Dawe, M. T., 866 (1-7). Garber, A. P., 59 (1-7).
Deam, C. C., 6-65 (2-1). Garcia-Barriga, H., 12174 (2-3).
Delgado, E., 288 (1-7). Gardner, C. A., 3342 .(2-3).
Diguet, s.n. (2-1). Gaumer, G. F., 338 (3-4), 437 (2-3), 1613 (3-4),
Dodge, C. et al., 16963 (3-2). 23289 (3-4), 23439 (3-4), 23440 (3-1),
Dombey, s.n. (2-3), s.n. (4-1), 353 (2-3), 23773 (2-3), 23804 (2-3).
391 (4-1), 874 (4-1). Gentle, P., 6-8 (2-3), 1606 (1-7), 1785 (1-7),
Donnell-Smith, J., 2096 (2-3), 2691 (3-1). 2433 (3-1), 2605 (1-2), 3093 (1-2), 3218 (1-2),
Dorantes, J., 843 (2-1). 4651 (1-7).
Dressier, R. L., 3679 (3-3?), 3943 (1-16), Gentry, A. H., 352 (2-3), 356 (2-1), 403 (2-3),
4901 (1-5). 754 (3-3), 757 (3-3), 813 (2-3 x 2-1),
Dryander, E., 2607 (1-7). 814 (2-1), 868 (2-3), 869 (2-3), 872 (2-1),
Duchassaing, E. P., s.n. (2-3). 1001 (1-14), 1182 (2-3), 1244 (1-7), 1280 (2-3),
Ducke, A., s.n. (2-2), s.n. (2-2), 552 (2-2). 1283 (1-7), 1290 (2-3), 1291 (1-7), 1293 (2-3),
Duke, J. A., 4820 (1-7), 8552 (1-7), 9649 (1-7), 1352 (2-1), 1368 (2-3), 1405 (2-3), 1478 (2-3),
12174 (3-3). 1742 (1-7), 2044 (1-14), 2053 (3-2), 2245 (2-3),
Dugand, A., 149 (2-3). 2488 (3-2), 2523 (2-3), 2802 (3-3), 2807 (2-3),
Dunlap, V. C., 421 (3-3). 2815 (3-3), 3658 (3-3), 3663 (3-3), 3690 (2-3),
Duss, A., 1236 (2-3), 3066 (2-3). 4121 (1-7), 4397 (2-3), 4505 (2-3), 5060 (3-2),
Dwyer, J. D., 444 (1-14), 606 (1-7), 1586 (1-7), 5692 (1-14), 5769 (1-16), 7379 (1-7),
2272 (1-7), 4550 (3-3), 7694 (1-14), 7443 (1-5), 7762 (3-1), 7850 (2-3), 7851 (2-3),
12983 (1-7). 8324 (3-1), 8872 (1-5), 9232 (1-5), 9435 (3-7),
10093 (2-3), 12279 (2-3), 12696 (2-3),
Echeverria, J. A., 62 (2-3), 329 (4-1). 13167 (2-3), 13415 (1-6), 17494 (1-7),
Edwards, J. B., P-127 (2-1), P-376 (2-1), 21162(2-2).
663 (3-1). Gentry, H. S., 5043 (2-1).
Egler, F. E., 42-23 (2-3). German, M., 176 (2-1), 464 (2-1).
Eggers, H. F. A., 657 (1-7), 873 (2-3), 5292 (2-3). Ghiesbreght, A. B., s.n. (1-13).
Ekman, E. L., H6933 (2-4), H7039 (2-4), Gill, R. C., 56 (2-3).
H8822 (2-4), H 10856 (2-3), 15535 (2-5). Gillis, W., 11921 (2-3).
Elias, Bro., 141 (1-7), 1043 (2-3). Gilly, C. L., 69 (2-1).
Emrick, G. M., 141 (2-1), 156 (2-1). Glassman, S. F., 1573 (2-1).
Enriquez, O. G., 309 (3-4), 821 (3-4). Godfrey, R. K., 66377 (1-7), 66416 (2-3).
Erlanson, C. O., 4 (1-7). Goldman, E. A., 315 (2-1), 542 (2-3),
Eyerdam, W. J., 227 (2-4). 575 (3-1), 672 (2-3).
Goll, G. P., 148 (3-1).
Feddema, C., 1309 (2-1), 1327 (3-1), 1643 (2-1), Gomez-Pompa, A., 20(1-1), 5156 (1-18).
1857 (3-7), 2058 (1-7). Gonzalez O., J., s.n. (1-13), 732 (2-1).
Fendler, A., 210 (1-7), 780 (1-7), 782 (2-3). Goudot, J., s.n. (4-1).
Fernandez Perez, A., 246 (1-7). Graham, G., 275 (4-1).
Ferreira, A., 58-252 (2-2). Gregg, J., 944 (2-1), 1208 (2-1).
Ferreyra, R., 1535 (4-1), 3881 (4-1), 39-4 (4-1), Grisebach, A., 1325 (1-7).
4018 (4-1), 4-47 (4-1), 11847 (4-1). Guttierez K., L., 92 (3-1).
Ferris, R. S., 5129 (2-1), 6850 (2-3).
Fisher, G. L., 46116 (3-1). Hagen, C. von & W. von Hagen, 1295 (2-3),
Fishlock, W., 208 (2-3). 1400(2-3).
List of Exsiccatae 121

Hahn, M., 120 (1-1), 889 (2-3), 1394 (1-7), Johnston, I. M., 10(1-7), 248 (2-3), 1179(1-7).
1436 (1-7)(2-3). Jones, M. E., 34 (2-1), 22354 (2-1), 24011 (2-1).
Hammel, B., 2236 (1-13). Justin, s.n. (1-7).
Handover, W. P., s.n. (2-3).
Hardy, 302 (2-3). Kalbreyer, W., 1015 (2-3).
Harris, W., 5983 (1-7), 5986 (1-7), 6533 (2-3), Kayap, R., 1037 (2-3).
9382 (2-3). Kellerman, W. A., s.n. (2-3), 5011 (2-3),
Harlow, J., 97 (1-7). 5710 (2-1), 7050 (2-1), 7168 (1-9).
Harmon, W. E., 2293 (3-1), 5955 (2-1). Kelsall, H. J., 467 (1-7), 547 (1-7), 1258 (1-6),
Hartweg, K. T., 547 (4-1). 1261 (1-6).
Hassler, E., 12016 (2-3). Kemp, W., GC204 (2-3).
Hatheway, W. H., 1447 (2-1). Kennedy, H., 2147 (1-5).
Haught, 0, 3950 (4-1), 4919 (3-7). Kenoyer, L. A., A347 (3-1).
Hayes, S., s.n. (3-1), s.n. (3-2), 29 (2-3), Killip, E. P, 14072 (2-3), 27428 (2-3),
44 (1-7), 141 (3-2), 604 (3-2), 826 (1-7), 33012 (1-7), 33122 (1-7), 39134 (1-7),
862 (1-7). 43846 (2-3), 44827 (1-7), 45550 (1-7).
Hazlett, D., 2287 (1-11). King, R. M., 210 (2-1), 767 (2-1), 1166 (2-1),
Heilborn, O., 748 (4-1). 1465 (2-1), 4379 (3-1), 5305 (1-14).
Heinrichs, E., 351 (2-3), 584 (4-1). Kinloch, J. B., 39 (1-7), 40 (1-7).
Heller, A. & Heller, 334 (2-3). Kirkbride, J. H., Jr., 26 (3-2), 615 (1-6),
Hernandez M., R., 501 (3-1), 576 (3-1), 1089 (3-3).
1344(3-1). Klug, G., 1480 (2-3).
Heyde, E. T. & Lux, 3109 (3-1). Knobloch, I. W., 548 (2-1), 887 (2-1).
Hinds, A. J., s.n. (1-7). Krapovickas, A., 18706 (4-1).
Hinton, G. B., 755 (2-1), 1016 (3-1), 1165 (2-1), Krukoff, B. A., 1551 (2-3), 6036 (2-3).
5804(2-3 x 2-1), 5805 (3-1), 5840 (2-1), Kubicek, 3741 (2-1).
5941 (2-1), 6315 (2-1), 7111 (2-3 x 2-1). Kupper, W., 291 (4-1).
7112(2-3), 7126 (3-1), 7276 (3-1).
Hioram, Bro., 7635 (1-7). Lamb, F. H., 107 (1-2), 426 (2-1), 587 (3-1).
Hitchcock, A. S., s.n. (2-3), 2624 (1-7), Lankester, C. H., 295 (1-14), 296 (4-1).
7048 (2-1), 21125 (4-1), 21442 (4-1), Langlasse, E., 197 (2-1).
21772 (4-1). Lao, E., 250 (1-7).
Holdridge, L., 460 (2-6). Lathrop, E., 5129 (2-3).
Holt, E. & Blake, 677 (2-2). Laughlin, R., 862 (3-1).
Howard, R. A., 4976 (2-3), 14273 (2-3), Leavenworth, W. C. & H. Hoogstraal,
15664 (1-7). 1595 (2-1), 1709 (2-1).
Howell, J. H., 40 (1-7). LeDoux & Dunn, 2315 (3-1).
Hoyos, J., 2214 (1-7). LeGallo, C. J., 960 (2-4), 967 (2-4).
Humboldt, F. W. H. A. & A. Bonpland, Lent, R. W., 469 (1-7), 1231 (1-14), 2427 (1-6),
26 (2-3), 1300 (2-3), 3858 (2-1). 2548 (2-1), 3247 (4-1).
Hunnewell, F., 11580 (1-7). Le6n, J., 808 (1-14).
Hunt, D., 405 (1-2). Le6n, Bro., 13408 (1-7).
Hunter, J. & P. Allen, 307 (3-3). Leonard, E. C. & Leonard, s.n. (2-3),
12635 (1-7), 14027 (1-7), 14264 (1-7),
Iltis, H. H., G-65 (2-3), G-153 (4-1). 14411 (1-7), 15324 (2-4).
Levy, P., 1383 (2-1).
Jack, J. G., 4044 (2-3), 5014 (1-7), 5400 (2-3), Lewis, W., 21 (3-2), 372 (4-1), 1780 (3-3),
7340 (1-7). 1842 (1-7).
Jaeger, A., 88 (2-3), 207 (1-7). Liebmann, F. M., 587 (1-1), 8022 (3-1),
Jansen & Ives, 1122(2-3). 8023 (3-1), 8024 (3-1), 11348 (3-1),
Janzen, D., 10476 (3-1). 11349(2-1).
Jarzuin, E. & L. A. Lot, 41 (3-1). Liesner, R., 75 (1-5), 1443 (3-1).
Jativa, C. D. & C. Epling, 658 (2-3). Linden, J., 5a (1-1), 113 (1-7), 228 (2-1),
Jim6nez, A., 1679 (1-7), 2047 (4-1), 2879 (1-7). 1608 (1-1), 4135 (4-1).
Jimenez, J., 1991 (1-7), 2323 (2-4). Liogier, A. H., 2701 (1-7), 7137 (3-1),
Jim6nez L., O., 1192 (1-7). 6871 (2-4), 21867 (2-4).
Johansen, H., 16(3-2). Little, E. L., 6339 (1-7), 7759 (2-3), 13462 (2-3),
Johnson, H., 867 (1-15), 1106 (1-7), 1241 (1-4). 13622 (2-3), 16102 (1-7), 16402 (1-7),
122 Flora Neotropica

16463 (2-3), 20132 (2-1), 20293 (2-1), Oersted, A. S., s.n. (2-4), s.n. (2-3), s.n. (2-1),
21669 (2-4). s.n. (1-7), 1131 (1-7), 7961 (1-7), 7963 (2-4).
Lloyd, F. & Mueller, 3650 (3-1). Orcutt, C. R., 3474 (2-1).
L6pez, A., 216 (4-1). Ortega, J. G., 420 (2-1), 4533 (2-3), 6527 (2-1),
Lot, A., 810 (3-1), 1283 (1-7). 7302 (2-1).
Lundell, C. L., 340 (3-1), 1021 (3-4), 2079 (3-1), Ortiz, R., s.n. (2-3).
2325 (2-3), 3279 (2-3), 6618 (1-2), Ostenfeld, C. H., s.n. (2-3), 20(1-7), 478 (2-3).
8198 (3-4), 17709 (3-6), 17896 (3-6). Otto, E., 357 (1-7).
Luteyn, J., 1214 (3-3). Ozment et al., 390 (3-1).
Madriz V, A., AMV-37 (1-14).
Magallanes, J. A. S., s.n. (2-1). Palmer, E., 80 (3-1), 85 (2-1), 136 (3-1),
Maja, R. & S. Salas, 4-22 (2-3). 149 (2-3), 150 (2-1), 269 (3-1), 434 (2-1),
Marquez, W., 614 (3-1). 630 (2-1), 1347 (3-4), 1532 (3-1).
Marshall, M. L., 28 (2-3). Paray, L., 560 (2-1).
Martinet, J. B., 367 (4-1). Peck, M. E., 969 (1-2).
Martinez, M., 54-4 (1-1). Penland & Summers, 154 (4-1).
Martinez-Calder6n, G., 149 (3-1), 1218 (3-1). Pennell, F. W., 4707 (2-3), 10843 (2-3).
Mathias, M., s.n. (1-5), 3620 (2-3). Pennington, T. & J. Sarukhan, 9133 (3-1),
Matuda, E., 1818 (1-11), 1947 (1-11), 9149 (2-3), 9192 (2-3), 9207 (2-1), 9251 (2-1),
2434 (4-1), 3209 (3-6), 3210 (3-6), 3388 (1-1), 9286 (3-1).
4044 (4-1), 42-6 (1-11), 4491 (3-1), Perez J., L., 551 (2-1).
16643 (2-3), 16818 (2-3), 17365 (2-1), Perez-Arbalaez, E., 2475 (2-3).
17649 (1-17), 17837 (1-11), 27908 (2-1). Pierotti, 1362 (4-1).
Maurice, Bro., 725 (5-1). Pinkley, H., 227 (2-3).
McKenzie, C., 49 G-79 (1-7). Piper, C. V., 5611 (3-2), 5641 (3-2), 6009 (3-2).
McVaugh, R., 14997 (2-1), 15339 (2-1), Pittier, H., s.n. (2-3), 212 (2-1), 295 (1-9),
1626 (3-4), 21002 (2-1), 25216 (2-1). 353 (3-1), 1860 (3-1), 1935 (3-1), 2237 (3-2),
Medrano, F. G., s.n. (2-1). 2782 (1-7), 3001 (1-14), 4252 (1-7), 4362 (1-7),
Mell, C. D., s.n. (1-7), s.n. (3-1). 4395 (4-1), 5922 (1-7), 7001 (4-1), 8262 (2-3),
Menendez L., F, 532 (2-3). 9071 (4-1), 9111 (4-1), 9239 (4-1),
Messer, E., 190 (3-1). 14251 (1-7), 16004(1-7).
Mexia, Y., 948 (2-1), 4035 (4-1), 8934 (2-1). Plee, s.n. (2-3), s.n. (1-7), 660 (1-7).
Meyer, W. C., 23 (2-3). Porter, L. C., s.n. (1-7).
Miller, G. S., 1621 (1-7). Prance, G., 21001 (2-3).
Millspaugh, C. F., 30 (3-1). Pringle, G., 5293 (2-1), 5297 (3-1), 7524 (3-1).
Miranda, F., 5154 (1-2), 6916 (1-2), 8226 (3-4), Prior, A., s.n. (1-7).
9118 (3-4). Proctor, G., 10190 (2-3), 18122 (2-3),
Moldenke, H., 721 (1-7). 24516 (1-7), 30354 (3-1), 32661 (1-7),
Molina R., A., 189 (1-2), 7452 (4-1), 12181 (1-9), 36493 (1-7).
15737 (3-1), 24295 (1-10), 24299 (1-10), Purpus, C. A., s.n. (3-1), 3950 (3-1), 4319 (1-8).
25791 (2-1).
Mocquerys, s.n. (2-3). Questel, A., 201 (2-3), 231 (2-3), 868 (2-3),
Morales, R., 1241 (4-1). 1408 (2-3).
Mori, S., 2234 (1-5), 2259 (3-5), 4-94 (1-5),
4967 (1-6), 5220 (1-5), 6399 (3-5), 7727 (3-5), Raunkiaer, s.n. (1-7), s.n. (1-7), s.n. (2-4).
7980 (1-5), 9214 (2-2). Raven, P. H., 21526 (1-7).
Moritz, E., 1953 (1-7). Reeder, J. & C. Reeder, 2374 (2-1).
Morton, C., 9692 (2-3). Rehder, A., 776 (1-7), 1204 (1-7).
Muller, F., 2201 (1-8). Renson, C., 317 (3-1).
Ridoutt, C., 12171 (4-1), 12172 (4-1),
Narvaez, M. & A. Salazar, 420 (2-1), 794 (2-1). 13033 (4-1).
Nash, G., 917 (2-3). Rimbach, A., 714 (4-1).
Nee, M., 14095 (1-6). Rodriguez, L., 7 (1-7), 3263 (2-3).
Nelson, E. W., 488 (2-3), 1683 (3-1), 2572 (2-1), Roig, J., 5364 (2-5), 5662 (2-5).
3585 (4-1), 6913 (2-1). Rolfs & Quaintance, s.n. (1-7).
Novelo, A. et al., 92 (2-3). Romero-Castafleda, R., 470 (1-7), 4682 (3-2).
Rose, J. N., 1529 (2-1), 1583 (3-1), 3432 (2-3),
Ochoa G, S., 1935 (3-1). 4308 (2-3), 4372 (3-1), 8590(2-1), 12910 (2-1),
List of Exsiccatae 123

13898 (2-1), 14021 (2-1), 14466(2-1), Sousa, M., 3102 (1-7), 3298 (1-1), 4427 (1-1).
21993 (2-3), 22028 (1-7), 22078 (1-7), Sparre, B., 15360 (2-3).
22088 (2-3), 22416 (4-1), 23477 (4-1). Spellman, D. L., 1872 (3-1).
Rovirosa, J. N., 171 (3-6), 388 (3-1), 728 (1-7). Standley, P. C., 131 (2-1), 5836 (2-1),
Rowell, 17M931 (2-1). 7551 (3-1), 9364 (2-1), 9732 (3-8), 10227 (3-8),
Rugel, 663 (2-3). 10239 (3-8), 10316 (3-8), 11234(2-1),
Ruiz, H. & Pav6n, s.n. (4-1). 11317(2-1), 11525 (2-1), 17751 (2-3),
Rutten-Pekelharing, C., 380 (2-3). 17962 (2-1), 19425 (2-1), 19477 (2-3),
Rzedowski, J., 10869 (2-1), 16647 (2-1). 19497 (2-3), 19498 (2-3), 21056 (3-1),
21685 (2-1), 22010 (3-1), 22145 (2-1),
Saenz R., J. A., 1076 (4-1). 22146 (2-1 x 2-3), 24490 (2-1), 25343 (2-3),
Sagot, P., s.n. (2-3), 451 (2-3). 27885 (2-3), 29979 (3-2), 30919 (1-7),
Sagra, de la, s.n. (1-7), s.n. (2-3), 324 (2-3). 31132(3-2), 31974 (2-3), 33708 (4-1),
Salas D., S., S-375 (1-7), S-549 (2-1), 34733 (1-14), 39186 (4-1), 40138 (2-3),
S-595 (1-7), S-994 (1-7). 41854 (4-1), 44081 (4-1), 46411 (3-9),
Salazar, C. F., s.n. (2-1). 51090 (4-1), 54402 (2-3), 59295 (3-1),
Sallee, E. H., ES75 (2-3). 59607 (2-1), 60505 (2-1), 63240 (4-1),
Sampson, H. C., 25 (3-1). 64002 (2-3), 64142 (2-3), 66605 (2-3),
Samuels, J., 117 (2-3). 66892 (4-1), 68402 (4-1), 68749 (4-1),
Santos, J. V., 2915 (3-1), 3644 (1-8). 70148 (1-15), 70325 (1-15), 71591 (1-9),
Saravia, A., 2355 (2-3). 74208 (2-1), 74386 (2-1), 75020 (2-1),
Sargent, C. S., s.n. (1-7), A67 (1-7). 77959 (2-1), 78670 (2-3), 78775 (2-1),
Saunders, 173 (4-1). 79285 (2-1), 80375 (2-1), 84585 (4-1),
Schiede, C. J. W., 1207 (3-1). 86200(1-11), 86700 (4-1), 87183 (4-1),
Schipp, W. A., 645 (1-7), 734 (1-2), 867 (1-2), 87726 (2-3), 88041 (3-1), 88538 (3-1),
1146(3-1). 89986 (1-9), 90114 (1-9), 91856 (1-9),
Schnee, L., 1166 (1-7). 19881 (1-9).
Schomburgk, R., s.n. (2-3). Stange, L., s.n. (2-1).
Schott, A., s.n. (2-3), 242 (3-1), 382 (3-4), Stearn, W., C-27 (2-3), 213 (2-3), 738 (1-7),
382a (3-4), 521a (2-3). 772 (1-7), 788 (2-3).
Schunke V., J., 1396 (2-3), 9785 (2-3). Stearns, J. L., 42 (1-7).
Seemann, B. C., s.n. (3-2), 644 (1-7), 1130 (2-1). Steggerda, M., 56 (2-3).
Seiberg, R. J., 1503 (3-2), 1694 (3-7). Stehle, H., 1080 (2-3).
Sess6, M. & J. M. Mociflo, 2227 (2-1), Stern, W. L., 891 (1-7).
2228 (1-1). Stevenson, J. A., 156 (1-7), 5566 (2-3).
Shafer, J., 27 (2-3), 29 (2-3), 343 (2-3), Steyermark, J. A., 29021 (2-1), 33245 (4-1),
1249 (1-7), 1355 (2-3), 1746 (1-7), 2528 (1-7), 36704 (1-11), 36987 (1-11), 36987 (1-11),
3245 (1-7), 11839(2-3), 12135 (1-7). 37012 (1-11), 37711 (4-1), 37927 (3-1),
Shank, P. J., 4305 (1-7). 38210 (1-4), 39130 (1-3), 39199 (1-4),
Sharp, R., s.n. (2-3). 39689 (1-7), 41587 (1-2), 41796 (1-3),
Shepherd, J. D., 62 (2-3). 44780 (1-9), 45080 (1-1), 45207 (1-4),
Shimek, B. & E. Smith, 105 (2-3). 45458 (1-1), 45575 (1-4), 45716 (3-1),
Shreve, F., 6398 (2-1). 45819 (1-1), 45894 (3-6), 46027 (3-6),
Sieber, F., 277 (1-7). 46786 (4-1), 48766 (1-11), 49177 (1-9),
Simpson, J., 540 (1-7). 51252 (1-17), 52159 (4-1), 87746 (2-3),
Sinclair, s.n. (2-3). 95961 (1-7), 105738 (4-1), 106890 (1-7).
Sintenis, P., 920 (1-7), 1898 (2-4), 2740 (2-3), Stimson, W. R., 52983 (2-3), 5333 (1-7).
2980 (2-4), 5586 (1-7), 6546 (2-3). Stoffers, A., 19 (2-3), 412 (2-3), 705 (2-3),
Skutch, A. F., 2420 (4-1), 4825 (1-5), 4940 (1-5). 791 (2-3), 817 (2-3), 879 (2-3), 1023 (2-3),
Small, J., 6767 (1-7), 7872 (1-7). 1081 (2-3), 3814 (2-3).
Smith, A., A134 (1-14), 576 (1-14), 1132 (2-3), Stork, H. E., 894 (3-3), 4135 (1-14).
1570(1-14), 1798 (3-3), 1948/23 (4-1),
P2190 (4-1), 4140 (4-1). Taylor, D., 118 (2-3), 119(2-3), 120(2-3).
Smith, C. E. et al., 4070 (3-1). Taylor, R. J., 4414 (2-1).
Smith, F. D., 233 (2-3). Templeton, B. C., 7492 (2-1), 8932 (2-1).
Smith, H., s.n. (1-7), 217 (2-3). Tessman, G., 3478 (2-2).
Solis R., F., 555 (1-7). Thiebaud, M.-A., 41(2-3).
Soukup, J., 797 (2-3), 1550 (4-1). Thompson, J. B., 467 (2-3). 1062 (2-3).
124 Flora Neotropica

Thurber,G., s.n. (1-7), 1859(1-7). Werckle, C., s.n. (4-1), 17419 (1-14).
Ton, A. S., 394(2-3), 3618(4-1). West, Dr., s.n. (1-7).
Tonduz,A., s.n. (2-3),997 (3-3),7097(4-1), West, J., 3508 (3-4), 3511(2-1).
7098 (2-3), 7384 (1-14), 7937 (1-14), White, G., 23 (4-1), 1027 (2-3).
9774(4-1), 11738(4-1), 12971 (4-1), White, P., 227 (1-14).
13644(2-3). White, S., 5269 (4-1).
Toro, R., 121(2-3). Whitefoord, 1478 (2-3).
Torres,R., R., 129(4-1). Wilbur, R., 9768 (2-1), 9867 (1-14).
Triana, J., s.n. (4-1), 4111 (4-1). Wiley, J. R., 212 (2-3).
Trujillo, B., 781 (4-1), 4129 (1-7), 5310(4-1), Williams, LI., 192 (2-3), 545 (2-3), 8394 (1-1),
10839 (2-2). 9443 (1-13), 9637 (2-1), 10856 (2-1),
Tuerckheim,H. von, II221(1-4), II2278(1-9), 15274 (2-2), 15757 (2-2), 17243 (1-7).
118723(1-9), 118724(2-3). Williams, L. 0., 10856 (2-1), 26915 (1-11),
TunOrtiz,R., 835(3-6), 1459(1-4). 29227 (4-1), 40695 (1-15).
Tyson,E. L., 2738(1-7), 3565(2-3),3797(1-7), Williams, R., 1476 (2-3).
5074 (1-7). Willink, 267 (4-1), 298 (4-1).
Wilson, C. L., 274 (1-9?), 369 (1-7).
Ugalde,A., 6358(2-3). Wilson, M., 40836 (1-15).
Uribe-Uribe,L., 3260(4-1). Wilson, P., 9265 (1-7), 9331 (2-3), 11377 (2-3),
Valerio,M., s.n. (3-1),358 (2-1),463 (1-7), Wing, E. S., 66 (2-3).
464(1-7). Winzerling, H. W., 1119 (3-1).
Vazquez,B., 67 (1-1), 105(1-13). Woodson, R. E. et al., 237 (4-1), 1116 (4-1),
Vaughnet al., 1143(3-1). 1554 (3-2), 1607 (3-2).
Vela,L., 452 (2-1),461 (3-1). Woytkowski, F., 6160 (2-3), 35155 (2-3).
Velasco& Ramia,451 (4-1),522(4-1). Wright, C., s.n. (2-3), s.n. (2-1), 7 (1-7),
Ventura,F. A., 3341(1-1), 3597(3-1). 361 (1-7), 361a (1-7)(2-3).
Versteeg, G., 443 (2-3).
Xoloctzi, X334 (1-11).
Wagoner,s.n. (2-3), 35 (3-2).
Weberbauer,A., 5246(1-4). Young, J. P., 413 (1-7).
Wedel,H. von, 561(1-7),988 (1-7), 1101(3-3), Yuncker, T. G., 8724 (1-6).
1421 (3-3), 2101 (1-7), 2258 (1-6), 2619 (3-3),
2769 (1-7), 2894 (2-3). Zizumbo & Colunga, 138 (3-1).
Index of VernacularNames 125

a-jo-nocht(1-1) higuero(2-3)
ayale(2-1;2-3) hojacruz(2-1)
blackcalabash(1-7) ho/o (2-3)
calabacero(1-14) huaz(2-3)
calabacillo(1-7) huingo(2-3)
calabacillode playa(1-7)
calabasa(2-3) jicara(2-1;2-3)
calabasillo(1-7) jicarilla(1-14)
calabash(1-2; 1-7;1-14;2-3) jicarita(1-7;2-1)
calabashtree(1-7) jicarito(2-1)
calabassier(2-3) jicaro(1-7;1-14;2-1;2-3)
calbas(2-3) jicarode estero(1-7)
calebassemarron(2-4) kabami(2-3)
callebasseboite(2-3) kalabash(2-3)
callebassecoricon(2-3) kat (3-1)
candletree(3-2) luch(2-3)
cat (3-1)
catcunc(3-4) magiiira(1-7)
catpxiu(3-4) malocacao(1-1)
cat-puc(3-4) mariquitade Maria(1-7)
cerial(2-3) mate(2-3)
chayote(3-1) matecillo(1-7)
chote(3-1) matecito(1-7)
chotes(3-1) matesillo(1-7)
cirian(2-1) mora(2-3)
cirianmazo(2-3) morade costa(1-7)
cow okra(3-1) moritode rio (1-7)
crucetillo(3-6) morrito(2-1)
cruzcruzan(2-1) morro(1-11;1-17;2-1;2-3)
cuajilote(3-1) morro-cimmaron (1-13)
cueira(2-3) o-hoch(1-9)
cuiamaraca(2-2) palode velas(3-2;3-7)
cuiapequena(2-2) pati(2-3)
cuiapequenado igapo(2-2) pegapega(4-1)
cuiupi(2-2) pilche(2-3)
espinade cuchumbde montafla(4-1) platanillo(3-6)
guacalillo(1-14) rivercalabash(1-7)
guajilote(3-1) sachahuingo(2-3)
gira(2-3) savanacalabash(1-7)
guiradel monte(2-3) sirian(2-1)
guirade olor (1-7) s-xom-qawa'uul(ourLordMountain'sGourd)
guisquilillo(4-1) (1-15)
guisquilito(4-1) syrial(2-1)
higuera(2-3) tapara(2-2;2-3)
higuerillo(1-7) taparamontanera(2-2)
higuerita(2-4) taparode agua(2-2)
higuerito(2-4) tapitode monte(1-6)
126 Flora Neotropica

tecomata(2-1) tree calabash (2-3)

tecomate(2-1;2-3) tsapa (2-3)
tecomatillo(1-1) tutunuto (1-7)
totumillo(1-7) xcatcunk (3-4)
totumito(1-7) xcatecuk (3-4)
totumocimarron(2-3) wild calabash (1-2; 1-7; 2-3)
Indexof ScientificNames 127


Note: Page numbersin bold-face indicateprimarypage reference;page

numberswith ' indicatepageswith illustrationsor maps.
Adenocalymma 6, 12, 13, 14, 16, 21,24 elegans46
purpurascens 21 japurensis15
Amphicome 6 mollissima23
Amphilophium 12, 15, 16, 17, 18, 21, 22, 24, revillae39
25, 29, 33, 34 verrucosa34', 36
paniculatum 25', 29, 34' Arthrophyllum 4
Amphitecna 13, 14, 15, 17, 19, 20, 21, 22, 24, Aspidospermum 4
26, 30, 32, 35, 39, 43, 59, 50-82 Astianthus5, 6, 13, 14, 19
apiculata 51, 52, 53, 54', 55, 56', 58, 81
breedlovei 52, 56', 57', 58, 62, 74 Bignonia2, 3, 4, 6, 8, 12, 14, 29, 40, 48
costata 21, 51, 54', 56', 58, 59', 60 capreolata29
donnell-smithii 51, 54', 58, 59', 60, 61', Bombacopsis46
62, 63',76, 82 Boutonia5
isthmica 52, 61 ', 62, 64, 76 Bravaisia5
kennedyi 51, 57 ', 64, 65', 66, 75
latifolia 26', 30, 35', 39, 43, 50, 51, 56', Callichlamys14, 22, 23, 29
63', 66, 67' latifolia29
macrophylla 50, 51, 53, 55, 57', 60, 61', Campsidium8, 23, 41,42
70, 72, 76 Campsis8, 23, 28, 41,42, 45
megalophylla 51, 53, 54', 56', 60, 71, grandiflora28
72, 76 radicans28, 45
molinae 52, 59', 63', 72, 73', 74 x tagliabuena28
montana 30, 51, 54', 56', 57', 75, 82 Catalpa3, 6, 13, 18, 22, 24, 27, 28, 42
nigripes 53, 55, 70 bignonioides27, 28
oblanceolata 60, 62 x erubescens28
obovata 66 ovata28
parviflora 32, 49, 51, 54', 57', 64, 75, speciosa28
76 Catophractes6, 18
regalis 51, 53, 57', 59', 72, 76, 77 Cedrella46
sessilifolia 52, 63', 64, 73', 75, 76, 77 Ceratophytum12, 13, 23
silvicola 52, 54', 56', 63', 78, 79, 80 Cespedezia72
spathicalyx 51, 63', 65', 80, 81 Chelone3
steyermarkii 51, 53, 54', 61', 63', 80, 81 Chilopsis6, 13, 24, 28, 42, 43
tuxtlensis 52, 63', 73', 81 linearis28
Anemopaegma 9, 12, 13, 14, 15, 16, 18, 19, Clytostoma13, 18, 20, 21, 24, 26, 29, 36, 39
20, 22, 24, 29, 33, 34',39 binatum24, 29, 39
chamberlaynii 29 convolvuloides24
chrysoleucum 39 costatum24
insculptum 24 pterocalyx17, 24
orbiculatum 29, 34' sciuripabulum 24
paraense 39 uleanum24
robustum 24 Cobaea4, 7
Argylia 5, 7, 11, 14, 23, 28, 31, 42, 43 Colea 4, 7
uspallatensis 28 Couralia22
Arrabidaea9, 10, 11, 12, 13, 14, 15, 16, 18, Craniolaria3
21,22, 23, 27, 29, 34,36,39,46 Cremastus13
bilabiata 39, 46 Crescentia2, 3, 4, 13, 14, 19, 22, 26, 30, 35,
brachypoda 11 37, 43, 46, 47, 82-96
bracteolata 39 aculeata 98, 100
candicans 34' acuminata90
chica 27 alata 13, 23, 26', 47, 49, 50, 53, 83, 87'
corallina 23, 29 amazonica43, 83, 87, 88', 89', 90
egensis 15 angustifolia90
128 Flora Neotropica

arborea90 steyermarkii 80
coriacea66 Deplanchea 13
cucurbitina66 Digomphia 7, 9, 14, 17, 19, 23
cujete3, 26', 30, 35', 43, 47, 83, 87, 90, Distictella 14, 21, 22, 24
94, 95, 96 magnoliifolia 24
cuneifolia90 Distictis 6, 14, 21, 22, 24, 29
cuspidata66 buccinatoria 29
donnell-smithii60 Dolichandra 20, 23
edulis98 Dolichandrone 21, 28
elongata66 spathacea 28
fasciculata90 stipulata 28
latifolia66 Dombeya 110
lethifera66 lappacea 110
linearifolia 83, 87', 88', 94, 95, 96 Doxantha 17, 22
microcarpa94 Eccremocarpus 4, 5, 7, 14, 23, 44
mirabilis83, 87', 88', 95, 96 Ekmanianthe 24, 42
musaecarpa98 Enallagma 20, 50
nigripes53, 54 cucurbitina 50, 66
obovata66 donnell-smithii 60
palustris66 latifolia 66
plectantha90 obovata 66
portoricensis83, 87', 96 macrophylla 70
regalis76 sessilifolia 78
regia70 Esterhazya 4
ternata84 Ferdinandia 70
toxicaria66 superba 70
trifolia83 Ferdinandusa 5
Curatella77 Fernandoa 36
Cuspidaria16, 18, 19, 21, 24 Fridericia 23
Cybistax6, 23, 46 Gardnerodoxa 24
antisyphilitica 46 Gelsemium 4
Cydista9, 12, 13, 15, 20, 21,24, 29, 31, 34, Gibsoniothamnus 20, 43, 44
36 Glaziovia 17, 24, 33
aequinoctialis15, 24, 29, 31 Godmania 23, 27
decora24 Grias 72
diversifolia24, 29 Guaiacum 45
heterophylla21, 24, 34' Gustavia 72
potosina24 Haplolophium 6, 12, 17, 24
sarmentosa15 Haplophragma 15, 37
Heterophragma 19, 28
Delostoma5, 6, 13, 17, 24, 28, 31, 32, 43 adenophylla 28
lobbii28 Hieris 31
Dendrosicus50 Holoregmia 5
isthmicus62 Incarvillea 6, 11, 14, 23, 28, 31,43
kennedyi64 compacta 28
latifolius66 delavayi 28
macrophyllus70 mairei 28
molinae72 olgae 28
montanus75 semiretschenskia 28
saxatilis50, 66 sinensis 28
silvicola79 Jacaranda 3, 7, 9, 14, 17, 18, 19, 22, 23, 27,
spathicalyx80 28, 31,42, 44
Index of ScientificNames 129

caucana27 cuspidata 21
hesperia28 Oroxylum 6, 13, 15, 18, 29, 31, 37
mimosifolia28, 44, 45 indicum 29
Otophora 84
Kigelia15, 17, 20, 22, 30, 37 paradoxa 84
africana30 Oxera 5
Kuhlia7 Pachyptera 6, 10, 13, 14, 18, 21, 22, 23, 24,
Leucocalanthe13, 23 alliacea 23, 24
Levya15 hymenaea 27, 30
nicaraguense15 parvifolia 14
Lundia7, 15, 16, 17, 18, 19, 20, 22, 23, 29 standleyi 27, 35'
corymbifera29 Pajanelia 28
densiflora15 longifolia 28
stricta15 Pandorea 8, 28, 31, 45
jasminoides 28, 45
Macfadyena6, 12, 14, 17, 21, 22, 23, 29, 31, Parabignonia 20, 23
39 Paradolichandra 23
uncata39 Paragonia 6, 10, 13, 23, 27, 30, 33
unguis-cati29, 31 pyramidata 6, 30
Macranthisiphon 22, 24, 25, 40 Paratecoma 23, 45, 46
longiflorus25' peroba 45, 46
Manaosella14, 23 Parmentiera 7, 12, 15, 17, 19, 21, 22, 23, 26,
Mansoa12, 13, 16, 21, 24, 27, 29 30, 46, 47, 49, 97-110
difficilis29 aculeata 17, 30, 46, 49, 97, 98, 100
Markhamia21, 28 alata 84
hildebrandtii28 cereifera 30, 47, 97, 100, 101'
platycalyx28 edulis 97, 98, 100
Martinella22, 24, 29, 35, 36, 46 foliolosa 98
obovata24, 29, 35', 46 lanceolata 98
Martynia3, 5 macrophylla 21, 26', 30, 98, 101', 102,
Medica110 103', 104
Melloa12, 17,20, 21,22, 23 millspaughii 97, 103', 104
Memora9, 12, 14, 16, 20, 21, 24, 39 morii 97, 103', 105, 106'
cristicalyx24 parviflora 97, 103', 107
flavida24 stenocarpa 97, 103', 106', 107, 108
imperatoris-maximilianii 24 trunciflora 98, 103', 108
patula30 valerii 97, 103', 109
pedunculata24 Paulownia 7, 44
schomburgkii39 Pedalium 3
tanaeciicarpa20 Peltospermum 4
Metternichia5 Perianthomega 12, 15, 20, 24, 25, 42
Millingtonia3, 29, 31 vellozoi
hortensis29 Periarrabidaea 23
Mussatia12, 13, 18,23 Periblema 5
Perichlaena 42
Neojobertia24, 25 Petastoma 13
candolleana25' Phryganocydia 17, 20, 22, 24, 30, 36, 39
Neotuerckheimia 71 corymbosa 30
megalophylla71 phellosperma 39
gonoclada77 uliginosa 39
Nestoria9 Phyllarthron 4, 13, 30
Neves-armondia19 comorense 30
Newbouldia28 Phylloctenium 12
laevis28 Piriadacus 21, 23
Nyctocalos18, 21, 31 Pithecoctenium 12, 14, 16, 18, 19, 21, 22, 24,
130 Flora Neotropica

30, 33, 34 Tabebuia6, 8, 11, 14, 15, 16, 18, 19, 22, 23,
crucigerum30, 34' 27, 28, 32, 42, 45, 46, 94
Platycarpum4, 5 aquatilis 39
Pleonotoma12, 13, 22, 23, 30 billbergii 45
jasminifolium30 cassinoides 39
Podranea45 chrysantha 28, 45
ricasoliana45 guayacan 28, 45
Potomoganos23, 40 heptaphylla 29
Pseudocalymma10, 22, 27 heterophylla 29, 45
Pseudocatalpa18,23, 40 impetiginosa 29, 46
Pseudopaegma9, 13 obtusifolia 39
Psilogyne4, 5 palustris 39
Pteromischus83 rosea 29, 42, 45
alatus83, 84 serratifolia 29, 45, 46
Pyrostegia14,22, 23, 24, 30, 35, 36, 45 sessilifolia 77
cinerea30 Tanaecium 4, 6, 17, 21, 22, 23, 26, 27, 30, 35,
dichotoma24 36, 39, 46
venusta23, 30, 45 apiculatum 23, 24
crucigerum 39
Radermachera 28 excitiosum 46
xylocarpa28 jaroba 17, 30, 39
Rhigozum4, 6, 18 nocturnum 23, 24, 35', 46
Rhodocolea18 Tecoma 3, 6, 8, 14, 23, 29, 31,42, 43
Roentgenia9, 23, 24 garrocha 29
Romeroa24 sambucifolia 29
Roseodendron16 x smithii 29
stans 29
Saldanhaea7, 16, 18 Tecomanthe 29, 31, 42
Saritaea24, 30, 40, 45 dendrophila 29
magnifica30, 45 speciosa 29
Schizopsis7 Tecomaria 29, 31, 36, 42, 43, 45
Schlegelia6, 7, 11, 20, 23, 24, 30, 43, 44 capensis 29, 45
parviflora30 Theobroma 72
Schrebera4, 7 cacao 72
Sesamum3 Tourrettia 4, 5, 7, 8, 14, 23, 26, 30, 36, 44,
Setilobus23 110-114
Sparattosperma 6, 22, 24, 25 lappacea 26', 30, 44, 110, 111 ', 112'
leucanthum25' volubilis 110
Spathicalyx23 Trigonocarpus 5
Spathodea6, 17, 28, 36, 37, 45 Tynanthus 7, 12, 17, 19, 20, 21,27, 33, 34, 46
campanulata28, 45 croatianus 34'
Stenolobium8 Urbanolophium 24
Stereospermum 5, 22, 23, 28
chelonoides28 Vitex 5
personatum28 Wightia 4
Stizophyllum10, 12, 14, 18, 23, 30
perforatum30 Xylophragma 23, 24, 30
riparium18, 30 seemannianum 24, 30
Synapsis24, 43, 44 Zeyheria 23