Flora Neotrópica - Trigoniaceae

Organization for Flora Neotropica
Trigoniaceae
Author(s): Eduardo Lleras
Source: Flora Neotropica, Vol. 19, Trigoniaceae (Apr. 28, 1978), pp. 1-73
Published by: New York Botanical Garden Press on behalf of Organization for Flora
Neotropica
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FLORA NEOTROPIC
MONOGRAPHNO. 19
TRIGONIACEAE
by
Eduardo Lleras
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/ / C f
~TROPIC 0O CANCRt
FLORA
NEOTROPICAI
TrlOPIC OF CAPRICORN
Published for
Organization for Flora Neotropica
by
The New York Botanical Garden
New York
1978
Issued 28 April 1978

Copyright( 19 78
The New York Botanical Garden
Publishedby
The New York BotanicalGarden
Bronx, New York 10458
Libraryof CongressCatalogCardNumber 77-91706

InternationalStandardSerialNumber 00 71-5 794
InternationalStandardBook Number0-8932 7-1 98-5
Lleras, Eduardo, 1945-

Trigoniaceae.
(Flora neotropica; monograph no. 19)

Bibliography: p.
Includes index.
1. Trigoniaceae. 2. Euphronia. 3. Botany--
Classification. 4. Botany--Tropics. I. Title.
II. Series.
QA495.T73L53 583' .143 77-91706
ISBN 0-89327-198-5
All material subject to this copyright may be photocopied for the non-commercial purpose
of scientific or educational advancement.
MONOGRAPHOF THE FAMILYTRIGONIACEAEI
EDUARDOLLERAS2
INTRODUCTION
The presentstudy includes all of the known Trigoniaceae,both neotropical

and paleotropical.Althoughthe family is primarilyneotropical,it is representedby
two monotypic generafound in Malaysia(Trigoniastrum)and Madagascar (Humber-
todendron).The neotropicalgenus, Trigoniais representedby 24 species of treelets,
shrubs,or scandentshrubs,and rangesfrom the southernmostportion of Mexico to
the south of Braziland easter Paraguay.
As will be seen in the section on taxonomic history, no comprehensivetreat-
ment has been undertakenfor the family in the past one hundredyears;Warming's
treatmentfor Flora Brasiliensiswas publishedin 1875. In his monographon the
Trigoniaceae,Warmingrecognizedonly two genera,one of which is here returned
to the Vochysiaceae.Since 1875, two new generahave been recognizedas belonging
in the family, and severalnew specieshave been published.
This paperwas undertakenin an attempt to delimit the family, and to deter-
mine taxonomicallythe numberof species known at present.AlthoughWarming
had a good workingknowledgeof the group,the smallamount of materialavailable
at the time of his publicationled to errorsthat can now be corrected.His incomplete
knowledgeof type materialhas also led to a duplicationof names for taxa that are
morphologicallyindistinguishable.This treatmentis a preliminarysurvey of the
Trigoniaceaeand is necessarilythe first step in any comprehensivestudy of this
family.
The taxonomic position of the genusEuphroniahas been reconsidered.
TAXONOMICHISTORY
In his Histoire des Plantes (1775), Aublet describedthe genus Trigoniain

whichhe includedtwo species, Trigoniavillosaand T. laevis. He placed these species
in DecandriaMonogynia(10 stamens, 1 ovary)based on the Linnaeansystem of
classification.Lamarck(1786) erroneouslydescribedanother species of Trigonia
as Crotoneriospermum,based on fruit characters.
De Jussieu(1789) placed TrigonianearMalpighiain the Malpighiaceae.The
only subsequenttreatmentsworthy of note until 1824 were the publicationsof
two illegitimate generic names for Trigonia:Hoeffnagelia,published by Necker
(1790) in ElementaBotanica, and Sprengel'spublicationof Nuttallia in 1821.
De Candolle(1824) in his treatmentfor the Prodromusplaced Trigoniain
the Hippocrateaceae.This position was accepted by Cambess6desin 1829. In 1825
Martiusand ZuccariniproposedEuphronia,a genus which was placed in Kunth's
family Spiraeaceae;this assignmentwas ignored by later authors, or at least by
Based in part on a dissertation submitted to the Graduate Faculty in Biology in partial

fulfillment of the requirements tor the degree of Doctor of Philosophy, The City University of
New York, 1975.
2 Instituto Nacional de
Pesquisas da Amazonia, Caixa Postal 478, 69000 Manaus,
Amazonas, Brazil.
2 Flora Neotropica
those who have undertakensystematictreatmentsof the Trigoniaceae.Vellozo, in

1825, publishedthe last superfluousname for Trigonia,Mainea.
The first applicationof the familialname, Trigoniaceae,was by Martius
(1835), who publishedthe name without a descriptionin his Conspectus.The name
was taken up, and againpublishedwithout a descriptionby Endlicherin 1840 in
his GeneraPlantarum,and was validatedthe following'yearwhen he provideda
descriptionof the family in his EnchiridionBotanicum.
Robert Schomburgk(1847) publishedthe genusLightia,based on material
from his own collections. Lightiaprovedto be a synonym for Euphronia.His name
was not only unnecessary,but was also invalidatedby his prioruse of the name
Lightia(1844) for whatis now known as Herraniain the Sterculiaceae.Schomburgk
placedhis second Lightiain the Vochysiaceae.Althoughillegitimate,Lightiahas
persistedto the presentday in the literaturein spite of occasionalreferencesto the
correctname.
The only authorto treat the Trigoniaceaeas a separatefamily in literature
appearingbetween 1841 and 1875 was Grisebach(1849) in a partialrevisionof
Trigonia.Agardh(1858) placed Trigoniain the Polygalaceae,a view that was
followed by Miquelwhen he publishedthe Malaysiangenus Trigoniastrumin 1862
and thus showed that Trigoniaceaeis representedon more than one continent.
Wallich,in 1832, listed a specimen of Trigoniastrumunder the name Isopteris
penangiana,but this name was nevervalidlypublished.
Benthamand Hooker(1867) and Baillon(1874) placed both Trigoniaand
Lightiain the Vochysiaceae,and retainedTrigoniastrumin the Polygalaceae.
The Trigoniaceaewas finally establishedas a family with Warming'smon-
graphictreatmentfor Flora Brasiliensis(1875). He treatedtwo generain the family,
Trigoniaand Lightia,and provideda detailed discussionof the reasonsfor regard-
ing the groupas a separatefamily.Warming'streatmentwas the most comprehensive
study of the family until then, and has remainedso until the present.Althoughhe
studied in detail only the Brazilianspecies, he listed all those known until then,
recognizing26 species in Trigonia,of which 10 were new, and 2 species in Lightia,
one of which was new. He also establishedthree sections within Trigonia,based on
the ultimate arrangementof the flowers:Racemosae,Cymosaeand Cincinnatae.He
recognizedthat these sections were purely artificialand did not reflect relationships
within the genus. In the presenttreatmentthese sections are not given taxonomic
recognition.
In 1895 and 1896 there were two importantpublicationson the Trigoniaceae.
Chodat (1895) transferredTrigoniastruminto the family, and suggestedthat
Lightia did not belong there. In the following year, Barth,one of Chodat'sstudents,
gave anatomicalsupportto Chodat'ssuggestions.
Chodat'sconclusionswere only partiallyfollowed in later treatments;thus
Trigoniastrumwas definitely recognizedas trigoniaceous(althoughnot in time to
be consideredso by Petersenin the first edition of Engler& Prantl,Das Pflanzen-
familien, 1896), but Lightiaremaineda memberof the family, and has remainedso
until the presenttreatment.
Chodatalso establishedthree tribeswithin the Trigoniaceaeto accommodate
the three known genera.The tribeswere Trigoniae,Trigoniastrae,and Ligthiae(sic).
The first two are consideredunnecessaryin the present work, and Lightia (ie
Euphronia)is transferredto Vochysiaceae.
No detailed study of the family has been made in the present century,
GeneralConsiderations 3
althoughit has been treatedin severallocal revisions,in all cases includingonly one
genus. The only importantoccurrencebetween 1897 and the presentwas Leandri's
(1949) descriptionof Humbertiodendron(Humbertodendron)from Madagascar,
addinganothercontinent to the known distributionof the family.
Trigoniawas treated by Standley (1924) for North AmericanFlora, by
Stafleu (1951) for the Flora of Suriname,by Reitz (1967) for Flora Iustrada
Catarinense,and most recently by Austin (1968) for the Flora of Panama.
Trigoniastrumwas revisedby Van Steenis (1949) for FloraMalesianaand by
Ng (1972) in TreeFlora of Malaya.Apartfrom the originalpublicationin 1949,
Humbertiodendronwas studied by Perrierand Leandri(1955) for the Flora de
Madagascar,in which they changedthe orthographyto Humbertodendron.This
second name is acceptedhere as the correctorthographicvariation,based on the
recommendationof the InternationalCode of BotanicalNomenclature(Stafleu,
et al 1972).
GEOGRAPHYAND ECOLOGY
The Trigoniaceaeare found in the Neotropics and in the Paleotropicsof

Malaysiaand Madagascar,with one genus occurringin each of these three geograph-
ical areas(Fig 1). The family occursin tropicallowlands,usually at altitudesbelow
500 meters. The paleotropicalgenera, Trigoniastrum(Malaysia)and Humberto-
dendron(Madagascar)are found as elements of the tropicalwet forest. Trigonia,
the neotropicalgenus, is known from periodicallyflooded riverineforests (called
varzeasin Brazil),galleryforests, the edges of primaryand secondarywet forests
(in the ecotone with adjacentopen areas) and along roadsides.Thus two main
modes of adaptationwithin the family occur;in the Paleotropicsmembersof the
Trigoniaceaeare actualelements of the forests, while in the Neotropicsthey are
found in open but protected habitats.
This dual mode of adaptationis correlatedwith important differencesin
habit and dispersalmechanismsthat separate the paleotropicalgenera from
Trigonia.Trigoniastrumand Humbertodendronare tall trees (ca 25 m), while
species of Trigoniaare treelets, shrubs,scandent shrubs, or lianas. In the paleo-
tropicalgenerathe seed is dispersedwith the fruit, an indehiscenttriwinged
samara.This type of dispersalmechanismis fairly effective for tall trees, and
permitsthe seeds to be disperseda moderate(but not very long) distancefrom the
parent plant. The fruit acts as a parachute,and a moderate breeze will insure
adequate dispersal.In Trigonia,the fruit is a trilocularcapsule that opens with
progressivedryingof the fruit. In the only species I have observedin fruit in the
field (T. spruceana),the capsuleopens at the top, so that the seeds cannot fall out.
A moderatewind is probablyrequiredto carrythe seeds away from the fruit. Of
the 24 species treatedhere in Trigonia,two have echinate trichomeson the seed
and are probablylimited to dispersalby water(or animals?).The other 22 species
have long silky trichomesthat are adequatefor both wind and water dispersal.The
dispersalmechanismof Trigoniaprobablyaccounts for dispersalover much greater
distancesthan that of the paleotropicalgenera.
The originaltrigoniaceousstock probablywas formed of elements that share
more charactersin common with present day Trigoniathan with Trigoniastrum
or Humbertodendron.A largenumberof connate stamenswere probablypresent,
4 Flora Neotropica
_3
. ..
FIG 1. Geographic distribution of Trigoniaceae. , Range of the genus Trigonia. 2, Range of the
genusHumbertodendron(Humbertodendron
saboureaui).3, Rangeof the genus Trigoniastrum
(Trigoniastrum
hypoleucum).
and the fruits were probably capsules with severalvillous seeds. (The seeds of
Trigoniastrumand Humbertodendronare relictuallyvillous, but the trichomes
serveno apparentfunction, for in Trigoniastrumthe seeds are reportedas germinat-
ing in situ by Ng, 1972.) It is probablethat the pubescenceon the seeds of the
ancestralstock was not as functional for wind dispersalas that found in present
Trigonia;it could have been much shorterand still have providedadequatedispersal
for tall trees (acting as a parachutein the same manneras the fruit of the paleo-
tropicalgenera).Witha progressivechangein habit from tall trees to lower shrubs,a
more effective dispersalmechanismwas probablyselected for in the ancestryof
Trigonia.The leaves of the ancestralstock were probablyopposite, with the alter-
nate leavesof Trigoniastrumrepresentingthe derivedcondition. This can be postu-
lated, based on the presenceof opposite leavesin TrigoniaandHumbertodendron,
and suggestsan ancestralstock for these generawith opposite leaves, as the neo-
tropicaland paleotropicalmembersof Trigoniaceaewere probablyseparatedbefore
the ancestorsof Trigoniastrumand Humbertodendron.The arborescentcondition
in Trigoniastrumand Humbertodendronis probably relictual, and the shrubby,
vininghabit of Trigoniaderived.
Although membersof the Trigoniaceaeand especially Trigoniaare fairly
common in some areas,the groupis, with some exceptions, a poorly collected one.
Thereis probablyno one reason for this; Trigoniaas a whole is found close to river
banks in periodically flooded forests, and in drier areas along gallery forests. In
general,species representedin galleryforests are collected more often than those
found along periodicallyflooded riveredges. These collecting practicescould be
due to severalcauses. First, gallery forests are usually botanically diverse,while

there is a certaintendency to ignore riversidecollections as repositoriesof common
(probablyoften collected) species in areaswhere periodicallyflooded forests are
found along well-travelledwaterways.Thus, in many ways, the ease of access to the
localities where this family is representedmay account for the scarcityof material
representedin herbaria.This has been recognizedby presentcollectors, and areas
that have been ignored in the past are now being sampledwith very rewarding
results.
Othercausesmay be responsiblefor the scarcityof collections in the family.
Some species probablyhave very low densities.In the course of my field work in
Brazil(1973), I had the opportunityto observeindividualsof four speciesin the
field. Only one of these had more than one individualat any givenlocality.
Also duringmy field work, I was able to obtain seeds of one species (T.
spruceana).The seeds were subjectedto severaldifferenttypes of treatmentat the
researchgreenhouseof the New York BotanicalGarden.Some seeds were cold- or
heat-treated;others were placedin water at differentpH (5.6, 6.0); otherswere
plantedin sand or differenttypes of soil at differentpH to simulatethe natural
conditions presentat the site where they were collected. To the present,it has been
impossibleto breakdormancyand induce germination.
The Trigoniaceaeis regardedby Thorne(1972) as havingan Amphi-Pacific
tropical distribution.Togetherwith Elaeocarpaceae,Chloranthaceae,and sixteen
generain other families,Trigoniaceaeis what Thornecalls a groupwith tropical
Amphi-Pacific-Madagascar distribution. The modes by which these and other
familieshave achievedtheir presentdisjunctdistributionhave been the subjectof
much speculation.Van Steenis (1962) has postulatedthe existence of land-bridges
acrossthe Pacific. Others(Wolfe, 1969; Thorne, 1972) have favoredthe Beringian
land-bridgeas a more probablepath for the dispersalof disjuncts.Thorne(1972)
also indicatedthat a similarland-bridgecould have existed in Antarcticathat would
account for groupswith australdispersalpatterns.ContinentalDrift has also been
suggestedas a possible cause (eg Raven& Axelrod, 1972, 1974), a possibilitywhich
Thorne(1972) considersas improbable.
In spite of the lack of fossil evidence, I find it possible to hypothesize a
possibleexplanationfor the disjunctdistributionof the Trigoniaceaebased on the
currentviews on ContinentalDrift. The presentviews on the originof the Angio-
spermsplace the date for this event (or seriesof events) in the lower Cretaceousca
125 million years ago (Axelrod, 1970; Raven & Axelrod, 1974). By the Middle
Cretaceous(ca 110 million years ago) many generaand severalextant familieswere
alreadypresent (Raven & Axelrod, 1974). Africa, South America,Antarctica,
Madagascarand Indiawere in close contact at that time and remainedin contact
until 90 to 100 million years ago, accordingto the currentviews on continental
drift (Green, 1972; Raven& Axelrod, 1972, 1974). By the end of the Mid-Creta-
ceous, Africaand South Americastartedto drift apart,and Madagascar and India
became separatedfrom the Africanmainland.Madagascar and Indiaeither started
to drift apartfrom each other (Raven& Axelrod, 1974) or remainedin contact
for a longer period of time (Cracraft,1973 in Raven & Axelrod, 1974). Direct
migrationbetween Africa and South America(West Gondwanaland)was last
possible between 90 and 110 million years ago (Raven& Axelrod, 1974).
Raven& Axelrod (1974) note that the familiesfor which a relationshipwith
the Trigoniaceaeis suggestedin this paper,Polygalaceae,Sapindaceae,Malpighiaceae,
6 Flora Neotropica
Dichapetalaceaeand Vochysiaceae, all presumablymigratedbetween Africa and

South Americaduringor priorto the Paleocene(before 54 million years ago). Of
these, Dichapetalaceaemore closely resemblesTrigoniaceae;the basic differencein
distributionis the absenceof Trigoniaceaefrom Africaat present.A similarsitua-
tion to that postulatedfor the above-citedfamiliescan be hypothesized for the
Trigoniaceae.
If the Trigoniaceaeoriginatedin WestGondwanaland(Africa-SouthAmerica),
and if the initial migrationoccurredwhile the continents were still together, the
ancestralstock givingoriginto the extant Trigoniaceaewas alreadypresentin the
UpperCretaceous.Subsequentmigrationinto Madagascar-India probablyoccurred
before these two areaswere separated.This migrationprobablyoccurredvia Africa,
where the Trigoniaceaewere probablypresentin the past. The seriesof climatic
changesthat have affected Africanflora, with conditionsbecomingprogressively
more arid,probablyaccount for the extinction of the family in that continent.
This case is not unique, and has been reportedfor many groupsfor which there is
adequatepaleobotanicalevidence(Thorne, 1972; Raven& Axelrod, 1974). The
climate in Africais characterizedby havingseveredry periods,even in the tropical
wet forests. Presentday membersof Trigoniaceaeare characteristicof humid
environments(with the exception of Trigoniarugosafrom Guanacaste,Costa Rica)
in which there is no extreme dry period. If this characteristicwas inherentin the
family from the beginning,it would help explain their limited success and disapear-
ance from the Africanmainland.
WhenMadagascar-India became separatedfrom the Africanmainland,the
membersof Trigoniaceaethat migratedinto this land massbecame isolated from
those presentin the Africanmainland.After a period of uncertaindurationduring
which Madagascarand Indiawere isolated together, and duringwhich a certaindegree
of evolution probablyoccurred,the ancestralstocks for Humbertodendronand
Trigoniastrumwere separatedwith the separationof India from Madagascar. The
hypothetical common ancestralstock that was to give rise to these two generawere
trees, with (probably)opposite leaves, a few (5-6) connate stamens,and fruit in
triwingedsamaras.Whetherthis ancestralstock had developedmost of these characters
while still on the Africanmainland,or whetherthese characterswere evolved on
the Madagascar-India land massin uncertain.Whenthe Indiansubcontinentcame
into contact with Asia, the floras of these two areascame in contact with each other,
with the subsequentmigrationof elements (includingTrigoniaceae)from one areato
the other. Subsequentextinction of Trigoniaceaefrom the Indiansubcontinent
must be hypothesized.
The above hypothesis, based on ContinentalDrift, is not the only possible
explanationfor the presentdistributionof disjunctsin the family. Long distance
dispersalmay have accountedfor some or all of the disjunts.Thorne(1972) notes
that of the 89 generaknown to be Amphi-Pacific,25 are known to a have migrated
successfully over great distances of water. The dispersalmechanismof Trigonia,
if presentin the ancestralstock, makesit possible to envisionlong-distancedispersal
by either wind or water.WhenI was trying to induce seeds of Trigoniaspruceanato
germinate,I floated some seeds in water,in hope of breakingtheir dormancy.I got
no such results.After three months most of the seeds were still floating, and had
absorbedlittle or no water, as the trichomes retainedair and kept water from
reachingthe seed coat.
It is now believedthat a seriesof drasticclimaticchangessimilarto those
reportedfor Africatook place in South Americaduringthe Pleistocene.Vuillemier
(1971), Haffer(1969) and Vanzolini(1970), using palynologicaland climatological

data have postulated the existence of a seriesof alternatingdry and wet periods.
Palynologicaldata obtainedby van der Hammen(1966, 1974 in press,cited by
Haffer, 1974) indicate that severalaridconditions prevailedduringthe peaks of the
glaciationperiods. Haffernotes that the fluctuation of the temperaturebetween
glacialand interglacialperiodswas of 3 to 4 C in the tropicallowlands,and of 7 to
8 C or more at high altitudes.This suggeststhat at high altitudesglacialperiods
caused a greaterimpact on the environmentin regardto the temperaturethan at
lower elevations.It can be hypothesized that in the lowlandsthe criticalfactor that
was affected by glacialperiodswas humidity and not (to any largeextent) tempera-
ture. This led to a reduction of the rainforestsand a gradualinvasion by more
xerophytic types of vegetation(Haffer 1969; Vanzolini 1970; Vuillemier1971).
Studies on distributionpatternin birds(Haffer, 1969, 1974), Anolis lizards
(Vanzolini, 1970, 1973) and angiosperms(Prance,1973) have led these authorsto
postulatethe existence of forestrefugesin the Amazon Basinduringthe Pleistocene.
These refugeswere restrictedareasof forest that were isolated by invasionof the
savannas.The species of forest taxa that survivedin the refugeswere isolated until
the more humid interglacialperiods,when the forests re-invadedthe areasthat had
been claimedby the savannas.Geologicaland palynologicaldata for AndeanSouth
Americaindicate that at least three majorglaciationsoccurredduringthe pleisto-
cene (van der Hammen,1966; Haffer, 1974). This would suggestthat refugeshave
existed at least three times in the past 1.2 million years. The present-dayrefugesas
postulated by Haffer (1969, 1974), Vanzolini (1970, 1973) and Prance(1973)
probablyonly reflect the refugesformed duringthe last dry-glacialperiod(10,000-
15,000 years ago) and are not necessarilythe same as those formedin previous
glaciations.
The present distributionof Trigoniais furtherevidence in support of the
theory of Pleistoceneforest refugesfor the Amazonbasin. I have obtainedsimilar
resultsto those reportedby Prance(1973) for Caryocaraceae,Chrysobalanaceae,
Dichapetalaceaeand Lecythidaceae.For the purposeof this paper,I have adopted
the concepts and nomenclatureused by Prance(1973) to delimit the refuges
(Fig 2). Withfurtherpaleobotanicalwork in the Amazon Basin,a somewhatdiffer-
ent distributionof refugesmay have to be postulated.
Althoughlittle work has been done on the ecology of Trigonia,my observa-
tions in the field, as well as the examinationof herbariumspecimens,provideenough
data to infer some of the possible reasonsfor the distributionpatternin the genus.
It is difficult here to refer to populations,as it has provento be difficult to deter-
mine what constitutes a populationfor any species in Trigonia.In most cases, I
have been unableto observemore than one individual,or to find more than one or
two individualsthat are apparentlypart of the same population.The data presented
in this treatmentreflect data collected on individualsonly, and from these data one
must extrapolatethe possibleimpact of the environmenton Trigoniaspecies.
The papilionaceousflower of Trigonia(and all Trigoniaceae)is suggestiveof
pollinationby insects. The presentationmechanismof the stamensand the stigma
to the pollinatoris essentiallythe same as that found in the Fabaceae,and I believe
it works in a similarmanner.Whetheror not the attractantis nectarproducedby
the glandsis not knownat present,as I havebeen unable to find any tracesof nectar
on either dry or the few fresh flowers available.Althoughthe flowers are described
as white for all species, a careful examinationof the standardin fresh material
shows the presenceof tenuous purpleor yellow nectarguides.AlthoughI have not
8 Flora Neotropica
00 .00 .....0
/ N
.'
FIG 2. Forest refuges proposed by Prance (1973). 1. Choc6; 2. Nechi; 3. Santa Marta; 4.
Catatumbo;5. Rancho Grande;6 Paria; 7. Imataca; 8. Guiana; 9. Imeri; 10. Napo; 11. Olivenca;
12. 'Tefe; 13. Manaus; 14. East Peru; 15. Rond6nia-Aripuana; 16. Belem-Xingu. The areas
surrounded by a heavy black line correspond to areas flooded during the pleistocene.
been able to observepollinationin the field, the label on one herbariumspecimen

of Trigoniapaniculataindicatesthat it "wasvisited by a largebee."
A distributionpatternwith few, widely separatedindividualscan be correlated
with pollinationby largebees; Janzen's(1971) work on flight patternsand rangeof
euglossinebees indicates that those bees are capableof coveringlong distanceson
their normal forage flights. It is improbablethat Trigoniais self-pollinated;the
small number of fruits in contrast with the large number of flowers produced
suggeststhat this is not the case.
The presenceof buds, flowers and fruits at variousstages of developmenton
the same herbariumspecimenfor most of the species of Trigoniasuggestsa lack of
seasonalityin floweringand fruiting.This is suggestiveof a more or less homogene-
ous environmentthroughoutthe year. Furtherevidence for the above statement is
providedby the types of environmentwhere species of Trigoniahave been collected.
These ecological observationsmay help in understandingwhy species of
Trigoniaare useful indicatorsof the refuge theory. The genusis adaptedto a humid
environmentand seems to have a specialized pollination mechanism.These two
factors would help explain why there has not been significantmigrationof the
differentelements cited here when the savannaswere reclaimedby the rainforests

and also help explain the largenumberof endemic taxa.
The present distributionof Trigoniain South Americashows a phytogeo-
graphicaldichotomy. Thereis a southeasterngroupwith severalspecies found in
the vicinity of Rio de Janeiro,and with distributionsextending towardsthe north
in the easterncoastal forests, and towardthe northwest of Brazilalong galleryand
edges of forests. These species probablysurvivedthe dry period of the last Pleisto-
cene recessionof the forests in the easterncoastal forests, or in the galleryforests,
and only the northernmostpopulationswere influencedand in some cases survived
in refuges,as has been shown by Prance(1973) for other families.A second, north-
centralgroup,presentsa largenumberof speciesin the coastal forests of the Guianas
and Venezuela,andalso alongthe riverineforests of centraland northernAmazonia.
This groupwas the most stronglyaffected by the Pleistocenedry periods,and the
species were isolated in refuges (Prance 1973). Severalspecies were in different
refuges throughout their ranges.Some have distributionsin eastern coastal and
galleryforests in the south, and in refugesin the northernpart of their ranges.
At presentit is impossibleto determinethe extent to which galleryforests
were destroyed or survivedwith the invasionof the forests by savannasduringthe
Pleistocene. Thereare no data to indicate what the successionpatternwas in the
reclamationof grasslandsby the forests once conditions becamemore humid.
Severalspecies are found in the easterncoastal forests: Trigoniarotundifolia
(Fig 6), T. rytidocarpa(Fig 6), and T. paniculata.Trigonianiveavarfasciculata
(Fig 7) is also endemic to this area, while var nivea is also found in Pariaand
Guianarefugesand varpubescensis found in the Manausrefuge. Trigoniaerio-
spermasubsp eriospermaand subspsimplex are endemic to the easterncoastal
forests, while T. membranacea(Fig 3) is found only in northernSouth America
and CentralAmericaand was probablyassociatedwith the Nechi and SantaMarta
refuges.Trigoniavillosa(Fig 5) varmacrocarpais endemic to the easterncoastal
forest, while var villosais also a component of the Guianarefuge.
Threespecies are endemic to the forests adjacentto the easternlimit of the
Andes: Trigoniaboliviana(Fig 4), T. floccosa (Fig 4) and T. echiteifolia.
The Guianarefugeharborsthe largestnumberof species of Trigonia.Four
species are endemic to this area: T. hypoleuca (Fig 5), T. coppenamensis(Fig 6),
T. subcymosa(Fig 4), and T. candelabra(Fig 4). Severalother species are repre-
sented there, althoughthey are not endemic,notably T. nivea, T. villosaand T.
laevis(Fig 6).
The Napo refugeis also importantin numberof species. Two species are
endemic there, Trigoniamacrantha(Fig 5) and T. prancei(Fig 4) (this species
could also be from the East Perurefuge;it is a borderlinecase). Two others are
represented:T. sericea(Fig 5), which is also found in the Napo and Nechi
refuges,and T. virens(Fig 5), which is also found in the Rondonia-Aripuana
refuge.
Trigoniaspruceana(Fig 5), a moderatelywidespreadspecies, is found in the
Manausrefuge, and possibly in the Imeri. Trigoniacostanensis (Fig 5) is found
in the Rancho Granderefuge.
The Nechi refugecontains Trigoniarugosa(Fig 3), T. sericea,and T. erio-
spermasubspmembranacea;the Santa Martarefugecontains T. rugosaand
possibly T. eriospermavarmembranacea.Trigoniabracteata(Fig 4) is found
close to the Rancho Granderefuge.The Catatumborefugecontains T. rugosaand
T. reticulata(Fig 6), and T. killipii (Fig 5) is found in the East Perurefuge.
10 Flora Neotropica
FIG 3. Known distributionof species of Trigonia.0 T. rugosa; v T. eriospermasubsp

membranacea.
0,
FIG4. Knowndistributionof speciesof Trigonia. 0 T. prancei; T. boliviana; T.floccosa;

x T. candelabra; T. eriospermasubsperiosperma;* T. eriospermasubspsimplex; A T.
subcymosa; T. racteata.
80 70 60 50 40
FIG 5. Known distribution of species of Trigonia. N T. macrantha: 0 T. killipii; * T. virens;

A T. sericea; 0 T. hypoleuca; l T. echiteifolia; A T. spruceana; * T. villosa var villosa;
T. villosa var macrocarpa; * T. paniculata; * T. costanensis.
ANATOMICALCONSIDERATIONS
The anatomy of the Trigoniaceaewas studied thoroughlyby Barth(1896),

who treatedmost of the known species of Trigoniaas well as Trigoniastrumand
Euphronia.The secondary xylem was also treated by Heimsch(1942) and the
Trigoniaceaewere studied by Metcalfeand Chalk(1951).
12 Flora Neotropica
80 0
,
FIG 6. Known distribution of species of Trigonia. * T. reticulata; [ T. coppenamensis:

- T. laevis var laevis; T. laevis var microcarpa; r T. rotundifolia; I T. rytidocarpa.
In the presentstudy five randomlyselected specimensof each of the more

commonly collected species and one specimeneach of those poorly known from
collection were surveyedfor anatomicalstudy. Smallstem segments,petioles, and
leaf bladeswere sectioned using different techniques.Most were sectioned with the
aid of a freezingmicrotome, but some sections were made by the freehandmethod,
and a smallnumberwere treatedusing paraffintechnique. The dyes used in all cases
were saffraninand fast green.
I examined enough materialto confirm the data given by Barth (1896),
Heimsch(1942) and Metcalfeand Chalk(1951). I also studiedHumbertodendron,a
genus that previouslyhad not been treatedanatomically(Fig 8). In the leaf, I found
trichomesto be simple and unicellular;the epidermisoften with mucilaginouscells
is uniseriatein Trigonia(except T. rugosa) and 2-layeredin Trigoniastrumand
Humbertodendron.Stomata are paracytic, and confined to the lower surface.
it"..
pubescens; o T...ni.......ci.l..
FIG 7. Known distribution of species of Trigonia. * T. nivea var nivea; (3T. nivea var
pubescens; 0 T. nivea var fasciculata.
Mesophyllisis with
with branchedsclereids
sclereids in Trigoniastrumand Hu
Trigoniastrum
in Humbertodendron;
and
rbertodendron;
palisadeparenchymais usually l-layered but sometimes2-layeredin Trigoniastrum
and Humbertodendron;rhomboidor irregularlychamberedcrystalsof calciumare
presentinand
niastrum Trigoniastrum, Humbertodendron
other Trigonia and several speciesisofsimple
Trigonia.
species. In the
The petiole has a simple stem,
all
the epidermis
three
and often
epidermis in genera;the trichomes,when
present,are simple and unicellular.The phloem completely surroundsthe xylem in
Humbertodendron and some species of Trigonia, or is open at the top as in Trigo-
has tannins.The cork i from subepidermalphellogen,and the cork cells
species
has tannins.The cork is derivedfrom subepidermalphellogen,and the cork cells
14 Flora Neotropica
often have tannins, and the regionjust below the peridermisis often subcollen-
chymatous. Corticalsclereids are often present and are especially common in
Trigoniastrumand Humbertodendron.Calciumoxalate crystalsare frequentin all
three genera (lacking in some species of Trigonia).The secondaryphloem has
sclereids. Intraxylaryphloem is absent, and the vessels are solitary (although
Heimsch1942, reportsporemultiplesin some speciesof Trigonia).The perforations
are simple and elliptic (sometimes double in T. eriosperma,fide Barth).The paren-
chyma is apotrachealin all three genera(1 specimenof Trigoniastrumreportedas
havingparatrachealparenchymaby Metcalfe& Chalk). The rays are 1-5-seriate,
heterogeneous(Kribs' type IIa and IIb). The wood is diffuse-porous.The pith
often has tannins,with or without crystals;in Humbertodendron,rhomboidcrystals
arevery common.
Cross sections and dissections of the flower of Trigoniaspruceanaand T.
prancei were made in order to determinethe nature of the floral glands. The
vascularizationof the glandsis the same as that found in the staminalring,with
2-3 vascularstrandsattachedto (ie entering)each gland.This supportsmy views
based on morphologythat the glandsare actuallymodified stamens.
Anatomically,the familyis fairlyuniform,with no single characterfacilitating
differentiationamongthe genera,althoughthere are trendsthat are more noticeable
in some groupsthan in others.
Humbertodendronis anatomicallyvery close to Trigoniastrum; in fact, the
anatomy of the leaf blade is so similarin the two generathat laminarmaterial of
the two generaeasily can be confused in the laboratory.The petioles are somewhat
different,however,especiallyin the position of the vascularbundles.As indicated
in the anatomicaldescription,the phloem in Humbertodendrontotally surrounds
the xylem, but is open in Trigoniastrum.In Humbertodendronsclereidsare present
only alongthe rays,while they are randomlydistributedin Trigoniastrum.Although
rhomboidcrystalsoccur in the pith in both genera,they are much more abundant
in Humbertodendron.
Barth(1896) consideredthe Trigoniaceaeas relatedto the Dichapetalaceae
on anatomicalgrounds.Heimsch(1942), based on his work on secondaryxylem,
consideredthe Trigoniaceaeclose to the Polygalaceae,the Tremandraceae,the
Zygophyllaceae,the Malpighiaceaeand the Vochysiaceae.Metcalfe and Chalk
(1951) consideredthe family closest to the Vochysiaceae.
The Trigoniaceaediffers from the Vochysiaceaein the absenceof intraxylary
phloem, the absenceof libriformfibers, and in havingapotracheal(vs paratracheal)
parenchyma.
Anatomically, the Trigoniaceaehave undoubtedly close affinity to the
Vochysiaceaeas well as the Polygalaceae.The suggestedrelationshipswith Dicha-
petalaceaemay also be correct. The taxonomic positions and familialrelationships
of the familiesin both Cronquist's(1968) Sapindalesand Polygalales,as well as the
relationshipof these familieswith the ancestralRosales,remainratherunclearat
present.
On the basis of anatomy, which corroboratesthe morphologicalobservations,
I agree with Barth that Euphroniashould be removed from the Trigoniaceae.
Anatomically,thereis no seriousobjectionto returnEuphroniato the Vochysiaceae,
with which it has many charactersin common. The data used to support this
decision are discussedelsewhere(Lleras, 1976).
'4 S7l
t: "'. '
~ ...*,
g,* 4
A B
Wi 35
.
eIf'j : '-t?
'I,r~ *...
' ?
' 9.. ? .
E F
FIG 8. Anatomy of Humbertodendron saboureaui (Perrier & Louvel 14896bisJ. A, young stem,
longitudinal section X 9.2;B, young stem, longitudinal section showing rhomboid crystals X 34;
C, young stem, cross section X 9.2;D, petiole, cross section X 18; E, Lamina, cross section X
66; F, lamina
la :.inacentralnervesX 18.
16 Flora Neotropica
POLLEN
The terminologyand formatused here to describethe pollen of the Trigo-

niaceaeare taken from Erdtman(1952) and Walker(1971).
Pollen Morphology:(Fig 9). The grainsare solitary, square or suboblate,
sometimesoblong;34 (rarely5)-porate;25-60 I on the longest axis (highly variable
within the same species). The exine stratificationis obscure,and the surface,when
observedwith the ScanningElectronMicroscope,is tenui-exinousand psilate. The
sexine is probablythickerthan the nexine, which has (fide Erdtman,1952) small
projectionsdirectedtowardsthe interiorof the grain.
In Trigonia,T. spruceana(42-60 p) and T. nivea (40-55 ,t) have the largest
grains.Trigoniarugosa(28-40 ,u)and T. prancei(25-38 ,u)presentthe smallestgrains.
The majorityof the species have grainsrangingbetween 35 and 45 gu(eg T. villosa,
T. virens,T. rotundifolia,T. echiteifolia, T. laevis and T. reticulata).The pollen of
Humbertodendron(only one specimenavailablefor sampling)has a size of ca 55
,; Trigoniastrumhas grainsrangingbetween 35-43- ,I. The high degreeof overlapin
grainsize in Trigoniamakesit impossibleto separatethe pollen into discretesize
groups.Withoutsuch discretegroupsit is impossibleto suggestpolyploids within
the genusbased on pollen evidence.
Althoughpollen charactershave not proveduseful taxonomicallywithin the
family due to the high degreeof overlapin size and to the very similarmorphology
(practicallyidentical for all three genera),pollen has provedto be of greatvalue in
distinguishingthe Trigoniaceaefrom other families,and has been useful in establish-
ing this family'srelationshipwith the Sapindaceae.Erdtman(1952) noted a similar-
ity of pollen in the Trigoniaceaewith the pollen of Paullinia(Sapindaceae),an
observationwith which I concur. There is a slight tendency in the Trigoniaceae
towardsmultiporatepollen, as can be seen in the progressionfrom 3-porateto 5-
porate grains(as in Trigoniaprancei, Fig 9E), and this might indicate a relationship
with the Polygalaceae,which have multiporatepollen. One might considertrigoni-
aceous pollen as somewhatintermediatebetween that of the Sapindaceaeand the
Polygalaceae,but this does not mean that I considerthe Trigoniaceaeas an inter-
mediate family.
EVOLUTIONOF CHARACTERSWITHINTHE FAMILY
Althoughit is difficult at presentto discerngeneralizedtrends of evolution

within the Trigoniaceae,the possible evolution of individualcharacterscan be
discussed.It is acceptedhere that the majortrendsin Angiospermfloral evolution
have involved progressivereduction and simplification,as postulated by Bessey
(1915), Tajktajan(1959, 1969) and Cronquist(1968).
One of the most interestingfeaturesis the changefrom a basicallytrilocular
ovary (4-locularin Trigoniarotundifolia) to a unilocularovary. The unilocular
ovaryoccursin T. virensand T. hypoleuca. This reductioncan also be observed
in Humbertodendronand severalother species of Trigonia,although in a less
advancedstate, with the septa presentbut not fused at the center. This processhas
apparentlyoccurredby a gradualreduction of the lateral septa of the locules:
first, as in Humbertodendronand T. sericea,the septa are complete but fail to fuse
at the center;then, asin both T. virensand T. hypoleuca, a gradualshorteningof the
septa occurs. Although severalof the intermediatesteps can be observedin the
A B
E F
FIG 9. Pollen of Trigoniaceae.A-C,ScanningElectronMicrographs,X 436. D-F, with light

microscope,X 171. A, Humbertodendron saboureaui(Perrier& Louvel14896 bis); B, Trigonia
villosa (Pires& Cavalcante52283); C, Trigoniarugosa (Burger7841); D, Trigoniaprancei
(Lleras,Stewardet a 17115); E, Trigoniaprancei(Lleras,Stewardet al 1 7115); F, Trigonia
rugosa(Haught4227).
18 FloraNeotropica
ovariesof the last two species, by the time the fruit reachesmaturityno trace of
the lateralsepta remains.
Evidenceof a gradualreductionof the androeciumcan be seen in livingspecies
of the Trigoniaceae.The most primitiveexisting condition occurs in Trigonia
macranthaand T. nivea, whose flowershave fairlylargenumbersof stamens(ca 12),
some of which are reducedto staminodes.The intermediatecondition is found in
species in which the staminodeshave been lost, and all of the stamensare fertile,
as in T. virens.The condition which I considerto be the most evolved, where there
areno staminodesand a relativelysmallnumberof stamens(ca 6), is presentin both
Trigoniastrumand Humbertodendron.
If we acceptan ancestorin the Rosales(as discussedundersystematicposition
of the Trigoniaceae)as the precursorfor Trigoniaceae,it was probablya groupof
plants with a largenumberof indefinite or definite stamensdisposedin a staminal
ring.The reductionprocessprobablyoccurredas follows: first, a complete reduc-
tion of a whole section of the staminalring,with subsequentevolution of the disc
glands. I interpret these glands as reduced and modified stamens, based on the
laciniaewhich are morphologicallyvestigialfilaments;anatomicalevidence for this
is discussedin the section on anatomy. Next there was a loss of antherson some of
the remaining(outer) stamensof the portion that remained;and finally, a loss of
staminodes,giving rise to the extant taxa displayingfew fertile stamens and no
staminodes.
The obvious differencein the dispersalmechanismbetween the neotropical
and paleotropicalTrigoniaceaerepresentsan importantevolutionarychange. In
both Humbertodendronand Trigoniastrum,the seeds remainin the winged fruit,
and are dispersedwith it. In Trigonia,however,the fruits are dehiscent,and the
seeds are adaptedto wind and water dispersal.This transferenceof function for
seed protection and dispersalfrom the seed to the ovary and vice-versais common
in the Angiosperms(Stebbins, 1974).
As is discussedin the section on geographyand ecology, and based on geo-
graphicaland ecologicalevidence, the plants of the ancestraltrigoniaceousstock
were probablytrees, with many-seededcapsules.The seeds were probablyvillous,
although the trichomes were probably shorter than those found on present-day
species of Trigonia.
Figure 10 is a schematic representationof my opinion of similaritiesand
differencesin Trigonia.The distancesdo not representcloseness of relationshipand
only serveto separatetaxa of dubiousposition. It is not my intention to suggest
phylogeny, as I believedthat presentevidenceis insufficient to do so. It is also
difficult to determinewhich species are primitiveor advanced,as in almost every
case a species that presents a characterthat I consider primitivealso presents
charactersthat I consideradvanced.
SYSTEMATICPOSITIONOF THETRIGONIACEAE
This section on Taxonomic Historypresentsthe taxonomic positions previ-

ously suggestedfor the family and in some cases for individualgenera.As can be
seen from this account, there has been considerabledivergenceof opinion as to the
systematicposition of the Trigoniaceae.A summaryof the most importantrelation-
ships suggestedfor the family follows:
,iiE,~i
FIG 10. Schematicrepresentationof similaritiesand differenceswithin Trigonia.
Aublet (1775) Decandria Monogynia

De Jussieu (1789) aff. Malpighia
De Candolle (1824) in Hippocrateaceae
Cambessedes (1829) in Hippocrateaceae
Martius (1835) fam. in Malpighinae
Grisebach (1849) with Polygalaceae & Euphorbiaceae
Agardh (1858) in Polygalaceae
Bentham & Hooker (1867) in Vochysiaceae
Baillon (1874) in Vochysiaceae
Petersen in Engler & Prantl (1896) Polygalales
20 Flora Neotropica
Hallier(1921) with Dichapetalaceae& Chrysobalanaceae

in Linales
Leandri(1949) Humbertodendronclose to Sapindaceae
Croriquist(1968) Polygalales
Tahktajan(1969) Polygalales
Hutchinson(1973) Polygalales
Breteler(1973) with Dichapetalaceae& Malpighiaceae
Most frequentlythe Trigoniaceaehave been associatedwith the Vochysiaceae
and the Polygalaceae(Polygalales);alternativelythe family has been linked with the
Dichapetalaceaeand Malpighiaceae.These two suggestedrelationshipsare less
contradictorythan may be supposed.Evidenceis presentedhere suggestinga closer
relationshipfor these familiesthan is commonly assumed.
Anatomicalandmorphologicalevidencehas been cited to supportboth points
of view; for example, Barth(1896), on anatomicalgrounds,suggestedthe relation-
ship between Trigoniaceaeand Dichapetalaceae.He consideredEuphronia(placed
in the Vochysiaceaein the presentwork) as anatomicallyintermediatebetween the
two families.Heimsch(1942), who basedhis decision on anatomy of the secondary
xylem, placed Trigoniaceaewith the Polygalaceaeand Malpighiaceae,as well as
with the Tremandraceaeand Zygophyllaceae,all familiesincludedin Cronquist's
(1968) Sapindalesor Polygalales.Metcalfe& Chalk(1951) consideredVochysiaceae
to be anatomicallythe family closest to the Trigoniaceae.These opinions based on
anatomy perhapsdemonstratethat the family is not clearly defined anatomically.
It also suggeststhat severalof the familiessuggestedas relativesfor the Trigoniaceae
have about the same level of anatomicalspecialization.
Palynologicalevidence cited by Erdtman(1952), and with which I agree,
indicatesa relationshipwith the Sapindaceae;the smooth, 3-5-poratepollen in the
Trigoniaceaeis very similarto that of Paullinia.It is also somewhatsimilarto the
pollen of the Polygalaceae,althoughthis family is characterizedby a multiporate
type of pollen grain.
Morphologically,the Trigoniaceaebear a strikingsimilarityto severalfamilies;
Breteler(1973) and Prance(1972) have both indicatedpossible relationshipsof the
Trigoniaceaeand the Dichapetalaceae,based on both anatomicaland morphological
data. Prancehas cited the tendency towardszygomorphy,the lack of endosperm,
and the polygamousflowers of the Dichapetalaceaeas possibleevidence of relation-
ships of this family with the Polygalales;Bretelerhas interpretedthe apparent
corolla of both the Trigoniaceaeand the Dichapetalaceaeas not being a true corolla
but as a structurederivedfrommodifiedstamens.He felt this to be furtherevidence
for consideringthese two families as closely allied. This may be true for the
Dichapetalaceae,but I find no evidence to supportthis hypothesis for the
Trigoniaceae.Nevertheless,the reductionof the disc glands,and the presenceof
connate stamens(if they can be regardedas such in Stephanopodium),seem to me to
be a significantcharactersin consideringthese two familiesas closely allied, as they
suggestsimilarevolutionarytrends.
The relationshipsof Trigoniaceaewith both Sapindaceaeand Polygalaceae
are probablymore evident at first sight. Both the Trigoniaceaeand the Polygalaceae
have the papilionaceouscorolla that makes them distinct. The Sapindaceae,although
not papilionaceous,have severalcharactersthat have probablyevolvedin the
same direction;the stamensare in many cases very much like those of Trigoniaceae,
and some petals are very similar(almost identical) to those of the Old Worldgenera
of the latter family. Another significantcharacteris the occurrenceof winged
samarasin all three families.
TableI presentsa comparisonof morphologicalcharactersof the Trigoniaceae,
the Vochysiaceae, the Polygalaceae,the Sapindaceaeand the Malpighiaceae.A
comparisonof charactersin the table is indicativeof the difficultiesencountered
when trying to establishrelationshipsbased on morphologicaldata.
The evidencecited above supportsCronquist(1968) who accepted a general
relationshipamongthese families,which he believesare derivedfrom the common
Rosalean stock, but I am not entirely convinced that the degree of relationship
amongthe familiesis clearat present.Thereis no doubt that the familiesincluded
in the Polygalalesare related,and are close to the Sapindales(Cronquist,1968) and
possibly the Celastrales(Prance, 1972). However,the relationshipsamong the
familiesin these three ordersare complex. Indicationsfor this have been givenby
Prance(1972), and Breteler(1973) on morphologicalgrounds,and anatomical
evidencehas been providedby Barth(1896) and Heimsch(1942).
The evidencepresentedabove supportsa common originin the Rosalesfor
the Polygalales,Celastralesand Sapindalesas has been suggestedby Prance(1972).
This would explain a numberof familiesthat seem to have affinitiesin all three
orders.Thus, the Trigoniaceaetogetherwith the Vochysiaceaeand the Dichapetal-
aceae and probablythe Malpighiaceae(fide Breteler,1973) would occupy positions
close to the point of divergenceof the three orders.
The Trigoniaceaehave (as have the other three families)evolvedin a direction
of their own, and probablyarrivedat the papilionaceoustype of flower independ-
ently. This suggeststhat this distinctiveflower type has evolved at least three times
in responseto insect pollinators(the Fabaceae,the Polygalaceaeand the Trigoni-
aceae). At presentit is difficult to establishthe closest relativesof the Trigoniaceae,
but it would be misleadingto give undue emphasisto the papilionaceouscorolla.
It is significantthat the relativessuggestedfor the Trigoniaceaeall belong to
familiesthat have significantlymodified corollas(Breteler,1973). This suggestsa
highly adaptablegroupin relationto the corolla, anotherfactor that would suggest
a common ancestry.
Taxonomically, I maintainTrigoniaceaein the Polygalalesas defined by
Cronquist(1968), as most of the possible relativessuggestedin this paper are
includedin that order.
TAXONOMICPOSITIONOF EUPHRONIA
As discussedin the section on TaxonomicHistory,Euphroniawas placedin

the Trigoniaceaeby Warming(1875), a decisionlater questionedon morphological
groundsby Chodat(1895) andon anatomicalgroundsby Barth(1896). Nevertheless,
until now Euphroniahas remainedin the Trigoniaceaeof all authors.Thereare,
however, many marked anatomical and morphological differences between
Euphroniaand Trigoniaceaesens. str. A comparisonof some charactersin Trigo-
niaceae,Euphroniaand Vochysiaceaeis givenbelow:
TABLE I
Trigoniaceae Dichapetalaceae Malpighiaceae Sapindace
position of leaves opposite or alternate alternate opposite (u) alternate,

opposite
type of leaves simple simple simple simple or

compound
stipules + + + (-) + (R)
floral symmetry zygomorphic actino (-zygo) actinomorphic actino, zygo
number of sepals 5 5 5 5
arrangement of sepals imbricate imbricate imbricate imbricate or

valvate
number of petals 5 (papil.) 5 5 3-5
arrangement of petals convolute imbricate convolute imbricate
number of stamens (total) 5 - 13 5 10 Ca 8
staminodes + (-) + -
filaments connate free or connate connate (u) free
number of locules on anthers 2 2 2 2
type of dehiscence of anther introrse introrse introrse introrse

TABLEI (continued)
Trigoniaceae Dichapetalaceae Malpighiaceae Sapindac
glands + + + (ext) +
position of ovary superior superior- superior superior

inferior
number of locules 3(14) 2-3 3 3(14)
type of style simple simple divided simple or di
number of ovules per locule 1 - many 2 1 1-2 (many)
orientation of ovules epitropous epitropous epitropous epitropous
placentation on intruded placentas axile-pendulous ascending- axile or par

pendulous
type of fruit capsules or samaras drupes samaras or various

drupes
endosperm - - -
embryo straight straight straight or plicate or tw

curved
24 Flora Neotropica
Trigoniaceae Euphronia Vochysiaceae
Pollen3-5 porate Pollentricolporate Pollentricolporate
Petals5 Petals3 Petals 1-5
Stamensall connatein one Stamensin 2 or 3 groups Stamensin 1-severalgroups

structure
StaminodesO-several Staminode1 Staminodesseveral
Disc glandspresent Disc glandsabsent Disc glandsabsent
Ovarylackinga central Ovarywith a central Ovarywith a central

column column column
Placentationon innerends Placentationaxile Placentationaxile

of the lateralsepta
Fibersnot libriform Fiberslibriform Fiberslibriform
Parenchymaapotracheal Parenchymaparatracheal Parenchymaparatracheal

Pith lackingsclereids Pith with sclereids No Data
Foliarbundlesimmediately Foliarbundlesextending Foliarbundlesextending

fused with the stele some distance(down the some distance(down the
stem)before fusingwith stem) before fusingwith
the stele the stele
Petiole epidermissimple Petiole epidermis No data

multipleand
collenchymatous
Hypodermisabsentin leaf Hypodermispresentin Hypodermispresentin

leaf someleaves
Palisadeparenchymaof Palisadeparenchymaof No data

1-2 stratifiedlayers 2-severalirregularly
disposedlayers
Usinganatomicalevidence, BarthconsideredEuphroniaa possibleintermedi-

ate between Trigoniaceaeand Dichapetalaceae,but at the same time noted that
anatomicallyit could be accommodatedin either family. Metcalfe& Chalk(1951)
observedthe anatomicalsimilaritybetween Trigoniaceaeand Vochysiaceae;several
importantanatomicalcharactersare sharedby Euphronia(considerdby Metcalfe
& Chalkas Trigoniaceae)and Vochysiaceaebut are not found in Trigoniaceae.For
example, both Euphronia and Vochysiaceaehave libriformfibers, paratracheal
parenchyma,and very small cells on the upperepidermisof the leaf. Intraxylary
phloem, the charactergivenmost emphasisby Metcalfeand Chalkto distinguish
Trigoniaceaeand Vochysiaceae, is not found in Euphronia (Heimsch, 1942).
Althoughintraxylaryphloem is very common in Vochysiaceae,it is not present
in all of that family (Metcalfe& Chalk, 1951). This holds true for many families
in which this characteroccurs;that is, it usually occursin a largenumberof species
or generaof a family, but not in all.
Morphologicalcomparisonbetween Euphroniaand Vochysiaceaeshows that
the only majordifferenceis in the numberand arrangementof stamensand stami-
nodes; the Vochysiaceae,as currentlydelimited,usuallyhas only 1 fertile stamen

and severalstaminodes,while Euphroniahas severalfertile stamensand one stami-
node. The staminodein Euphroniaoccupies the position of the fertile stamenin
Vochysiaceae.This comparativelysmall differencein stamennumberand arrange-
ment does not seem to me to be sufficient evidence to maintainEuphroniaapart
from Vochysiaceae.It is a relativelyminor evolutionarystep to changethe stamen
numberin responseto selective pressure.It is highly possible that Euphroniaand
Vochysiaceaehave divergedfrom a common ancestorall of whose stamenswere
fertile, but have divergedin degreesof reductionin the numberof stamens.
Duringthe course of my researchon the Trigoniaceae,I have come to agree
with Chodat (1895) and Barth (1896) that Euphroniadoes not belong in the
Trigoniaceae.I believethat the relationshipsof Euphroniaarewith the Vochysiaceae,
and consider that it is superfluousto create a new family to accommodatethis
unusualgenus. I propose to returnEuphroniato the Vochysiaceae,with which its
affinities are closest.
SPECIFICCONCEPTS
The conceptsusedto delimitspeciesin the Trigoniaceae,especiallyin Trigonia,

have been determinedby the data availablefor the family. As noted in the section
on Geographyand Ecology, it has proved difficult to determinepopulationsin
Trigonia(as is true for many other tropicaltaxa). The scarcityof collections, as
well as the lack of successin obtaininglive materialfor work in the laboratory,have
made it very difficult to obtain data that would provideclues to the biology of
Trigonia.Most of the evidencecited here and elsewherein this treatmentabout the
ecology and biology of Trigoniahas been obtained from label data and personal
observationson a few individualsof four species.
I choose here to consider the species recognizedin Trigoniaas taxonomic
species, a concept proposedby Grant(1963), primarilyon morphologicalevidence.
The data availableat presentare insufficient to establishall of the biologicaltaxa
in the genus. However,I am inferringthat the species proposedhere do (for the
most part) representself-perpetuatingpopulationsor groupsof populationsrepro-
ductivelyisolated from other such groups.
I have recognizedboth subspeciesand varietiesas infraspecificcategoriesin
Trigonia.Although these two termshave been the objects of continued discussions
(eg Grant, 1963; Cronquist,1968) no agreementhas been reachedas to the usageof
these terms. I have used the term subspeciesto separatetaxa that are allopatric
and almost completely distinct morphologically,but that I believe are elements of
the same species. The term varietyhas been used here to separatetaxa that are
geographicallysympatric or allopatric,and in which a morphologicaldistinction
is not alwayspossible.
TO THESYSTEMATICTREATMENT
INTRODUCTORYCOMMENTS
The following brief comments are given to aid the user of the key and to
clarify the descriptions.
Since sterile materialis not adequatelyrepresentedin collections, and sterile
shoots and vegetative branchesoften have much largerleaves, leaf size measure-
26 Flora Neotropica
ments are taken from fertile branchesonly.

In the species descriptionsand keys, the surfaceof the venation is treated
separatelyfrom the descriptionof the intercostalpubescence;thus, "leaf glabrous"
will referto intercostalsurfaces,but not to the venation unless otherwisestated.
The petals are treated using nomenclature applicable to papilionaceous
flowers, thus the termsstandard,keel, and wings;when the throat is mentioned
in relationto the standard,this indicatesall of the areasurroundingthe mouth of
the pouch (Fig 11).
The termpeduncle refersto the primaryaxis from which the pedicels of the
flowers arise.The term'panicleis here used to describeelongate mixed panicles,
often leafy-bracteatebelow. These inflorescenceshave the primarybranchesand
the terminalpart of the main axis well spaced, elongate, narrowand raceme-like.
Thyrseis used to describeinflorescenceshaving the primarybranchesand the
terminalpart of the main axis with evident ramifiedsecondarybranches.These
thyrsoidinflorescencesare more compact or dense than the panicles.
Dichasiumhere describesultimate inflorescences(or flower groups) having
what has traditionallybeen considered a complete or a compound dichasium.
Cymuleis here used to describemodified or reduceddichasiain which some of
the flowershave been lost. The term cincinnusis here appliedto helicoid cymes
(or flower groups)in which a spiralor helicoid arrangementof the flowers can be
inferredeven when only 2 or 3 flowers are present(Fig 11).
This descriptiveterminology for inflorescencesis not wholly in agreement
with the usage of other authors(Lawrence,1951; Rickett, 1944; Gray, 1875), as
the inflorescencesof the Trigoniaceaedo not fit any standarddefinition. The terms
used are thereforeslightly modified usagein orderto meet the needs for the family.
' '
?.
.. , :.. - . ,'? .i'.'..
"... ?: .
FIG 11. Some floraland inflorescencecharactersin Trigoniaceae.A. Globosestandardpetal.

B. Nasiform standard petal. C. Dichasium. D. Cymules common in Trigonia. E. Cincinni common
in Trigonia.
Laciniae,when used in respectto the floralglands, referto the fairly short,

often solitary, filamentousstructuresfound at the apex.
In species with few collections, all herbariummaterialexamined has been
cited; in common species, only selected specimensare cited in the text.
Abbreviationsused.
fl = flowering
fr = fruiting
st = sterile
bds = buds
Herbariaabbreviationsare those of Index Herbariorum(Holmgren& Keuken,
1974).
28 Flora Neotropica
SYSTEMATICTREATMENT
TRIGONIACEAE Endlicher,Enchir.Bot. 570. 1841; Martius,Conspec.51. 1835

(nomen nudum); Endlicher,Gen. P1. 5659. 1840 (nomen nudum);
Grisebach,Linnaea22: 27. 1849; Warming(Trigoniaceae)Mart. Fl.
Bras. 13(2): 122. 1875; Chodat,Bull. Herb.Boiss. 3: 136. 1895; Barth,
Bull. Herb. Boiss. 4: 481. 1896; Petersen, (Trigoniaceae)Engler&
Prantl,Nat. Pflanzenfam.3(4): 309. 1896; Standley, (Trigoniaceae)
North Am. Fl. 25(4): 297. 1924; Van Steenis, (Trigoniaceae),Fl.
Malesiana1(4): 58. 1949; Stafleu, (Trigoniaceae)Pulle, Fl. Suriname
3(2): 174. 1951; Perrier& Leandri,(Trigoniaceae)Fl. de Madagascar
108(bis): 1. 1955; Reitz, Fl. Ilustr.Catarinense1(13) 3: 1967; Austin,
(Trigoniaceae)Fl. Panama, Ann. MissouriBot. Gard. 54(3): 207.
1967; Ng, (Trigoniaceae)Tree Fl. Malaya1: 448. 1972.
Trigoniaceae tribe Trigoniae Chodat, Bull. Herb. Boiss. 3: 138. 1895.

Trigoniaceae tribe Trigoniastrae Chodat, Bull. Herb. Boiss. 3: 138. 1895.
TrigoniaceaetribeLigthiae(sic) Chodat,Bull. Herb.Boiss. 3: 138. 1895.
Flowers obliquely zygomorphic,hypogynous to subperigynous,plane of

symmetry through the third sepal, receptacle of varied shape and size, slightly
gibbousat base. Calyx gamosepalous,quincuncial,the base cupulate;sepalsimbri-
cate in bud, unequal.Corollapapilionacious;petals 5, contorted in bud; the two
anteriorpetals forminga keel, often saccate, the posterioror standardpetal saccate,
the two lateral petals or wings spathulate. Stamens 5-8, monadelphousand 0-4
staminodes, unilateral,opposite the keel petals; filamentaltube subperigynous;
anthersintrorse, 2-locular,longitudinallydehiscent. Pollen 34-porate, the exine
smooth or slightlyverrucose.Disc glandsopposite the standard,sometimeslaciniate.
Ovarysuperior,basicallytrilocular,seldom 4-locular,if 1-locular,then by reduction
of the parietalsepta;centralcolumn absent. Ovules 1-numerous,biseriate,anatro-
pous, attached to the interior ends of the lateral septa. Style terminal,simple;
stigmacapitate. Fruit a septicidalcapsuleor 3-wingedsamara.Seed exalbuminous;
embryo straight,longitudinalor transverseto the length of the seed, cotyledons
plane, thin, the radiclevery short.
Trees,treelets,shrubs,or scandentshrubs.Branchesterete, lenticellate. Leaves
simple, entire, alternateor opposite, often subcoriaceous,glabrousor pubescent,
pinnately veined. Stipules present, interpetiolar, often connate, deciduous or
caducous. Inflorescencesthyrses,paniclesor racemes,sometimesreducedto cymes;
flowers bibracteolate.
Twenty six species in three generafound in lowland wet forests or humid
riverineforests or forest edges in the Neotropicsand Paleotropics.
Type genus. TrigoniaAublet. The name is derived from the Greek words
Trias-(three) and Gonos-(angles),and refersto the three-sidedor angularfruit.
Key to the Generaof Trigoniaceae

1. Leaves opposite; New World and Madagascar.
2. Fruit a loculicidal capsule; ovary lacking lateral ridges; ovules more than one
per locule; tropical and subtropical America. 1. Trigonia.
SystematicTreatment 29
2. Fruit a three-winged samara; ovary with lateral ridges; ovules one per locule;
Madagascar. 2. Humbertodendron.
1. Leaves alternate; Malaysia. 3. Trigoniastrum.
1. TrigoniaAublet, Hist. P1.Guian.Fr. 387. 1775; H.B.K.,Nov. Gen. 5: 141.

1821; de Candolle,Prodr.1: 571. 1824; Cambessedesin St. Hil., Fl.
Bras. Mer. 2: 112. 1829; Endlicher,Gen. PI. 5659. supl. 4 (3): 82.
1840; Grisebach,Linnaea22: 27. 1849; Bentham& Hooker,Gen. P1.
1: 977. 1867; Baillon,Hist. P1.5: 103. 1874; Warming,(Trigoniaceae)
Mart. Fl. Bras. 13(2): 122. 1875; Barth, Bull. Herb. Boiss. 4: 481.
1896; Petersen, (Trigoniaceae)Engler & Prantl, Nat. Pflanzenfam.
3(4): 309. 1896; Standley, (Trigoniaceae)North Am. Fl. 25(4): 297.
1924; Stafleu, (Trigoniaceae)Pulle Fl. Surin.3(2); 174. 1951; Reitz,
(Trigoniaceae)Fl. Ilust. Catar.1(13): 3. 1967; Austin, (Trigoniaceae)
Fl. Panama,Ann. MissouriBot. Gard.54(3): 207. 1967.
Hoeffnagelia Necker, Elem. Bot. 3: 68. 1790.

Nuttallia Sprengel,Neue Entdeck. 2: 158. 1821.
Mainea Vellozo, Fl. Flum. 275. 1829.
Treelets,shrubs,or scandentshrubs,the branchesterete, and conspicuously

lenticellate. Leavessimple, opposite, petiolate, the marginsentire; stipules inter-
petiolar, deciduousor caducous,often connate, simple or bifid at the apex. Inflor-
escencesthyrses,paniclesor racemes.Flowersultimately disposedin dichasia(simple
or compound), cymes or cincinni;bractsglandularor eglandularat the margins;
sepals 5, quincuncial;unequal;petals 5, contorted, unequal,papilionaceous,fertile
stamens5-8, staminodespresentor absent,when presentto 4, the filamentsconnate
at base, the anthersbasifixed, bilocular,introrse, dehiscingalong a central slit;
pollen 3-4 porate, smooth; glandsusually2, lobed, sometimeslaciniate;ovary 1-,
3- or 4-locular,when 1-locularby reductionof the lateralsepta, these when present
fused or open at the center;centralcolumn absent;ovules numerousto few in each
locule, biseriate,anatropous,attachedto the interiorends of the lateralsepta. Fruit
a septicidalcapsule, dehiscing(in situ) from the apex towardsthe base. Seeds 2 to
severalper locule, pubescent.
Type species. Trigoniavillosa Aublet
Distribution.SouthernMexico to northernParaguay,usuallyin low-altitude
forests. The pubescentseeds, suggestiveof wind dispersal,as well as collection data
and personalobservations,seem to indicate that this genusis restrictedto open
forest edges or galleryforests. It is usually recordedfrom rivermargins,edges of
clearings,and along roadsidesand other disturbedareas.
Key to the Species of Trigonia

1. Leaves rotund or subrotund, length/breadth ratio ca 6:5-6:4.
2. Branches densely golden-strigose; inflorescence a thyrse; flowers disposed
in dichasia; peduncles 4-8 mm long; bracts and bracteoles navicular.
1. T. coppenamensis.
2. Branches white-lanate, becoming glabrous; inflorescence a panicle; flowers
30 Flora Neotropica
disposed in cincinni;peduncles0-1.5 mm long; bractsand bracteolessub-

ulate or linear. 2. T. rotundifolia.
1. Leavesovate to obovate, elliptic or oblong-elliptic,length/breadthratio ca 2:1.
3. Leavesglabrousbeneath.
4. Ultimateinflorescencesdichotomousdichasia;wingpetalsglabrousat
base;glandslaciniate. 3. T.prancei.
4. Ultimate inflorescencescymules or cincinni;wing petals barbateat
base;glandsnot laciniate.
5. Flowersdisposedin cymules;standarderect;style glabrous.
6. Petioles 5-8(-10) mm long;bracts1.5-3 mm long; peduncles
2-4 mm long; stamens8, staminodesabsent;ovaryuniloc-
ular. 4. T. virens.
6. Petioles(10-)12-18mm long;bracts34 mm long;peduncles
0-1 mm long; stamens 10-12; staminodes present; ovary
trilocular. 5. T. macrantha.
5. Flowers disposed in cincinni; standardrevolute; style villous.
7. Secondary leaf veins 4-7 pairs;standardpouch nasiform;
stamens 8-9, staminodes present; fruit 0.6-3 cm long;
seedsvillous. 6. T. laevis.
7. Secondary leaf veins 9-11 pairs; standardpouch globose;
stamens 6-7, staminodes absent; fruit 7-9 cm long; seeds
echinate. 7. T. spruceana.
3. Leavespubescentbeneath.
8. Inflorescenceof thyrses.
9. Flowersin dichasia.
10. Leaf marginsrevolute;tertiaryand quaternaryleaf venation
veryevident;bracts,bracteolesand sepalspapillose-glandular.
8. T. reticulata.
10. Leaf marginsplane; tertiary and quaternaryvenation not
especiallyevident; bracts, bracteolesand sepals not papil-
lose-glandular.
11. Leaf pubescence tomentellous (sometimes almost
glabrous); wing petals glabrous at base; staminodes
lacking appendages,anthers 0.8-1.5 mm long, glands
laciniate;fruit 6.5-8 cm long, velutinous-tomentose.
9. T. candelabra.
11. Leaf pubescence adpressed-lanate(sericeous) some-
times almost glabrous;wing petals barbate at base;
staminodes often appendiculate;anthers 0.4-0.5 mm
long;glandsnot laciniate;fruit 2-3.2 cm long, glabrous.
10. T. rugosa.
9. Flowersdisposedin cymulesor cincinni.
12. Branches densely golden-strigose,lenticels never visible;
bractsand bracteolesnavicular. 1. T. coppenamensis.
12. Branches not densely golden-strigose,becoming glabrous
with age, lenticelsvisible;bractsandbracteolesnot navicular.
13. Standardpouch nasiform;antherssubglobose,0.2-0.3
mm long, staminodesappendiculate;petiole 2-5 mm
long; (thyrsesoften reducedto paniclesor racemes).
11. T. eriosperma.
13. Standardpouch globose; anthers elliptic, 0.8-1 mm
long; staminodesnot appendiculate;petioles5-10 mm
long; (thyrsesneverreducedto paniclesor racemes).
14. Leaveswithmixedintercostalpubescence beneath,
lanate & strigose;pedunclesabsent or nearly so;
bractsand bracteoles5-10 mm long;sepalspapil-
lose-glandular at the margins;style villous.
12. T.floccosa.
14. Leaveslanate, with only 1 type of pubescence;
peduncles24 mm long;bractsandbracteolesless
than 5 mm long; sepalsnot glandular;style glab-

rous. 13. T. costanensis.
8. Inflorescencespaniclesor racemes.
15. Leavespubescentabove.
16. Flowersdisposedin cymules.
17. Inflorescencesterminalpyramidalpanicles;wingpetals
glabrousat base;stamens8. 14. T. subcymosa.
17. Inflorescences terminal and axillary panicles and
racemes,not pyramidal;wing petals barbateat base;
stamens9-11.
18. Secondaryleaf veins4-7 pairs;staminodesappen-
diculate;glandslackinglaciniae. 11. T. eriosperma.
18. Secondaryleaf veins 10-16 pairs;staminodesnot
appendiculate;glandslaciniate. 15. T. nivea.
16. Flowersdisposedin cincinni.
19. Secondaryleaf veins 14-16 pairs. 16. T. killipii.
19. Secondaryleaf veins4-12 pairs.
20. Leavesoblongto elliptic,the leaf pubescencedark
olive-greenbeneath (or seemingly so) in exsic-
catae, distinctly white-lanateat the margins;fruit
truncateat base. 17. T.paniculata.
20. Leavesbroadlyelliptic to obovate,the leaf pubes-
cence light yellowish-greenor white beneath in
exsiccatae, glabrous at the margins;fruit not
truncateat base.
21. Bracts 2-3 mm long; standard revolute;
stamens 10-11; style 2.5-2.8 mm long; fruit
4.5-11 cm long; the valves not cornate at
apex. 18. T. villosa.
21. Bracts3-5 mm long;standarderect;stamens
8-9; style 1.9-2.1 mm long; fruit 2-2.5 cm
long, the valvesdistinctlycornateat the apex.
19. T. boliviana.
15. Leavesglabrousabove.
22. Staminodesabsent;inflorescences,when terminal,borneon
shortaxillarybranches;whenaxillary1 per node, unilateral.
4. T. virens.
22. Staminodes present; inflorescences, when terminal, not
borne on short axillarybranches;when axillary2 per node,
bilateral.
23. Flowersnoticeablycaducous,the bractsand bracteoles
thenformingrosetteson the axes;bractsand bracteoles
the wingpetalsglabrousat base.
papillose-glandular;
20. T. bracteata.
23. Flowersnot noticeablycaducous,bractsand bracteoles
not formingrosetteson the axes;bractsand bracteoles
not papillose-glandular;wing petals barbateat base.
24. Leavessparselyarachnoid-pubescent beneath.
25. Petioles 12-18 mm long;flowersdisposedin
cymules,bractsneverlaciniate,34 mm long,
not sheathing the peduncles or pedicels;
anthers 0.8-1 mm long; fruit 7-9 cm long;
exocarpvelutinous-tomentose. 5. T. macrantha.
25. Petioles 5-10(-12) mm long; flowers dis-
posed in cincinni; bracts often laciniate;
1-2.5 mm long, sheathingthe pedunclesand
pedicels;anthers0.2-0.3 mmlong;fruit2.8-3
cm long;exocarpglabrous. 21. T. rytidocarpa.
24. Leaves lanate or adpressed lanate (sericeous)
beneath.
32 Flora Neotropica
26. Ovary 1-locular;staminodesappendiculate;

fruit with the exocarp flush with the endo-
carp. 22. T. hypoleuca.
26. Ovaryimperfectly3-locular;staminodesnot
appendiculate;fruitwith the exocarpshorter
than the endocarp.
27. Flowersdisposedin cymules.
28. Leaves densely brownishyellow-
lanate beneath; peduncles 0.5-1
mm long;bractssubulateorlinear,
2.5-3.5 mm long; glandsnot laci-
niate; ovary with septa not fused
at center. 23. T. echiteifolia.
28. Leaves white adpressed-lanate
beneath;peduncles2.54 mm long;
bracts ovate, 1.5-2 mm long;
glands laciniate;ovarywith septa
fused at center. 24. T. sericea.
27. Flowersdisposedin cincinni.
29. Stamens 6-7, all fertile; glands
never laciniate; style 2.8-3 mm
long, villous; fruit 7-9 cm long;
seeds echinate pubescent; leaves
sometimesbecomingglabrous.
7. T. spruceana.
29. Stamens 10-11; staminodespres-
ent; glands often laciniate; style
1-2 mm long, glabrous;fruit 3-7
cm long, seedsvillous. 15. T. nivea.
1. TrigoniacoppenamensisStafleu, Rec. Trav.Bot. Neerl. 42: 70. 1950; Stafleu,

Med. Bot. Mus.Utr. 105: 70. 1950; Stafleu, in Pulle Fl. Surin.3(2):
176. 1951.
Scandent shrub, the branches striate, (almost angular), not conspicuously

lenticellate, ferruginous-strigose. Stipules 4.0-7.0 mm long. 2.0-3.0 mm wide,
triangular, strigose; petioles 10.0-22.0 mm long, ca 1.0 thick, striate to terete,
strigose; lamina elliptic to obovate, sometime almost circular, 5.5-15.0 cm long,
2.5-12.0 mm wide, subcoriacous, the margins entire, the apex acute to acuminate,
the base subrotund; venation eucamptodromous, densely golden strigose (including
tertiary veins beneath), the midrib prominulous above, prominent beneath, second-
ary veins 6-9 pairs, plane above, prominulous beneath, the tertiary nervation im-
pressed above, prominulous beneath. Inflorescences terminal and subterminal
axillary thyrses, to 12 cm long. Flowers in dichasia of 4-6; peduncles 4.0-8.0 mm
long, ca 0.5 mm thick, strigose; pedicels 2.5-5.5 mm long, ca 0.5 mm thick, strigose,
the bracts and bracteoles of equal size, 3.0-6.0 mm long, 1.0-2.0 mm wide, linear-
ovate (almost navicular), revolute at the margins, curved upwards towards the apex
of the flower, strigose. Flower buds to 6.0 mm long; sepals ovate to oblong, ca 5.0
mm long, thickened at the nerves (appearing striate), strigose; standard with the
pouch extending to /2 of the length, barbate at the throat, the wings spathulate,
barbate at the base, the keel petals saccate; stamens 9-10, 6 fertile, staminodes
3-4, the anthers with a short knob-like projection at the apex; glands 2, 2-3-lobed;
style glabrous;the stigmatrilobate;ovarysubglobose,denselyvillous, the ovules

numerous.Fruit unknown.
Type. Maguire24857, Suriname,CoppenameRiverHeadwaters,Mt. Schmidt,
buds (holotype U; isotypes F, NY).
Distribution.Known only from the type gathering.
This very distinct species can be recognizedeasily by its striatestems, petioles
and sepals,as well as by its very largenavicularbractsand bracteoles.
Thereis insufficient data to indicate the relationshipsfor this species. Stafleu
has suggestedan affinity with Trigoniasubcymosabased on the inflorescenceand
bracts,but without knowledgeof the structureof the fruits, which are still unknown
for the two species, it is prematureto assignrelationships.
2. TrigoniarotundifoliaLleras,sp nov Fig 12.
Rami juveniles adpressi-lanati,glabrescentes,lenticellati. Stipulae caducae

(nv). Folia opposita, petiolo 0.7-3.0 mm longo, ca 1.0 mm crasso,adpresso-lanato,
glabrescenti;laminaerotundae,6.0-15.0 cm longae, 5.0-11.0 cm latae, chartaceae,
margineintegrae, apice rotundato usque emarginatovarianti,basi obtusa, supra
glabrae,infra adpresso-lanatae;costa media supra plana,glabra,infraprominula,
adpressa-lanata, costis secundariis6-9-jugis,eucamptodromis.Inflorescentiaein pani-
culis terminalibusvel axillaribusdispositae,5.0-15.0 cm longae. Floresin cincinnos
1-2 (raro 3) dispositi;pedunculi 0.0-1.5 mm longi, 0.5-0.7 mm crassi, strigoso-
tomentosi, bracteis2.5-3.0 mm longis, ca 0.4 mm latis, strigosis;pedicelli 1.0-2.5
longi, 0.4-0.6 mm crassi,strigoso-tomentosi,bracteolissubulatis,1.0-2.0 mm longis,
strigosis;sepala ovata vel leviter oblonga, 3.5-4.2 mm longa, 1.5-2.0 mm lata,
strigoso-tomentosa;vexillum 5.0-5.5 mm longum, 2.0-2.8 latum, usque ad medium
longitudinalitersaccatum,apicerevolutum,intus barbato-tomentosum,alaeconcavo-
spathulatae,5.0-5.2 longae, 1.0-1.5 mm latae, ad basin barbatae,carinaepetala
saccata,3.5-4.0 mm longa, 2.0-2.2 mm lata, apice cristato;stamina9-10, fertilia
6-7, sterilia3-4, appendiculata,filamentisad mediumconnatis, 1.8-2.2 mm longis,
antherisoblongis, 0.8-1.1 mm longis, (0.3-)0.4-0.6 mm latis, castaneis;glandulae2,
bilobae, reniformes,ca 0.5 mm longae, 0.8 mm latae, lanatae;stylus erectus,
strigosus, 1.8-2.1 mm longus, stigmate circulari,albo, ca 0.4 mm lato; ovarium
subglobosum,ca 1.0 mm diametro,4-loculare,ovulisin quoque loculo numerosis.
Fructusmihi ignotus.
Type. L. B. Smith &McWilliams15441, BrazilRio de Janeiro,Mangaratiba,
fl (holotype MO;isotype US).
Distribution.Known only from the type gathering.
This distinctive species is characterizedby its almost round leaves, and
especiallyby the 4-locularovary.At presentit is impossibleto establishits relation-
ships within the genus, as no fruitingmaterialis availablefor comparison.
3. TrigoniapranceiLleras,sp nov Fig 13.
Frutex, ramulisjuvenilibustomentellis,glabrescentibus,lenticellatis.Stipulae
caducae,ca 5.0 mm longae, ad basinconnatae. Folia opposita,petiolo 8.0-15.0 mm
longo, 0.8-1.5 mm crasso, leviter tomentello; laminae ellipticae vel obovatae,
6.0-11.0 cm longae, 3.0-5.8 cm latae, subcoriaceae,margineintegrae,apice acuto
34 Flora Neotropica
Vs.2
F G
D E
J K L
FIG 12. TrigoniarotundifoliaA, habit X0.35; B, crosssection of ovaryXI 3.2; C, flower X3.3; D,
sepalsX3.3; E, ovaryX3.3; F, stamensX3.3; G, anthersX6.6; H, glandsX6.6; J, keel petal X3.3;
K, wingpetal X3.3; L, standardX3.3.
J?
FIG 13. TrigoniapranceiA, habit X0.35; B, flower X3.3; C, ovaryX3.3; D, keel petal X3.3; E,
wingpetal X3.3; F, standardX3.3; G, stamensX3.3; H, anthersX6.6; J, crosssection of ovaryX13.2.
36 Flora Neotropica
usque acuminatovarianti,basi obtusa vel obliqua,glabrae;costa media supraplana,

infra prominens,strigulosa,costis secundariis5-6-jugis,eucamptodromis.Inflores-
centiae in thyrsos terminalesvel axillaresdispositae,5.0-15.0 cm longae. Floresin
cymis 1-bifloris;ad extremum in inflorescentiassecundariasdispositi, a dichasiis
dichotomisusque cymis solitariisvariantes;axes inflorescentiarumsecundariarum
1.5-4.0(-7.0) mm longi, 1.0-3.0 mm crassi,tomentelli;pedunculi0.5-2.0 mm longi,
0.2-0.3 mm crassi, tomentelli, bracteis ovatis, 1.5-2.0 mm longis, tomentellis,
caducis;pedicelli 1.5-2.0 mm longi, 0.2-3.0 mm crassi,tomentelli, bracteolisovatis,
0.8-1.3 mm longis, tomentellis;sepalaovatavel leviter oblonga,3.44.0 mm longa,
1.3-2.2 mm lata, tomentellavel strigulosa;vexillum4.04.8 mm longum, 2.0-2.5
mm latum, usque ad medium longitudinalitersaccatum, apice erectum, intus
barbatum;alae spathulatae,3.5-4.3 mm longae, 1.0-1.8 mm latae, glabrae,carinae
petala saccata, 3.2-3.5 mm longa, 1.1-2.0 mm lata; stamina 8-10, sterilia 2-4,
fertilia6, filamentisusque ad 2/3 connatis, 1.2-1.5 mm longis, antherisoblongis,
0.5-0.7 mm longis, 0.2-0.3 mm latis; glandulae2, bilobae, reniformes,laciniatae,ca
0.3 mm longae, 0.4 mm latae; stylus erectus, 1.5-1.8 mm longus, villosus usque
glabervarians,stigmatecirculari,albo, ca 0.3 mm lato; ovariumsubglobosum,ca
1.0 mm diametro,3-loculare,ovulisin quoque loculo numerosis.Fructus8.5-9.0
cm longus, ca 1.2 cm latus, extus velutino-tomentosus,intus cartilagineus,glaber.
Seminaca 20 in quoque loculo, subglobosa,ca 3.0 mm diametro,pilis ca 7.0 mm
longis.
Type.Lleras,Stewardet alP 7115, Peru,Loreto,Rio Javari,AngamoGarrison,
fl, fr (holotype INPA;isotypes COL,F, K, MG,MO,NY, US).
Distribution.Known only from the type collection, which was from non-
flooded groundin an open disturbedareaat the edge of the forest.
This species is probably closely related to Trigoniasericea from which it
differs in the type of inflorescence(thyrses, ratherthan panicles in T. sericea),
shorterpeduncles,smallerbractsand bracteoles,largerfruit, and glabrousleaves, as
well as other minor characters.
I am dedicatingthis species to Dr. GhilleanT. Prance,underwhose auspices
the type materialwas collected and whose help and guidancein this study have
been invaluable.
4. TrigoniavirensMacbride,Publ. Field Mus.Nat. Hist. Bot. 11(2): 68. 1931;

Smith, Phytologia3: 130. 1935.
TrigoniacrassifloraA. C. Smith,Phytologia3: 129. 1935.
Scandent shrub, the branchesterete, lenticellate. Stipules caducous (nv);

petioles 5.0-8.0(-10.0) mm long, 1.0-2.0 mm thick, densely strigose;laminaelliptic
to broadlyelliptic, 8.0-16.0 cm long, 4.0-9.0 cm wide, chartaceous,smooth, the
marginsentire, the apex acute to acuminate,the base acute to obtuse;venation
eucamptodromus,slightly strigose-pubescent,the midribplane above, prominulous
beneath, the secondarynerves6-10 pairs;intercostalpubescenceabsent on both
surfaces, rarely with arachnoidpubescence beneath. Inflorescencescondensed
terminaland axillarypanicles,when axillary,presentonly associatedwith one of
the leaves of a pair, 2.0-15.0 cm long. Flowersin groupsof 1-3 (usually2); ped-
uncles and pedicels of equal length, 2.0-4.0 mm long, 0.7-0.9 mm thick, densely
strigose, the bracts 1.5-3.0 mm long, ovate, strigose, the bracteoles 1.0-2.5 mm
long, narrowlyovate, strigose;sepalsovate to deltoid, the inner ones usuallynar-

rower, 4.0-5.5 mm long, 2.5-5.0 mm wide, the exposed portions strigose, the
protected portions white-lanate-pubescent; standard5.0-6.0 mm long, 4.0-5.0 mm
wide, the pouch extending 2/3 of the length, the upperportion erect, the apex
irregular,barbate-pubescenton the inside of the throat, the wings spathulate,
4.0-5.0 mm long, 3.0-3.5 mm wide, barbateat the inside of the base, the keel
petals 4.0-5.0 mm long, 3.0-4.0 mm wide, the pouch extending down, the apex
irregular;stamens8, all fertile, the filaments 1.5-2.0 mm long, free at apex, the
anthersovate-elliptic,0.6-0.8 mm long, 0.4-0.5 mm wide, adhering4 on each side
of the style;glands2, 3-lobate,ca 0.9 mm high, 2.0 mm long, glabrous;style clavate,
1.8-2.0 mm long, ca 0.5 mm thick at the apex, the stigmatrilobate,white; ovary
ovoid, 1.0-1.2 mm high, ca 1.5 mm wide at base, barbate-pubescent,1-locular,the
ovules numerous.Fruit 7.0-8.0 cm long, the valvesca 10.0 mm per side;exocarp
thin, coriaceous,glabrous;endocarpcartilaginous.Seeds not seen.
Type. Killip & Smith 29539. Peru, Loreto, fl (holotype US; isotypes F, G, NY).
Distribution.Growingat low altitude,in humid periodicallyflooded forests
in Colombia,Peruand Brazil.
PERU. Loreto: Asplund 14281 fl (G, K, NY, P, US); Williams2395 fl (F, G). BRAZIL.
Amazonas: Krukoff 6149 fl (BR, F, G, K, M, NY, U, US, W); Krukoff 8424 fr (A, BR, G, MICH,
MO, NY, P, U, US). Parai:Black & Ledoux 501680 fl (IPEAN). COLOMBIA. Santander:
Haught1800 fl (COL);Haught1907 fl (US);Haught2211 fl (F, US).
This speciesis easily distinguishedby its large,glabrousleaves,its unilateral,
condensedinflorescences,the presenceof a clavatestyle, and the unilocularovary.
Trigoniacrassiflorais not taxonomicallydistinct and thus is here treated as a later
synonym.
5. TrigoniamacranthaWarming,(Trigoniaceae)Mart.Fl. Bras.13(2): 129. 1875.
Shrubor small tree, the branchesterete, lenticellate. Stipuleslorate, 6.0-8.0

mm long, 0.6-0.9 mm wide, tomentose, caducous;petioles 12.0-18.0 mm long,
0.6-0.8 mm wide, slightly tomentose or sometimes glabrous,lamina elliptic to
ovate-elliptic,9.0-13.0 cm long, 3.5-7.0 cm wide, chartaceous,slightly rugulose,the
marginsentire, the apex acuminate,the base obtuse;venationeucamptodromous,
slightly strigosepubescent,the midribplane to sunkenabove, prominulousbeneath,
the secondarynerves6-9 pairs;intercostalareasarachnoid-pubescenton both sur-
faces, sometimesbecomingentirely glabrous.Inflorescencesterminaland subtermi-
nal axillarypanicles,sometimesreducedto racemes,the subterminalones 34 pairs.
Flowersin cymules of 1-4 (usually 2-3); pedunclesand pedicels of equal length, to
1.0 mm long, tomentose, the bractslorate or subulate,3.04.0 mm long, 0.6-1.3
mm wide, the bracteolesnarrowlyobovate to subulate, 1.2-2.3 mm long, 0.3-0.6
mm wide, tomentose;sepals ovate to deltoid, 3.0-5.5 mm long, 2.04.0 mm wide,
tomentose, the apex rounded or acute, the base truncate;standard5.0-6.0 mm
long, 4.0-5.0 mm wide, the pouch extending to ?2the length, the upperportion
erect, the apex slightly irregular,barbate-pubescentat the inside of the throat, the
wings spathulate,the upperportion slightly convex, 3.0-4.5 mm long, 1.3-1.6 mm
wide, barbate-pubescenton the inside of the base, the keel petals 3.0-4.0 mm long,
the pouch extending along 1/3 of the length, the apex slightly irregular,barbate
pubescentat base;stamens 10-12, 6-7 fertile, with 4-5 staminodes,the filamentsto
1.5 mm long, free alongV?of their length, the antherselliptic, 0.8-1.0 mm long,
38 Flora Neotropica
0.4-0.5 mm wide; glands2, 1-lobed,slightly trapezoid,ca 1.0 mm high, 1.0 mm

long, fused to the staminodes;style 1.5-1.8 mm long, glabrous,the stigmatrilobate,
white, ca 0.4 mm in diameter,ca 0.3 mm high; ovarysubglobose, 1.0-1.4 mm in
diameter,barbate-pubescent,the ovules numerous. Fruit 7.0-9.0 cm long, the
valvesca 10.0 mm per side; exocarp thin (ca 0.5mm), fleshy, with a greenish-gray
velutinous-tomentosepubescence;endocarpcartilaginous.Seeds ca 20 per locule,
ovoid, 2.0-3.0 mm in diameter,barbate-pubescent,the trichomesto 10.0 mm long.
Type. Spruce 3871. Peru, Loreto, "Yurimaguasad flumen Huallaga,"fl
(lectotype W;isotypes BR, CGE,G, NY, OXF).
Distribution.An endemic species known only from the vicinity of the type
locality; found in rainforestat low elevationbetween Yurimaguasand Balsapuerto
on the Huallagariverbasin.
PERU. Loreto: Killip& Smith28136 fr (NY, US);Klug2954 fl (A, F, G, GH,MO,NY,
US); Klug3028 fl (A, F, G, GH,MO,NY, US).
I am designatingthe Vienna (W) specimenas lectotype, as there is no speci-
men deposited at Copenhagenand the W materialwas seen by Warming.
6. Trigonialaevis Aublet, Hist. PI. Guian.Fr. 1: 390 pl. 150. 1775; de Candolle,
Prodr.1: 571. 1824; Warming,(Trigoniaceae)Mart.Fl. Bras. 13(2): 131. 1875.
TrigoniakaieteurensisMaguire,Bull.TorreyBot. Club.75: 399. 1948.

Type. Maguire& Fanshawe23192, KaieteurPlateau,Guyana,fr (holotypeNY;
isotypesA, BR, F, MO,NY, P, US, VEN).
Scandentshrub,the branchesterete, lenticellate, the young ones strigulose,

becomingglabrouswith age. Stipules 1.5-2.0 mm long, subulate,strigulose;petioles
4.0-8.0 mm long, terete to canaliculate,strigoseor glabrous;laminabroadlyelliptic
to oblong-elliptic,sometimeslaterallyunequal,4.0-9.0 cm long, 2.24.5 cm wide,
chartaceousto subcoriaceous,the marginsentire, the apex acute to acuminate,the
base oblique to obtuse;venationeucamptodromous,glabrousor slightly pilose, the
midribprominulousabove, prominentbeneath, the secondaryveins 4-6(-7) pairs,
prominulouson both surfaces;intercostalpubescenceabsent. Inflorescencestermi-
nal and subterminalpanicles(sometimesthyrses), to 30 cm long. Flowersin groups
(cincinni) of 1-7; peduncles0.2-0.5 mm long, strigose,the bracts 1.0-3.0 mm long,
subulate to linear, strigose;pedicels 0.8-2.0 mm long, strigose, the bracteoles
0.3-1.0 mm long, subulateto linear,strigose;sepals deltoid to oblong, 2.0-3.0 mm
long, 0.8-1.5 mm wide, strigose;standard2.5-5.0 mm long, 2.0-3.0 mm wide, the
pouch extending along 2/3 of the length, barbateat the throat, the wings 2.5-5.0
mm long, spathulate,0.5-1.0 mm wide, barbateat the base, the keel petals 2.0-3.5
mm long,1.5-2.0 mm wide, sometimeslanate at the apex; stamens8-9, 5-7 fertile,
staminodes1-3, the filaments0.7-1.2 mm long, connate for 2/3 of the length, the
antherssubglobose,0.4-0.5 mm long, ca 0.4 mm wide, the staminodessometimes
with a small, terminal,laciniateappendage;glands2, 2-3 lobed, irregular,ca 0.3 mm
per side, glabrous;style slightly villous, 1.0-1.3 mm long, the stigmatrilobate,ca
0.2 mm in diameter;ovarysubglobose,0.6-0.8 mm in diameter,villous, the ovules
numerous.Fruit oblong or obovate, 0.6-3.0 cm long. 0.6-1.0 cm per side; exocarp
thin, yellow-velutinouswhen young, becomingglabrouswith age;endocarpwoody,
glabrous.Seeds 1-4 per capsule,subglobose,ca 0.3 mm in diameter,barbatevillous,
the trichomesca 10.0 mm long.
Systematic Treatment 39
This species is separatedinto two varieties.
Key to the Varietiesof Trigonialaevis
1. Flowers in groups(cincinni)of 1-2(-3);fruit oblong, 1.5-3.0 cm long. 6a. varlaevis.

1. Flowersin groups(cincinni)of (2-)3-6;fruit obovate,0.6-1.0 cm long. 6b. varmicrocarpa.
6a. Trigonialaevisvarlaevis
Flowersin groupsof 1-3; petals 2.5-3.5 mm long. Fruit oblong, 1.5-3.0 cm
long, ca 1.0 cm wide; seeds usually3-4 per locule.
Type. Aublet sn, French Guiana,Cayenne, fl, fr (holotype BM, isotypes
F,P).
Distribution.Known only from a few collections in FrenchGuiana,and one
collection (type of Trigoniakaieteurensis)from Guyana.
FRENCH GUIANA. Gabriel 1802 fl, fr (G); Soubirou sn fl, fr (P).
Withoutfruitingmaterial,it is practicallyimpossibleto separatethis variety
from varmicrocarpa.It is quite possible that some materialplaced by me in var
microcarpamight actually be varlaevis.
6b. Trigonialaevisvar microcarpaSagot, Ann. Sci. Nat. 6(2): 176. 1881.
TrigoniaparvifloraBentham,Hook. Jour. Bot. 3: 163. 1851. Type.

Sprucesn, Brazil,Para,Santar6m,fl, (holotype K; isotypesC, CGE,F,
G, GH, GOET,M, NY, P, W),nomen illegit.
TrigoniamicrocarpaSagot exWarming,(Trigoniaceae)Mart.Fl. Bras.13(2):
131. 1875; Stafleu,(Trigoniaceae)PulleFl. Surin.3(2): 174. 1951.
Type. Sagot 36, FrenchGuiana,Karouany,fl, fr (holotype P; isotypes
BR, F GH, GOET,P, W).
TrigoniabicolorSuessenguth& Overkott,Fedde Rep. Sp. Nov. RegniVeg.
51: 204. 1942. Type. Buchtien1631, Bolivia,Mapiri,fl (lectotype F;
isotype nv US).
Flowers in groups of 3-6; petals usually 3.5-5.0 mm long. Fruit obovate,

0.6-1.0 cm long, as wide as it is long; seeds usually 1-2(-3) per locule.
Type. Sagot 36, FrenchGuiana,Karouany,fl, fr (holotype P; isotypes BR,
F, GH, GOET, P, W).
Distribution.A rainforestvariety found from FrenchGuianato Bolivia,but
not yet reportedfrom Colombia.
VENEZUELA.DeltaAmacuro:Blanco426 fr (NY, VEN);Steyermark87550 fl (M, NY,
VEN, W);Wurdack& Monachino 39697 fr (F, NY, U, US). GUYANA. Fanshawe 4573 fl (U);
Sandwith 563 fl, fr (G, NY, P, U, US); A. C. Smith 2752 fl (A, G, MO, NY, U, US); Schomburgk
953 fl (G). SURINAME.Lan/ouw& Lindeman2416 fr (IPEAN,NY, US);Lindeman5785
fl (U); Stahel & Gonggryp 631 fl (U); van Donselaar 2894 fr (U). FRENCH GUIANA. Geav
1890 fl (P);Lambert sn fl (BR);Martin sn fl (P);Medlinon230 fr (P);Poiteau sn fl, fr (G, US).
PERU.Spruce4944 fl (BR, C, CGE,F, G, GOET,K, OXF,W). BRAZIL.Amazonas:Ducke
1208 fl (GH,IPEAN,MG,MO,NY, US);Ducke 1282 fl (F, GH, IPEAN,MG,NY, US). Para:
Black 4 7-932 fl (IPEAN, U); Fr6es 32914 fl (IPEAN, U). Rondonia: N. T. Silva 439 fl (IPEAN,
U). BOLIVIA.La Paz:Krukoff10338 (A, F, G, MICH,MO,NY, U, US); Krukoff10956 fr (A,
F, G, MICH,MO,NY U); Rusby 1220 fr (MICH,NY);Rusby2449 fl (E, G, GH,MO,NY, P,
US).
This widespreadvarietyincludes the taxa formerlyknown as Trigoniamico-
carpa, T. parvifloraBentham,and T. bicolor which are later synonyms. Although
40 Flora Neotropica
there is a tendency for an increaseof size of the inflorescencegoing from East to

West,I do not considerthis significantenough to merit taxonomic recognition.
TrigoniaparvifloraBenthamwas, when published, an illegitimatename, as
this name had been publishedby Schott in 1827; the name T. bicolor is clearly
superfluous,but must be used if T. parviflorais recognizedas a separatetaxon.
7. TrigoniaspruceanaBenthamex Warming,(Trigoniaceae)Mart. Fl. Bras.

13(2): 130. 1875.
Scandent shrub, the branchesterete, glabrous,lenticellate. Stipules deltoid

or subulate,0.8-2.0 mm long, 0.7-1.0 mm wide, pilose, caducous;petioles
(5.0-)6.0-12.0 (-13.0) mm long, 0.9-2.1 mm thick, sparselystrigose,laminaelliptic
to ovate-elliptic,sometimesovate, 5.0-14.0 cm long, 2.0-5.0 cm wide, subcoriaceous,
the marginsentire, the apex acute or acuminate,the base oblique;venationbrochi-
dodromousor seemingly so, sparselystrigose;the midrib prominulousabove,
prominentbeneath, the secondaryveins 9-13 pairs;intercostalpubescenceabsent
above, frequentlycaducousbeneath, sometimesbecomingentirely glabrous.Inflor-
escences terminaland subterminalaxillarypaniclesto 15 cm long. Flowersin groups
(cincinni) of 1-4 (usually 1-2), solitary on the axes of the ultimate inflorescences,
the axes 0.0-4.0 mm long, 0.6-0.8 mm thick, decreasingin length towardsthe apices
of the primaryinflorescences;peduncles 0.2-2.5 mm long, 0.6-0.8 mm thick,
strigose, often resemblingan articulatedpeduncle with the axes of the ultimate
inflorescences,the bracts2.0-3.0 mm long, 0.2-0.3 mm wide, subulateto linear,
strigose, caducous;pedicels 1.5-2.5 mm long, 0.6-0.8 mm thick, strigose, the
bracteoles0.7-1.4 mm long, 0.2-0.6 mm wide, deltoid or subulate,strigose;sepals
ovate or oblong, (4.0-)4.5-5.1(-5.8) mm long, 1.8-3.0 mm wide, tomentellousor
lanate;standard6.0-7.5 mm long, 2.0-3.5 mm wide, the pouch globose, extending to
2/3 of the length, revoluteat the apicalportion, barbateat the throat;the wings
spathulate, 5.8-6.2 mm long, 3.4-4.0 mm wide, barbateat the base, keel petals
5.0-6.0 mm long, 3.0-4.0 mm wide, the pouch excentric, barbateat the base;
stamens6-7, all fertile, the filamentsconnate for ?-1/3 of their length, 3.0-3.5 mm
long, the antherssubglobose, 0.8-1.0 mm long, 0.5-0.8 mm wide; glands2, 2-3
lobed, ca 0.3 mm long, 0.4 mm wide, glabrous;style 2.8-3.0 mm long, the stigma
ca 1.0 mm in diameter;ovary subglobose, 1.0-2.0 mm in diameter,villous, the
ovulesnumerous.Fruit7.0-9.0 cm long, 1.4-2.0 cm per side; exocarp fleshy, thin (ca
0.5 mm), with a whitish golden-brownvelutinous-tomentosepubescence;endocarp
cartilaginous,glabrous.Seeds ca 20 per locule, ovoid to globose, 5.0-6.0 mm long,
ca 4.0 mm wide, equinatepubescent,the trichomesca 2.0 mm long.
Type. Spruce 1501, Brazil, Amazonas,Manaus,fl (holotype C; isotypes
CGE,F, G, GH, GOET,M, NY, P, W).
Distribution.A common but poorly collected riverinespecies from the
basinof the Rio Negro. It growsin periodicallyflooded riversideforests, especially
alongblack-waterrivers,althoughit has once been collected along the Rio Madeira,
and is known to have been in cultivationin a gardenin Rio de Janeiro. Known
from Venezuelaand Brazil,probablyalso found in Colombia.
VENEZUELA. Territorio Amazonas: Gaillard 164 fl (P); Bunting, Akkermans & Rooden
4067 fl (U); Level 101 'fl (NY, VEN); Wurdack& Adderley 43194 fl (NY, VEN); Wurdack&
Adderley 43283 fl (NY, VEN); Wurdack& Adderley 43747 fl (NY, VEN). BRAZIL. Amazonas:
Cavalcante 552 fr (MG); Coelho, L. 1139 fl (INPA, IPEAN); Ducke 179 fl (A, F, MO, NY, US);
Fr6es21293a fl (F, IPEAN,NY, US);Froes22588 fr (IPEAN,U); Fr6es25056 bds (IPEAN);

Labroysn fl (P);Pranceetal 2657 fl (COL,F, INPA,MG,NY, US);Pranceetal 4945 fl (COL,
INPA,NY);Pranceet al 15200 fl (COL,MG,NY);Pranceet al 17700 fr (COL,INPA,MG,NY,
US);Rodriguez,W.726 fr (IPEAN);Silva,M. 950 fl, fr (MG,NY);Spruce2416 fl (CGE,F,
FHO,G, GH,GOET,P, W);Ule 6074 fl (G, L, MG);Ule 6140 fl (GOET,MG,W).
8. TrigoniareticulataLleras,sp nov
Frutex scandens,ramulisjuvenilibustomentellis,glabrescentibus,lenticellatis.
Stipulascaducae,haud visae. Folia opposita;petiolus 9.0-18.0 mm longus, 1.0-3.0
mm crassus,tomentellus;laminaeovataevel ellipticae,interdumobovatae,4.5-11.0
cm longae, 3.0-6.5 cm latae, subcoriaceae,margineleviter revolutae,apice acuto
usque acuminatovariante,basi obliquavel subrotundata,supraglabrae,infralanato-
tomentellae;costa media supraplana,infra prominens,strigulosa;costis secundariis
(8-)9-11(-12)-jugis,costis tertiariisquaternariisquereticulatis,evidentibusinfra.
Inflorescentiaein thyrsisterminalibusvel axillaribusdispositae,4.0-15.0 cm longae.
Flores in dichasiassimpliciausque composita dispositi; axes dichasiorum0.5-2.0
mm longi, 0.5-0.8 mm crassi,tomentelli;pedunculi0.3-1.0 mm longi, 0.5-0.8 mm
crassi,tomentelli, bracteissubulatis, 1.0-2.5 mm longis, margineglanduloso-papil-
latis, tomentellis;pedicelli 1.0-2.5 mm longi, 0.3-0.5 mm crassi,tomentelli, bracte-
olis subulatis, 1.0-1.5 mm longis, margineut in bracteis,tomentellis;sepalaovata
vel oblonga, 2.8-4.0 mm longa, 1.7-2.2 mm lata, nonnumquamglandulosa,
tomentella;vexillum 4.3-4.6 mm longum, 2.5-2.8 mm latus, usque ad medium
longitudinalitersaccatum,apice leviter revolutum,intus barbatum;alae spathulatae,
3.3-3.8 mm longae, 1.3-1.6 mm latae, ad basin barbatae;carinaepetala saccata,
2.8-3.0 mm longa, 1.0-1.8 mm lata; stamina8-10, sterilia24, fertilia6, staminodiis
interdumappendiculatis,filamentisad mediumconnatis, 1.0-1.4 mmlongis, antheris
oblongisvel ellipticis, 0.5-0.7 mm longis, 0.3-0.4 mm latis; glandulae2, unilobae, a
fronte visae labiatae, lanato-villosae;stylus erectus, 1.0-1.2 mm longus, villosus,
stigmatecirculari,ca 0.3 mm diametro,albo; ovariumsubglobosumusque pyrami-
dale varians,ca 0.6 mm latum, 3-loculare,villosum, ovulis in quoque loculo
numerosis.Fructusjuvenilisoblongus,dense villoso-tomentosus.
Type. Tillett & Tillett 45524, Guyana,UpperMazaruniRiverBasin,Kako
River,fl (holotype NY; isotypes F, GH, P, US).
Distribution.At edge of forest in open areasalong roadsor rivers.
VENEZUELA.Bolivar:Steyermark&Aristeguieta77 fl, imm fr (VEN,U). GUYANA.
Forest Department7992, KukuRiver,Fl (NY).
Due to lack of adequate fruiting material, the relationships of this species are
impossible to determine. It has been confused with Trigonia villosa var macrocarpa
(T. macrocarpaBentham)and somewhat resemblesit but differsin the reticulation
of the leaves, the more complex inflorescence,and severalother minor characters.
Trigoniareticulata and T. bracteataare unique in Trigoniain havingglandular
papillaeon the marginsof the bracts,bracteolesand sometimesthe sepals.
9. Trigonia candelabra Lleras, sp nov Fig 14, 15.
Frutex scandens, ramulis juvenilibus tomentellis, glabrescentibus, lenticellatis.

Stipulae caducae, non visae. Folia opposita, petiolo 8.0-15.0 mm longo, 1.0-3.0 mm
crasso, tomentello, postea glabro;laminae ellipticae vel obovatae, 8.0-21.0 cm
42 Flora Neotropica
?? r
FIG 14. Trigonia candelabra A, Habit, vegetative branch; B, Habit, flowering branch; C, Habit,
fruiting branch, X 0. 3?3.
P*~~~ '
D E F
FIG 15. Trigonia candelabra A, flower X3.3; B, ovary X3.3; C, anthers X6.6; D, Keel petal X
3.3; E, Wing Petal X3.3; F, Standard X3.3; G, compound dichasium in bud X3.3.
longae,4.0-10.0 cm latae, subcoriaceae,margineintegrae,apice acuto usque acumi-

nato variante,basi obtusa vel obliqua, supra glabrae,infra minute tomentellae
postea glabrae;costa media supraplana,infra prominens,strigulosa,costis secund-
ariis 6-9-jugis.Inflorescentiaein thyrsis terminalibusvel axillaribusdispostiae,
5.0-9.0 cm longae. Flores in dichasiacomposita dispositi;axes dichasiorum1.0-7.0
mm longi, 0.4-0.9 mm crassi,tomentelli; pedunculi0.8-2.0 mm longi, 0.5-1.0 mm
crassi, tomentelli, bracteis subulatis, 1.5-2.5 mm longis, tomentellis; pedicelli
1.0-2.5 mm longi, 0.3-0.5 mm crassi, tomentelli, bracteolis subulatisvel ovatis,
0.8-1.5 mm longis, tomentellis;sepalaovata vel oblonga,4.34.6 mm longa, 1.0-2.0
mm lata, tomentella;vexillum 4.5-5.0 mm longum, 2.0-3.0 mm latum, usque ad
medium longitudinalitersaccatum, apice leviter revolutum,intus barbatum;alae
spathulatae,4.04.5 mm longae, 1.3-1.6 mm latae, glabrae,carinaepetala saccata,
3.0-3.5 mm longa, 1.0-1.3 mm lata; stamina8-10, sterilia24, fertilia6, filamentis
ad mediumconnatis, 1.5-1.8 mm longis, antherisoblongis vel ovatis, 0.8 mm longis,
0.6-0.9 mm latis, apice acuminatis;glandulae2,2-3 lobae, rectangulatae,ca 0.5 mm
latae, laciniatae;stylus erectus, 1.3-1.6 mm longus, villosususque glabervarians;
stigmatecirculari,ca 0.3 mm diametro,albo; ovariumsubglobosum,ca 1.0 mm dia-
metro, 3-loculare,villosum, ovulis in quoque loculo numerosis.Fructus6.5-8.0 cm
longus, ca 1.0 cm latus, oblongo-obovatus,extus velutino-tomentosus,intus carti-
lagineus,glaber.Seminaca 10 in quoque loculo, subglobosa,ca 4.0 mm diametro,
pilis ca 7.0 mm longis.
Type. VanDonselaar2812, Suriname,Brokopondo,villageof Brokopondo,
at edge of secondaryforest, fl (holotype U).
Distribution.Known only from the type of locality, and from three collec-
tions in Brazil.
SURINAME. Brokopondo: Van Donselaar 2891, fr (paratype U). BRAZIL. Para: Froes
32897 fl (IPEAN); Pires & Silva 4734 fl (IPEAN).
44 Flora Neotropica
This very distinct species shows affinities with Trigoniahypoleuca, in the leaf
(shape, pubescence, color, etc) and the fruit. However,the inflorescenceis very
distinct. It is the only speciesin the genusin which the flowers are arrangedin
perfectcompounddichasia.The sterile (vegetative)branchesbear some of the largest
leaves in the genus, another characterthat separatesit from T. hypoleuca. The
pubescencein the youngerleaves cannot be seen with the naked eye, or even with a
dissectingmicroscope(100X).
I have selected floweringmaterialas the holotype for the species. It is quite
possible that the fruitingmaterialdesignatedas paratypeis from the same plant,
gatheredat a later date.
The specific epithet, candelabra,refersto the arrangementof the ultimate
inflorescences.
10. TrigoniarugosaBentham,Bot. Voy. Sulph. 74. 1844.
TrigoniafloribundaOrsted,DanskNaturh.Foren., Copenh.Vide. Meddel.38. 1856;

Standley,N. Am. Fl. 25: 297. 1924;Austin, Fl. of Panama,Ann. MissouriBot.
Gard.54(3): 208. 1967. Type. Orsted3721 Nicaragua,Granada,fl (holotypeC;
isotypes F).
TrigoniarigidaOrsted,DanskNaturh.Foren.,Copenh.Vide. Meddel.38. 1856. Type.
Orsted3 722 Nicaragua,Granada,fl (holotypeC).
Trigonia thyrsifera Donnell Smith, Bot. Gaz. 23: 3. 1897. Type. Biolley 2638 Costa
Rica, Alajuela,San Mateo,fl (holotype US;isotype CR).
Trigonia euryphylla Standley, N. Am. Fl. 25: 297. 1924. Type. Baker 2554 Nicaragua,
Granada,fr (holotype US;isotypesA, G, GH, K, MO).
TrigoniapanamensisStandley,Publ.Field Mus.Nat. Hist. Bot. 22: 346. 1940. Type.
Aviles 961 Panama,CanalZone, BarroColorado,fl (holotype F).
Scandentor subscandentshrub,the branchesterete, lenticellate, the young

ones densely tomentose, becoming glabrouswith age. Stipules caducous (nv);
petioles 5.0-25.0 mm long, 0.8-1.2 mm thick, subcanaliculate,tomentose or
glabrous;laminaoblong-ellipticto obovate, (3.5-)5.0-15.0 cm long, (2.5-)4.0-8.0 cm
wide, chartaceousor subcoriaceous,the marginsentire, the apex roundedor acute,
the base obtuse to acute; venation eucamptodromous,tomentose to strigose on
both surfaces,the midribprominulousabove, prominentbeneath, the secondary
veins 6-9 pairs;intercostalpubescenceabsent above, lanate-adpressedto arachnoid
when young, frequentlybecomingglabrouswith age. Inflorescencesterminaland
subterminalthyrses, 7.0-25.0 cm long. Flowers in dichasiaof 1-3, arrangedulti-
mately in secondaryinflorescencesvaryingfrom dichotomousdichasiato individual
cymules; axes of secondaryinflorescences4.0-7.0 mm long, ca 0.8 mm thick,
tomentose, the bractsovate, 2.0-3.5 mm long, tomentose;peduncles0.7-2.0 mm
long, ca 0.4 mm thick, tomentose; pedicels 1.0-2.0 mm long, ca 0.4 mm thick,
tomentose, the peduncularbractsand bracteolesof equal size, 0.8-1.5 mm long,
tomentose; sepals ovate or oblong, 2.2-3.5 mm long, 1.2-1.8 mm wide, lanate;
standard3.2-4.0 mm long, 1.2-2.2 mm wide, the pouch globose, extending to ? of
the length, the apex revolute,barbateat the throat, the wingsbroadlyspathulate,
2.8-3.2 mm long, 1.0-1.5 mm wide, barbateat the base, the keel petals 2.7-2.9 mm
long, 0.6-1.5 mm wide, sometimeslackinga pouch, glabrous;stamens8-10, with 6
fertile, staminodes2-4, sometimeswith knob-likeappendages,the filaments 1.0-1.2
mm long, connate to the middle, the anthersoblong, 0.4-0.5 mm long, ca 0.2 mm
wide; glands2, 2-3 lobed, irregular,ca 0.5 mm per side, villous or glabrous;style
1.2-1.3 mm long, slightly villous or glabrous,the stigmatrilobate,ca 0.2 mm in

diameter;ovary subglobose, ca 0.6 mm in diameter,short-villous,the ovules
numerous. Fruit oblong to elliptic, 2.0-3.2 cm long, 1.0-2.0 cm wide; exocarp
yellow velutinous-tomentosewhen young, glabrouswhen mature. Seeds 6-9 per
locule, ovate, ca 4.0 mm long, villous-barbate,the trichomesca 10.0 mm long.
Type. Sinclairsn, CentralAmerica,fl, fr (holotype, K).
Distribution. A well collected species known from Southern Mexico to
NorthernColombia,from the tropicaland subtropicaldeciduousforests along the
Pacific coast of CentralAmerica,and the Atlantic coast of northernSouth America.
MEXICO. Chiapas: Matuda 2517 fl (F, G, K, MICH, NY); Matuda 16548 fl (F, MICH,
NY). GUATEMALA. Retalheu: Standley 87755 st (F); Standley 88330 st (F); Standley 88718
st (F). Suchitepequez; Standley 62205 st (F). Escuintla: Molina & Molina 12477 fl (NY).
Santa Rosa: Standley 78860 st (F); Standley 79927 st (F). Zacapa. Standley 73760 f (F);
Standley 73850 st (F). Chiquimula: Standley 32044 st (F); Standley 74348 st (F). Jutiapa:
Standley 95137 st (F); Standley 75475 st (F). EL SALVADOR: Santa Ana: Standley & Padilla
3068 st (F). Salvador: Standley 20596 fr (GH, NY, US); San Vicente: Standley 21340 fr
(GH, NY, US); Standley & Padilla 3411 st (F). San Miguel: Calder6n 2124 fr (GH, NY, US).
HONDURAS: Octepeque: Molina 22436 fr (F, NY). Morazan: Glassman 1 723 fl (F, NY);
Molina 131 fl (F, GH, MO, US); Webster,Miller & Miller 12040 fl (GH, MO, U); Williams&
Molina 11262 fr (F, US). El Paraiso: Molina 7494 fl (F). Choluteca;Molina 14206 fl (F, NY).
NICARAGUA: Managua: Chaves 213 fl (US); GarnierA 1283 fr (A, F). Masaya: Maxon 7675
fl (US); Maxon 7686 fl (US). Granada: Levy 192 fl (C, G, P, US); Levy 1073 fl (G, P); Orsted
3723 fl (C, F). COSTA RICA. Guanacaste: Burger & Burger 7841 fl (F, NY); Jimenez 1575 fr
(F); Quir6z 848 fr (F); Tond0iz 13486a fr (NY, US); Webster,Miller & Miller 12479 fl (GH,
MO, U). Alajuela: Standley 40045 st (US). PANAMA. Canal Zone, Barro Colorado: Croat
16581 fl (NY); Standley 40949 st (US). Darien: Stern et al 163 fl (MO); Stern et al 224 fl (GH.
MO, U, US); Stern et al 993 fl (GH, U, US). COLOMBIA. Antioquia: Haught 4866 fl (COL,
US). Atlantico: Bro. Elias 258 fl (NY, US); Bro. Elias 357 fl (US). Magdalena: H. H. Smith 884
fl (K); H. H. Smith 886 fl (F, G, MICH, P, US); H. H. Smith 888 fl (F, MICH, MO, US). Cesar:
Haught 4186 fl (COL, F, US). Guajira: Haught 4227 fl (COL, NY, U, US).
Trigoniarugosahas been commonly known as T. floribunda,but the former
is the older, valid name. The name T. rugosahas been used only for the material
collected in the type gatheringand althoughthis materialis very poor, it belongs
together with what has been known as T.floribunda. Both the specimensused to
typify T. rugosaand T. euryphyllahave thickerleaves than most of the material
representingthe species, but I considerthis to be a local variationof no taxonomic
significance.
As the high variabilityof the species makesit impossibleto recognizesub-
specific categories,I agreewith Austin in consideringthe other epithets cited under
this species as being synonyms.
11. Trigoniaeriosperma(Lamarck)Fromm& Santos, Bol. Mus.Rio de Janeiro

42: 2. 1971.
Croton eriospermum Lamarck, Encycl. 2: 211. 1786. Type. Commerson sn Brazil, Rio
de Janeiro, fr (holotype P -Herb. Lamarck; isotype P).
Mainea racemosa Vellozo, Fl. Flum. 275. 1825; Vellozo, Iconog. 7: pi. 8. 1827; Netto,
Arch. Mus. Rio de Janeiro 5: 260. 1881.
Trigonia crotonoides Cambessedes, (Hippocrateaceae) St. Hilaire Fl. Bras. Merid. 2:
83. pl. 105. 1829; Grisebach, Linnaea 22: 31. 1849; Warming, (Trigoniaceae)
Mart. Fl. Bras. 13(2): 127. 1875. Type. St. Hilaire c102. Brazil, Rio de Janeiro.
fl, fr (holotype MPU; isotype P).
Trigonia crotonoides var incana Cambessedes, (Hippocrateaceae) St. Hilaire Fl. Bras.
Merid. 2: 83. 1829; Warming, (Trigoniaceae) Mart. Fl. Bras. 13(2); 127. 1875.
Type. St. Hilaire sn, Brazil, Rio de Janeiro, fr (holotype MPU).
46 Flora Neotropica
Trigoniacrotonoidesvar oblongifoliaCambessedes,(Hippocrateaceae)St. HilaireFl.

Bras,Merid.2: 83. 1829;Warming, Mart.Fl. Bras.13(2): 127.
(Trigoniaceae)
1875.Type.St. HilaireBl-385,Brazil,MinasGerais,fl (holotypeMPU;isotypeP).
micrantha
Trigonia Martius, Flora20(2): 102. 1837.Nomennudum.
TrigoniacrotonoidesCambessedesvarellipticaWarming,(Trigoniaceae)Mart.Fl. Bras.
13(2): 128. 1875. Type. Glaziou2114, Brazil,Rio de Janeiro,fl (lectotype C;
isotypes BR,P).
Trigoniaracemosa(Vellozo) Hoehne,Ind. Bibliog.Num. 256. 1951.
Shrubs or scandent shrubs,sometimes treelets, the branchesterete, lenti-

cellate, strigose, becoming glabrouswith age. Stipules linear, 1.54.0 mm long,
strigoseto almost glabrous,caducous;petioles 2.0-5.0 mm long, 0.3-0.5 mm thick,
strigose;laminaoblong-ovateto oblong-elliptic,1.5-9.0 cm long, 1.0-3.5 cm wide,
membranaceousto chartaceous,the marginsentire, the apex acuminate,the base
obtuse;venationeucamptodromousto brochidodromous,strigoseto very densely
strigoseon both surfaces,the midribplane above, prominentbeneath, secondary
veins 4-7 pairs,impressedon both surfaces.Inflorescencesterminalpanicles,thyrses
or racemes,1.5-10.0 cm long. Flowersin cymules of 1-3;peduncles 1.0-6.0 mm
long, ca 0.2 mm thick, tomentellous or strigose,the bractslinear,0.7-4.0 mm long,
strigose;pedicels 1.0-3.0 mm long, ca 0.2 mm thick, tomentellousor strigose,the
bracteolesdeltoid to linear,0.2-1.0 mm long; sepalsovate to oblong, 2.0-3.0 mm
long, 0.8-1.0 mm wide, membranaceous,tomentellous or strigose;standard4.0-5.0
mm long, ca 1.5 mm wide, the pouch extending along ? of the length, spurred,
barbateat the throat, the wings spathulate,2.5-3.5 mm long, 0.8-1.0 mm wide,
barbateat the base, the keel petals 2.8-3.5 mm long, ca 1.0 mm wide, the pouch
centeredon the petal, nasiform;stamens9-10, 6 fertile, 34 staminodes,the fila-
ments 0.9-1.2 mm long, connate for 4/5 of the length, the staminodeswith a small,
knob-like terminalappendage,the antherssubglobose,0.2-0.3 mm in diameter;
glands2, 2-lobed, irregular,ca 0.3 mm per side, villous;style glabrous,0.9-1.2 mm
long, the stigmatrilobate,ca 0.1 mm in diameter;ovarysubglobose,ca 0.6 mm in
diameter,densely villous, the ovules ca 4 per locule. Fruit oblong, 0.8-2.5 cm long,
the valvesca 0.7 cm perside, exocarpvelutinouswhen immature,becomingglabrous
with maturity;endocarpwoody, glabrous.Seedssubglobose,ca 1.0 mm in diameter,
villous, the trichomesca 6 mm long.
This speciesis separatedinto 3 subspecies.
Key to the Subspecies of Trigonia eriosperma
1. Inflorescencesusually longer than 4.0 cm, in paniclesor thyrses;cymes usually

2-3-flowered.
2. Inflorescencespanicles; cymules usually 3-flowered (sometimes 1); leaves
villousor lanate;from S.Brazil. a. subsperiosperma.
2. Inflorescencesthyrses;cymules 2-floWered;leavesglabrousor nearlyso; from
Central& N South America. c. subspmembranacea.
1. Inflorescencesshorterthan 3.5 cm, racemes;cymules 1-flowered;from S. Brazil.
b. subspsimplex.
1la. Trigoniaeriospermasubsperiosperma
Leaves1.5-7.0 cm long, 1.0-3.0 cm wide, villous-strigoseto lanate. Inflores-

cences terminalpanicles,4.0-8.0 cm long. Flowersin 3-floratecymes (sometimes
1-florate).Fruit 0.8-1.5 cm long.

Type. Commersonsn Brazil,Rio de Janeiro,fr (holotype P-LAM;isotype P).
Distribution.Known from forests in southernBrazil.
BRAZIL.Bahia:Belem 1185 fr (NY);Blanchet3593 fl (F, G, MO,P). MinasGerais:
Claussen1096 fl (G, P); Glaziou13480 fl (MPU,P); Glaziou13811 fl (B, C, F, G, IPEAN,MG,
P, US);Glaziou14689fl (BR,C,G,IPEAN,MPU,US);Mexia
4120 fl (F, G, NY,US).Espirito
Santo: Duarte4011 fl (NY);Pereira9835 fl (F, M). Rio de Janeiro;Brade11215 fl (GH);
Glaziou9417 fl (BR, E, P); Langsdorffsnfl, fr (P, US);Martius122 fl (BR, F, G, GH, MO,
NY, P, US);Pereira& Duarte1246 fl (NY); Vauthier522 fl (G, GH); Weddell684 fl (G).
Guanabara; Lindley 749 fr (BR);Luschnathsn fl, fr (BR, GH);Schenk1589 fr (C);Schenk
3061 fr (C).
This subspeciesis one of the most collected taxa in the genus. Becauseof the
high variabilityof characters,it has been treatedas severalspecies, or at best, as
severalvarieties.I agreewith Fromm& Santos (1971) in consideringit as only one
taxon; it is impossible to separatemorphologicallythe taxa delimited by other
authors.
The affinities of the species as a whole indicate close relationshipswith both
Trigonialaevis and T. paniculata.
Trigoniaeriospermasubspecieseriospermacanbe separatedfrom T. paniculata
by its shorterpanicles,the paniclesbeing terminal(vs terminaland axillaryin T.
paniculata),by its more glabrousstems and floweringbranches,by the lack of a
white "border"on the marginsof the leaves, and by the sparserpubescenceof the
leavesin general.
i lb. Trigoniaeriospermasubsp simplex (Warming)Lleras,stat nov
TrigoniasimplexWarming,(Trigoniaceae)Mart.Fl. Bras.13(2): 125. 1875.
Leaves 3.5-5.0 cm long, 1.5-2.5 cm wide, strigose. Inflorescencesterminal

racemes,1.5-3.5 cm long. Flowersin 1-floratecymules. Fruit (immature)ca 2.5
cm long.
Type. Warming596, Brazil,MinasGerais,LagoaSanta, fl, fr (holotype C, on
2 sheets as indicatedby Warming).
Distribution.Known only from the type locality.
BRAZIL.MinasGerais:Warming
sn fl, fr (topotypescollectedat differenttimes, C, US).
1lc. Trigoniaeriospermasubspmembranacea(A. C. Smith) Lleras,stat nov
membranacea
Trigonia A. C. Smith,Phytologia
3: 128. 1935.
rasaStandley&Steyermark,
Trigonia Publ.FieldMus.Nat.Hist.23(2):59. 1944.Type.
Standley78584Guatemala, SantaRosa,RioPanal,fr (holotypeF;isotypeF).
Leaves4.0-9.0 cm long, 2.5-3.5 cm wide, very slightly strigoseto glabrous.

Inflorescencesterminalthyrses3.0-10.0 cm long. Flowersin 2-floratecymes. Fruit
oblong, 1.5-2.0 cm long, glabrous.
Type. Killip & Smith 14396, Colombia, Bolivar,Turbaco, fr (holotype
NY; isotypes A, GH, US).
Distribution. Reported from fairly open areas or edges of forest from the
Atlantic Coast to CentralAmerica,and NorthernSouth America,from the South
of Mexico to NorthernColombia.
48 Flora Neotropica
MEXICO.Chiapas:Matuda17630 fl (F, NY). GUATEMALA. Bernoulli& Cario3139

fl (C, K);Standley87416 st (F); SnedakerC-24 fl (F); SnedakerC-25fl (F). BELIZE.Gentle
5251 fr (MICH,U); Lundell 7021 fr (MICH,U). PANAMA.Hayes 720 fl, fr (BR, E, K,
M). COLOMBIA. Atlantico:Bro. Elias1202 fr (US);Bro. Elias1651 fr (F, US);Bro. Elias 1105
fr (F, US).
Both subspeciesmembranaceaand subspeciessimplex have largerfruits than
subspecieseriosperma.Subspeciesmembranaceais in generalthe most glabrousof
the three subspeciesin the species, and subspeciessimplex is probablymore xero-
phytic than the other two.
12. Trigoniafloccosa Rusby, Bull. N. Y. Bot. Gard.4: 325. 1907.
Scandentshrub,the branchesterete, lenticellate, strigoseor adpressed-lanate,

becomingglabrouswith age. Stipulessubulateto lanceolate, 10.0-12.0 mm long,
tomentellous, caducous;petioles 5.0-15.0 mm long, ca 1.5 mm wide, strigulose;
lamina ovate to obovate, inequilateral,4.0-13.0 cm long, 2.0-6.0 cm wide, sub-
coriaceous,the marginsentire, the apex acute to acuminate,the base subcordate;
venation eucamptodromous,lanate to glabrousalong the midrib above, strigose
beneath, the midrib plane above, prominent beneath, secondaryveins impressed
above, prominulousbeneath;intercostal pubescence arachnoid-lanateto glabrous
above, adpressedwhite-lanateand golden-strigosebeneath. Inflorescencesterminal
and subterminalaxillarythyrses,to 12 cm long. Flowersin cymules of 1-2, ped-
unclesabsentor nearlyso;pedicels 1.0-1.5 mm long, ca 0.4 mm thick, tomentellous,
bracts and bracteoles of equal size, 2.5-10.0 mm long, subulate, tomentellous;
sepals ovate to oblong, sometimes deltoid, 3.5-5.2 mm long, 2.0-2.5 mm wide,
papillateat the margins,lanate or strigose;standard5.0-5.5 mm long, 2.5-3.0 mm
wide, the pouch extending along 1/3 of the length, slightly revoluteat the apex,
barbateat the throat, the wings spathulate, 5.0-5.5 mm long, ca 1.5 mm wide,
densely barbateat the base, the keel petals 4.04.5 mm long, ca 3.0 mm wide, the
pouch nasiform;stamens8-9, fertile ones 6-7, staminodes2, the filaments 1.8-2.0
mm long, connate for /2 of the length, anthersoblong, 0.8-1.0 mm long, ca 0.4 mm
wide; glands2, 2-3 lobed, 0.5-0.8 mm per side, acute at the apex, fused to the
staminaltube; style 1.8-2.0 mm long, villous, the stigmatrilobate,ca 0.3 mm in
diameter;ovarysubglobose,ca 0.6 mm in diameter.villous, the ovulesnumerous.
Fruit(immature)5.0-6.0 cm long, ca 1.2 mm perside; exocarpvelutinous-tomentose;
endocarpwoody, glabrous.Seeds8-10 perlocule, subglobose,ca 3.0 mm in diameter,
barbatepubescent,the trichomesca 12 mm long.
Type. Bang2191, Bolivia,Yungas,Coripati,fl (holotype NY; isotypes E, G,
GH, M, MICH,MO,NY, US, W).
Distribution.Known only from two localities in Bolivia.
BOLIVIA. Santa Cruz: Rusby 2450 fl (GH, MICH, NY, US); Rusby 2596 fr (MICH, NY).
This species can easily be recognizedby the dual pubescencetypes on the
undersideof the leaf, by the largeglandsfused to the staminaltube, and especially
by the papillaeon the sepals.
13. TrigoniacostanensisSteyermark& Badillo,Acta Bot. Ven. 6(14): 77. 1971.
Scandent shrub, the branches terete, lenticellate, slightly yellow-puberulent-

tomentose when young, becoming glabrous with age. Stipules caducous (nv);
petioles 15.0-20.0 mm long, ca 1.0 mm thick, puberulent-tomentose,sometimes
glabrous;lamina oblong-ellipticto oblong, 8.0-15.0 cm long, 4.0-8.0 cm wide,

coriaceous,the marginsentire, the apex obtuse or rounded,sometimesmucronate,
the base obtuse;venationeucamptodromous,glabrousabove,slightlyvillous beneath,
the midribe plane above, prominulousbeneath, the secondarynerves 6-8 pairs;
intercostalpubescenceabsent above, adpressed-lanate(sericeous)beneath. Inflores-
cences axillary thyrses 10.0-15.0 cm long. Flowers in groups of 1-3; peduncles
2.04.0 mm long, ca 0.4 mm thick, densely yellow-puberulenttomentose; the
bracts3.5-5.0 mm long, linear to subulate,puberulent-tomentose;pedicels 1.0-2.0
mm long, ca 0.4 mm thick, puberulent-tomentose,the bracteoles2.0-3,5 mm long,
1.0-2.0 mm wide, linearto ovate, puberulent-tomentose;sepals ovate or oblong,
4.5-6.0 mm long, 2.0-3.0 mm wide, puberulent-tomentoseon exposed portions,
adpressed-lanateon coveredportions;standard4.5-5.0 mm long, ca 3.0 mm wide,
the pouch extending to 2/3 of the length, the apex rounded,barbateat the throat,
the wings spathulate,4.5-5.0 mm long, 1.5-1.8 mm wide, barbateat base, the keel
petals 4.0-4.5 mm long, 2.0-2.5 mm wide, ovate-oblong,not saccate,ciliate (fide
Steyermark& Badillo)at the apex; stamens8-9, with 6 fertile, staminodes2-3, the
filaments 1.8-2.0 mm long, connate for most of their length, the anthersoblong,
0.8-1.0 mm long, ca 0.4 mm wide; glands2, 2-3-lobed, deltoidor trapezoid,ca 0.3
mm long per side, glabrous;style 1.8-2.0 mm long, glabrous,the stigmatrilobate,
ca 0.1 mm in diameter;ovarysubglobose,ca 1.5 mm in diameter,densely villous,
the ovules numerous.Fruit unknown.
Type. Madriz37, Venezuela,Yaracuy,north of Nirgua,fl (holotype NY).
Distribution.Confinedto the cloud forests of the coastal Cordilleraof the
states of Yaracuyand Carabobo.
VENEZUELA.Yaracuy:Steyermark,Bunting& Wessels-Boer 100290 fl (paratypesNY,
VEN): Carabobo:Steyermark& Steyermark95234 fl (paratypesNY, VEN).
This species is closely related to Trigoniasericea, from which it differsin
havingshorterand less ramifiedinflorescences,slightlylongerpetioles, and specially
in the much longer bractsand bracteoles,as well as severalother minor characters.
14. TrigoniasubcymosaBentham,Hook. Lond.Jour. Bot. 2: 373.1843; Grisebach,

Linnaea22: 31. 1849; Warming,(Trigoniaceae)Mart.Fl. Bras. 13(2):
127. 1875;Stafleu, in Pulle Fl. Surin.3(2): 176. 1951.
Largeshrub,the branchesterete, denselylenticellate, strigosewhen young,

becomingglabrouswith age. Stipulessubulate,4.0-5.0 mm long, ca 2 mm wide,
caducous;petioles 3.0-4.0 mm long, ca 0.5 mm thick, densely golden-brownstri-
gose; lamina elliptic to obovate, or oblong, 3.0-8.0 cm long, 2.04.0 cm wide,
chartaceous,the marginsentire, the apex obtuse, acute or mucronate, the base
cuneate to obtuse;venationeucamptodromous,sparselygolden-strigosepubescent,
the midrib plane above, prominulousbeneath, the secondarynerves 8-10 pairs;
intercostalpubescencevery sparselystrigoseabove, depressed-lanatebeneath, the
indumentumcreamor yellowish in most cases. Inflorescencesterminalpyramidal
paniclesto 15 cm long. Flowersin cymules of 2-4; peduncles 1.0-8.0 mm long,
diminishingin length towardsthe apex of the inflorescence;pedicels 1.0-2.0 mm
long, villous-strigose,the bracts and bracteoles of equal size, 2.0-3.0 mm long,
0.5-0.7 mm wide, ovate, curvedupwards,villous strigose;sepals ovate or oblong,
2.5-3.5 mm long, 0.7-1.5 mm wide, strigose or villous-strigosealong exposed
portions, lanate along protected portions; standard3.0-3.5 cm long, ca 2.0 mm
50 Flora Neotropica
wide, the pouch extending to 2 of the length, erect along the upperportion or
nearlyso, the apex revolute,barbateat the throat, the wings spathulate,2.5-3.1
mm long, 0.8-1.0 mm wide, glabrousat base, the keel petals 2.5-2.8 mm long, ca
2.0 mm wide, with the pouch extendingfrom 1/3 of the length up, the apex revolute,
glabrous;stamens 8, fertile ones 5-6, staminodes2-3, the filaments 1.0-1.5 mm
long, free for 1/3 of the length, the anthersovate or oblong, 0.4-0.5 mm long, ca
0.2 wide; glands2, 2-3 lobed; deltoid or trapezoid,ca 0.4 mm per side, glabrous;
style 0.8-1.1 mm long, glabrousor villous, the stigmatrilobateca 0.1 mm in diam-
eter;ovarysubglobose,ca 0.4 mm in diameter,villous-barbate,the ovulesnumerous.
Fruit not known.
Type. Schomburgk56 Guyana, fl (lectotype K - Bentham Herb.;isotypes
CGE,G, NY, W).
Distribution.Known only from Schomburgkcollections made in Guyana.No
localities are reported.
GUYANA. Schomburgk 63 fl (paratypes BR, CGE, G, OXF, U, W); Schomburgk 249 fl
(F, W); Schomburgk 373 fl (GOET, K).
Trigoniasubcymosais easily recognizedby its short, pyramidalpanicles,the
long peduncles, the typically curved bracts and bracteoles, and its fairly small
leaves and flowers.
Benthamcited both Schomburgk56 and 63 in his descriptionof this species.
These specimensarevery similarand either one could be selected to typify the
taxon. I have selected Schomburgk56 as the lectotype. Thereare other elements
labeled as 63 and 56 that are not referableto this speciesbut Trigoniavillosa,and
this mixturehas led to confusion in the past. The specimenfrom Bentham'sherb-
ariumat Kew has Schomburgkcollections 56, 63, and 373 mounted on the same
sheet. This could well mean that they constitute elements of the same gathering
that were numbereddifferently.The specimensare extremely similar.
The lack of fruit makesit extremely difficult to determinethe relationships
of this species.
15. Trigonianivea Cambessedes,(Hippocrateaceae)St. Hil. Fl. Bras.Mer.2: 81.

1829; Grisebach,Linnaea 22: 29. 1849; Warming,(Trigoniaceae)
Mart. Fl. Bras. 13(2): 134. 1875; Reitz, in Fl. Ill. Catar. 1(13): 10.
1967.
Trigonia candida Warming,(Trigoniaceae) Mart. Fl. Bras. 13(2): 139. 1875. Type.Glaziou
2505, Brazil, Rio de Janeiro, fl, fr (holotype C; isotypes A, BR, C, F, GH, IPEAN,
NY, US).
Trigonia ovalifolia Glaziou, Bull. Soc. Bot. Fr. 3(52): 34. 1905. Type. Glaziou 14690,
Brazil, Minas Gerais, fl (holotype P; isotypes C, F, G, P).
Trigonia nivea forma paniculata Chodat & Hassler, Bull. Herb. Boiss. 2(8): 801. 1903.
Type. Hassler 8416, Paraguay, Apa River Headwaters, fl, fr (holotype G; isotypes
BM, G, MO, NY).
Scandentshrub,the branchesterete,lenticellate,lanatewhenyoung, becoming

glabrouswith age. Stipules triangular,6.0-10.0 mm long, strigulose,caducous;
petioles 2.0-8.0 mm long, 0.5-1.0 mm wide, strigoseor lanate, sometimesbecoming
glabrous;laminaelliptic to oblong-elliptic,sometimesovate or obovate, 5.0-13.0
cm long, 1.0-4.5 cm wide, subcoriaceous,the marginsentire or revolute,the apex
acute to acuminate,sometimes mucronate,the base obtuse; venation eucampto-
dromous,slightly striguloseor glabrousabove, lanate or strigosebeneath, the mid-

rib plane above, prominentbeneath, the secondarynerves 10-16 pairs;intercostal
pubescence absent above, lanate to adpressedlanate (almost sericeous)beneath.
Inflorescencesterminaland axillaryracemes,or panicles5.0-10.0 cm long, varying
from highly congested to very open (sometimes on the same plant). Flowers in
groups(cincinni) of 1-4; peduncles0.0-5.0 mm long, ca 0.5 mm thick, strigose,the
bracts2.0-6.0 mm long, subulateor triangular,strigose;pedicels 1.0-5.0 mm long,
ca 0.5 mm thick, strigose, the bracteoles 1.04.0 mm long, subulate, strigose;
sepals ovate or oblong, 3.0-5.0 mm long, 1.5-3.0 mm wide, lanate on protected
portions, strigoseon exposed areas;4.5-6.5 mm long, ca 3.0 mm wide, the pouch
extending along ? of the length, barbateat the throat, the keel petals 3.5-5.0 mm
long, ca 3.0 mm wide, the pouch extending along 2/3 of the length;stamens 10-11,
fertile ones 6-7, staminodes3-4, the filaments 1.0-2.0 mm long, free for 1/ the
length, the anthersoblong, 0.4-0.7 mm long, ca 0.3 mm wide; glands2-3-lobed,
trapezoidor rounded,ca 0.4 mm per side, in some cases with short laciniaeon the
lobes, glabrous;style 1.0-2.0 mm long, glabrous,the stigmatrilobate, 0.2-0.3 mm
in diameter;ovarysubglobose, 1.0-1.5 mm in diameter,densely villous, the ovules
numerous.Fruits3.0-7.0 cm long, 1.0-1.5 cm per side; exocarp thin, velutinous-
tomentose or slightlystrigose;the endocarpseparablefrom the mesocarp,velutinous-
tomentose or partiallyso, sometimesglabrous.Seeds ovate, barbatepubescent,the
trichomesca 10.0 mm long.
This speciesis separatedinto 3 varieties.
Key to the Varietiesof Trigonianivea
1. Petioles shorterthan 3 mm;glandslaciniate. 15 b. varfasciculata.

1. Petioles4-8 mm long;glandsnot laciniate.
2. Peduncles,when present, to 1.5 mm long; leaves usually oblong to oblong-
ellipticwith dense,white pubescence;bractsusuallyas long as the pedicels.
15 a. varnivea.
2. Peduncles2-5 mm long; leaves usuallyelliptic or ovate-elliptic;bractsshorter
than the pedicels. 15 c. varpubescens.
15a. Trigonianiveavar nivea
Leavesoblong to oblong-elliptic,denselylanate to adpressed-lanatebeneath,

the marginsentire to revolute;petioles 4.0-8.0 mm long. Flowerbuds ca 5 mm
long; peduncles0.0-1.5 mm long, the bracts4.0-5.0 mm long. Fruit 5.0-7.0 mm
long; endocarpusuallypubescent.
Type. St Hilaire226 Brazil,fl, fr (holotype MPU).
Distribution.Extendingin a coastal and mediterraneanbelt from Venezuela
to Santa Catarinain southernBraziland adjacenteasternParaguay;the habitatsmost
frequentlyoccupiedby this varietyseem to be coastal forests and mesophilicgrass-
lands and forests, with some populationsfound in drier,more xerophytic areas.
VENEZUELA. Sucre: Aristeguieta 3988 fl (US, VEN). Anzoategui: Aristeguieta &
Agostini 5554 fl (VEN); F. D. Smith 32 fl (US). Monagas: Pursell et al 8911 fl (NY, US).
GUYANA.Rupununi:Irwin 679 fl (US). Quelch& MCDonnell222 fl (K). BRAZIL.Para:
Black 49-7957 fl (IPEAN); Froes & Black 24606 fl (IPEAN). Maranhao: Pires & Black 2481 fl
(IPEAN). Piaui: Ducke HG 814 fr (MG); Lisboa 2388 fr (MG; Ceara: Frd Allemdo 602 fl (P).
52 Flora Neotropica
Ducke 2453 fl, fr (IPEAN);Guedes483 fl (IPEAN,MG);Lisboa2433 fl (MG).Pernambuco:

Gardner943 fl (W);Schenk4250 fl (C);Pickel133 fl (F, IPEAN,US). Bahia:Belem 1170 fl
(IPEAN,NY);Belem3632 fl (IPEAN);Belem3652 fl, fr (IPEAN);Blanchet2029 buds (F,
G, M, NY); Guillotsn fl (U). MinasGerais:Barreto 7154 fl (A, F); Pires & Black 3314 fl
(IPEAN),NY);Sampaio7163 fl, fr (F). Rio de Janeiro:Glaziou10730 fl (C, G, P); Glaziou
Aparicio& Rizzini59 fl (NY); Trinta511 &
8670 fl, fr (G, P);Pereira4250 fl (F). Guanabara:
Fromm1587 fl (M). Sao Paulo:Humboldt& Bonpland1870 fl (NY);Hoehnesn, Sao Paulo
BotanicalGarden,fl (NY); Novdes1108 fl (US). Parana:Dusen8696 fl (L);Dusen12168 fl
(GH);Hatschbach6861 fl (L, W);Hatschbach19042 & Guimaraesl31fl (C). PARAGUAY.
Amambay:Fiebrig5133 fl, fr (A, G, GH,GOET,L, M, US, W);Hassler5588 fl (G, NY). Alto
Parana:Hassler5362 fl, fr (BM,G).
Commonname: Cipo de Paina(Santa Catarina,Sao Paulo).
This highly variablevariety includes the taxa formerly known as Trigonia
candida and T. ovalifolia, as well as T. nivea f paniculata.Warmingproposed
T. candidabasedprimarilyon leaf characters,and a more open inflorescence.An
examinationand a comparisonof the types of T. candidaand T. niveayield infor-
mation which makesit impossibleto maintainthem as separatetaxa. Furthermore,
had Warmingbeen able to examinemore sheets of the same materialthat he used as
a basis for T. candida,he would have found his specific charactersto be variable
even within the same specimen.The sheets examinedby Warminghave fairly open
inflorescences,whilematerialof the same collections depositedin other herbariawas
almost identical to the St. Hilairecollection of T. nivea.
This high variabilityis presentthroughoutthe whole species,but is especially
noticeablein this variety. AlthoughI have established3 varietieswithin the species,
this solution is not perfect. Thereis some overlapin charactersbetween the three.
On the other hand, I have retainedmaterialin varnivea that might possibly be
distinct, if there were consistencyin the characters.
15b. Trigonianiveavar fasciculata(Grisebach)Lleras,stat nov
TrigoniafasciculataGrisebach,Linnaea22: 29. 1849; Warming,(Trigoniaceae)Mart.

Fl. Bras.13(2): 139. 1875.
Leavesoblong to oblong-elliptic,sometimeselliptic, denselylanate beneath,

the marginsentire; petioles 1.5-3.0 mm long. Flower buds ca 5.0 mm long, the
bracts3.0-6.0 mm long. Glandslaciniate.Fruit ca 4.0 cm long; endocarpwith very
short velutinous-tomentosepubescence.
Type. Blanchet 2921, Brazil, Bahia, Rio Sao Francisco,fl (lectotype G,
isotypes BR, E, F, G, MG,P, W).
Distribution.Reportedonly from Bahiaand MinasGerais.
BRAZIL.Bahia:Dialer618 fl (M);Rose & Russell19950 fl, fr (NY, US);Rose & Russell
19968 fl, fr (GH,NY). MinasGerais:G.MendesMagalhdes5343 bds (IPEAN).
This variety,correspondingto what has previouslybeen known as Trigonia
fasciculata,can easily be distinguishedby its short petioles, the lanate pubescence
on the nervationof the undersideof the leaf, and by the presenceof laciniaeon
the glands.Althoughit merits taxonomic recognition,I do not considerit distinct
enough to justify maintainingit as a separatespecies.
15c. Trigonianiveavar pubescens(Cambessedes)Lleras,stat nov.

pubescensCambessedes,
Trigonia (Hippocrateaceae) St. HilaireFl. Bras.Mer.2: 114.
1829;Grisebach,Linnaea22: 28. 1849;Warming, (Trigoniaceae)Mart.Fl. Bras.
4. 1967.
13(2):135. 1875;Reitz,Fl. Ill.Catar.1(Trigoniaceae):
Leaveselliptic to ovate-elliptic,thinly lanate-pubescentbeneath (having a

grayishor brownishaspect), the marginsentire;petioles 5-8 mm long. Flowerbuds
ca 3.0 mm long; pedicels 2.0-5.0 mm long, the bracts 1.0-3.0 mm long. Fruit
3.0-4.0 mm long; the endocarpvelutinous-tomentose.
Type. St. Hilaire2205 Brazil,MinasGerais,fl (holotype MPU;isotype P).
Distribution.Found in coastal forests from Parato Santa Catarina;absent or
not known from Ceara,south to Bahia.
BRAZIL.Parf:Fr6es 29747 fl (IPEAN).Maranhao:Prance58579 fl (F, MO,NY, U,
US). MinasGerais:MendesMagalhaes765 fl (IPEAN,US); Hatschbach26974 fl (NY); St.
Hilaire27 fl (P). Rio de Janeiro:Schott 5982 fr (F, US, W). Sao Paulo:St. Hilaire1234 fl
(F, P). Parana:Hatschbach3722 fl (US);Hatschbach6782 fl, fr (US). SantaCatarina:Gevieski
62 fl (NY, US);Reitz 1 748 fl (GH);Reitz 2016 fl (US);Reitz & Klein2549 fl (US);Smith
5795 fr (US); Ule 1029 fl (G, US).
This varietycan be recognizedby its broaderleaves, the typical grayor brown

aspect of the leaf pubescence, and by the relativelylong peduncles. Although I
agreewith Cambessedesin recognizingit as taxonomicallydistinct, I considerthat
the distinctionsdo not merit specific status. This varietyincludesall the material
previouslyknown as Trigoniapubescens.
16. Trigoniakillipii Macbride,(Trigoniaceae)Fl. of Peru,Publ. Field Mus.Nat.

Hist. 13(3): 95. 1950.
Shrubor scandentshrub,the branchesterete, lenticellate,when young, lanate

to strigose,becomingglabrouswith age. Stipulesnarrowlytriangular,membrana-
ceous, 15.0-25.0 mm long, 1.5-2.5 mm wide, slightly strigose,persistent;petioles
6.0-8.0 mm long, ca 1.5 mm thick, densely strigose;lamina elliptic to obovate-
elliptic, 6.0-13.0 cm long, 3.0-7.5 cm wide, subcoriaceous,smooth, the margins
entire, the apex acute to acuminate,the base obtuse;venationeucamptodromous,
the midribplane above, prominulousbeneath, strigoseon both surfaces,the secon-
darynerves 14-16 pairs,usually opposite to subopposite,usuallyless than 10.0 mm
from one another;intercostalpubescencevery slightlyvillous above, lanatebeneath.
Inflorescencesterminaland subterminalaxillary panicles, sometimes reduced to
racemes,to 10.0 cm long. Flowersin groups(cincinni) of 1-3 (usually 1); peduncles,
when present,0.1-0.5 mm long, strigose,the bractsovate, 4.04.5 mm long, 1.5-2.5
mm wide, the pedicels 0.5-1.0 mm long, ca 0.5 mm thick, strigose;sepalsoblong to
deltoid, 4.0-6.0 mm long, 2.0-4.0 mm wide, strigose on the exposed portions,
lanate on the protected areas,acute to roundedat the apex; standard6.0-6.5 mm
long, 4.5-5.0 mm wide, the pouch extending 1/3 of the length, the upperportion
erect to revolute, barbateat the throat, the wings spathulate, 5.5-6.0 mm long,
2.5-3.0 mm wide, the apex rounded,densely barbateat the base, the keel petals
4.5-5.0 mm long, 2.5-2.8 mm wide, the pouch extending along most of the length,
the apex revolute;stamens8, 6-7 fertile, 1-2 staminodes,the filaments2.5-3.0 mm
long, free along /? of the length, the anthersoblong to obovate, 0.6-1.1 mm long, ca
54 Flora Neotropica
0.6 mm wide, one of them smaller;glands2, 2-lobed, more or less rectangularto

trapezoid,ca 0.6 mm high, 1.0 mm long, glabrousto slightly pilose; style 2.5-3.0
mm long, barbate,the stigmatrilobate, approximatelytriangular,0.5-0.6 mm per
side; ovary subglobose, 1.0-1.5 mm in diameter,barbate-pubescent,the ovules
numerous.Fruit 10.5-11.0 cm long, the valves9.0-10.0 mm wide; exocarp thin,
fleshy, velutinous-tomentose;endocarpcartilaginous.Seeds not seen.
Type. Macbride5513, Peru,Junin, La Merced,fl (holotype F, isotype G).
Distribution.Tropicalrainforestsin westernAmazonia.
PERU.Killip& Smith 23846 fl (F, NY, US);Schunke2299 fr (MO,NY). BRAZIL.
Amazonas:Lleras,Stewardet al 16995 fl (INPA,MG,NY). Rond6nia:Pranceet al 5195 fl
(INPA,MG,NY);Pranceet al 6711 fl (INPA,MG,NY).
This speciesis a close relativeof Trigoniaechiteifolia Rusby, from which it
differsin its largerflowers, broaderbracts and bracteoles,largernumberof secon-
darynerveson the leaves, and in the shorterpeduncles.
17. TrigoniapaniculataWarming,(Trigoniaceae)Mart.Fl. Bras. 13(2): 132. 1875.
TrigoniaschottianaWarming,(Trigoniaceae)Mart.Fl. Bras. 13(2): 133. 1875. Type.

Schott 1677, Brazil,Rio de Janeiro,fl (lectotypeW;isotypes F, W).
Shrub or treelet, the branchesterete, lenticellate, densely tomentellous,

becomingglabrouswith age. Stipulescaducous(nv); petioles 5.0-8.0 mm long, ca
0.3 mm thick, densely tomentellous;laminaoblong to elliptic, 5.0-10.0 cm long.
2.5-5.0 cm wide, the marginsrevolute,the apex acuminate,the base roundedto
slightly cordate;venation eucamptodromous,densely tomentellous along the mid-
rib above, densely tomentellousbeneath, the midribprominentabove, prominu-
lous beneath, secondaryveins 8-9 pairs,tertiaryvenation reticulate,visible beneath;
intercostal pubescence sparselylanate or arachnoidon the upper surface of the
young leaves, becoming glabrouswith age, villous beneath, impartinga typical
olive-greencoloration, white-lanate along the revolute margins,in exsiccatae.
Inflorescencesterminaland subterminalaxillary pyramidalpanicles, 5.0-15.0 cm
long. Flowersin groups(cincinni)of 1-2; pedunclesshort or absent, to 0.4 mm long;
pedicels 1.0-1.9 mm long, ca 0.2 mm thick, densely tomentellous, the bractsand
bracteolesof equal size, linear, 1.5-2.5 mm long, tomentellous;sepalsoblong or
ovate, 1.5-2.5 mm long, 0.5-0.8 mm wide, tomentellous;standard3.54.0 mm long,
ca 2.0 mm wide, the pouch extending along ? of the length, nasiform,barbateat
the throat, the wings narrowlyspathulate,2.5-3.0 mm long, ca 0.5 mm wide,
barbateat the base, the keel petals 2.0-2.5 mm long, ca 2.0 mm wide, the pouch
centered on the petal; stamens 9-10, 6-7 fertile, 34 staminodes, the filaments
0.9-1.2 mm long, connate for most of the length, the antherssubglobose,0.2-0.3
mm in diameter;glands2, 2-3-lobed,irregular,ca 0.3 mm per side, glabrous;style
glabrous,0.9-1.1 mm long, stigma trilobate, ca 0.2 mm in diameter;ovary sub-
globose,ca 0.5 mm in diameter,villous,the ovules numerous.Fruit oblong, truncate
at base, 1.5-2.0 cm long, the valvesca 1.0 cm wide; exocarpyellow-villouswhen
immature,turningglabrouswith age; endocarpwoody. Seeds 1-2(-3) per locule,
subglobose,ca 0.5 mm in diameter,reddish-brownpubescent,the trichomesca 5.0
mm long.
Type. Glaziou 2938, Brazil, Rio de Janeiro, Larangeiras,fl (lectotype C;
isotypes BM,BR, P).
Distribution.Known from forest edges and gallery forests from eastern

central Brazil.
BRAZIL.MinasGerais:Irwin2195 fl (F, NY, US);Irwin2713 fl buds (F, NY);Mexia
4516 fl (G, GH, MO,NY, P. US);Mexia4705 fr imm (F, G, GH,MICH,MO,NY, P). Rio de
Janeiro:BarthP2 fl (US); Glaziou3882 fl (C, P). Glaziou5793 fl (P, US);Glaziou10728 fl
Houlletsn fl (BR);Luschnath
(C, P);Pereira4256 fl (F); Vauthier168 fl (G, P). Guanabara:
sn fl (BR). Sao Paulo:Burchell4029 fl buds (BR, G);Burchell4608 fr imm (BR, G, P);Hoehne
28821 fl (A, NY, US).
This species is closer to Trigoniaeriospermathan to any other. It can be
separatedfrom T. eriospermaby the presenceof axillarypanicles,the largerinflor-
escences, and especiallyby the typical white-lanateborderon the leaf margins,a
characterwhich is found in no other species of the genus.
Of the materiallisted by Warming,I have selected Glaziou2938 as the lecto-
type for this species. I cannot find the differencebetween Trigoniapaniculataand
T. schottianaindicatedby Warming;the two cannot be distinguished.Schott 1677
is consideredas the lectotype of the name T. schottiana.
18. Trigoniavillosa Aublet, Hist. P1.Guian.Fr. 1: 338. pi. 149. 1775;Poiret,

Encyc. Meth. Bot. 8: 98. 1808; de Candolle,Prodr.1: 571. 1824;
Grisebach,Linnaea22: 28. 1849; Warming,Mart.Fl. Bras. 13(2): 137.
1875; Stafleu, in Pulle Fl. Surin.3(2): 176. 1951.
Trigoniavillosavarobtusatade Candolle,Prodr.1: 571. 1824. Type.Perrottet261,

FrenchGuiana,fl (holotype G; isotype G).
Trigoniavillosavarcuneata de Candolle,Prodr.1: 571. 1824. Type. Perrottet259,
Trigoniavillosavar oblonga de Candolle,Prodr.1: 571. 1824. Type. Perrottet260,
Trigoniamollis de Candolle,Prodr.1: 571. 1824. Type Martius179 Brazil,Guanabara,
Rio de Janeiro,Corcovado,fl (holotype G, isotype M).
Trigoniacepo Cambessedes,(Hippocrateaceae) St. HilaireFl. Bras.Merid.2: 115. 1829.
Type. St. Hilaire125, Brazil,Guanabara,Rio de Janeiro,fl (Holotype MPU;
isotypes F, G, P, US).
Shrub,the branchesterete, slightly lenticellate, densely golden-brownstrigose

when young, becoming glabrouswith age. Stipules subulate, 3.0-6.0 mm long,
1.5-2.0 mm wide, coriaceous, caducous;petioles 3.0-10.0 mm long, ca 1.0 mm
thick, very densely golden-brownstrigose;lamina broadly elliptic to obovate,
sometimes oblong, inequilateral,5.0-14.0 cm long, 2.0-8-5 cm wide, subcoriaceous,
the marginsentire, the apex acute to acuminate,the base cuneate to obtuse; vena-
tion eucamptodromous,densely golden-strigosepubescent,the midribplane above,
prominulousto prominent beneath, the secondarynerves 6-9 pairs, inserted at
angles of ca 70? to midrib;intercostal pubescencevery sparselystrigose above,
yellowish or greenish-whitelanate beneath. Inflorescencesterminaland subterminal
axillarypaniclesto 25.0 cm long. Flowersin groupsof 1-3, commonly 1-2;ped-
uncles 0.5-3.0 mm long, 0.7-0.9 mm thick, strigose,the bractssubulateto linear,
sometimes ovate, 2.0-3.0 mm long, 0.5-1.5 mm wide, strigose,the pedicels 1.8-2.8
mm long, 0.5-1.5 mm thick, strigose,the bracteolessubulateto linear, 1.0-2.0 mm
long, 0.4-0.7 mm wide, strigose;sepals ovate or oblong, 4.0-5.5 mm long, 2.0-3.5
mm wide, acute or roundedat apex, lanate along protected portions, strigosealong
exposed areas;standard5.0-7.0 mm long, the pouch extending to ?i of the length,
56 Flora Neotropica
with the upper half revolute, irregularat apex, barbateat the throat, the wings
spathulate,6.0-6.5 mm long, 2.0-3.0 mm wide, barbateat the base, the keel petals
4.0-5.0 mm long, 3.04.0 mm wide, the pouch extending along 2/3 of the length,
barbateat the base;stamens 10-11(-12), 6-7 fertile, staminodes3-4, the filaments
2.5-3.0 mm long, free for 1/3 of the length, the anthersobovate or oblong, 0.5-0.7
mm long, ca 0.4 mm wide; glands2, 2-3 lobed, the lobes deltoid, 0.2-0.3 mm per
side, glabrous;style 2.5-2.8 mm long, glabrousor slightly villous, the stigmatrilo-
bate, ca 0.2 mm in diameter;ovarysubglobose,ca 1.0 mm in diameter,barbate
pubescent,the ovules numerous.Fruit4.5-11.0 cm long, the valves6.0-20.0 mm
per side; exocarp thin (ca 0.7mm) fleshy, with reddish-brownor yellowish-brown
velutinous-tomentosepubescence;mesocarpwoody, separablefrom the endocarp;
endocarpthin, sometimesdensely coveredwith long, brown velutinous-tomentose
pubescence,to glabrousor nearlyso. Seeds ca 20 per locule, ovoid, barbatepubes-
cent, the trichomesto 20.0 mm long, spirallydisposedaroundthe seed.
This speciesis dividedinto two varieties.
Key to the Varietiesof Trigoniavillosa
1. Petioles over 5.0 mm long;fruitsunder7.5 cm long;endocarpdenselypubescent.

18 a. var villosa
1. Petioles 3.04.0 mm long; fruitsover9.0 cm long;endocarpglabrousor nearlyso.
18 b. var macrocarpa
18a. Trigoniavillosavar villosa
Leaves with petioles 5.0-10.0 mm long. Fruit 4.5-7.5 cm long; exocarp

usuallyreddish-brown;endocarpdenselyvelutinous-tomentosepubescent.
Type. Aublet sn, French Guiana,"Cayenne 1775," fl, fr (lectotype BM).
Distribution.Known from coastal areasin Guyana, French Guiana,the
states of Amapa and Para, Brazil, and as a significant range disjunction, from the
vicinity of Rio de Janeiro.
GUYANA. Jenman 1083 fl (U); Jenman 6736 fl (NY, U). FRENCH GUIANA. Leblond
36 fr (G); Poiteau 1826 fr (P); Herb. Moquin-Tandon "Fragment d'Aublet" fl (P); Broadway
662 fr (GH, NY, US); Lemee sn fl, fr (P). BRAZIL. Amapa: Pires & Cavalcante 52283 fl
(IPEAN, MG, NY, U, US), Pires & Cavalcante 52522 fl (IPEAN, MG); Para: Fr6es 20368 fl
(IPEAN, NY, US), Pires & Silva 4570 fl (IPEAN); Prance & Silva 58711 fl (IPEAN);Silva, N.
3016 fl (IPEAN). Guanabara; Guillemin 696 fl, fr (F, G, P);Domingues 239 fl (NY);Dusen 108
fl (GH).
This variety includes all of the material formerly known as Trigonia mollis
and T. cepo, as well as the three varieties recognized by de Candolle.
De Candolle's varieties based on leaf characters are untenable, as it is possible
to assign varietal epithets to portions of the same collection following his usage.
Trigonia cepo has long been known to be a synonym for T. mollis. Although there
is a very significant range disjunction between those elements formerly referred to
as T. mollis (Rio de Janeiro), and T. villosa (the Guianas and northern Brazil), I
find no taxonomically significant distinction.
After careful evaluation of the material, I can find no justification for main-
taining so many varieties within the species, or for splitting this taxon into several
species, as it has been done in the past.
I have designatedthe materialdeposited at the BritishMuseumas the lecto-

type, there is no collection of Trigoniavillosain Aublet'sherbarium,nor, to my
as
knowledge,in the generalcollection of Paris.Two specimens,one from the Poiret
Herbarium,and the other from the Moquin-TandonHerbarium,are annotatedin
pencil as "Fragmentsd'Aublet."althoughthey match the lectotype fairly well, I
hesitate to designatethem as isotypes.
18b. Trigoniavillosa var macrocarpa(Bentham)Lleras,stat. nov.
TrigoniamacrocarpaBentham,LondonJour.Bot. 2: 373. 1843.
Leaveswith petioles 3.0-4.0 mm long. Fruit 9.0-11.0 cm long; exocarp

usuallyyellowish-brown;endocarpglabrousor nearlyso.
Type. Schomburgk54, Guyana,EssequiboRiver,fl, fr (holotype K, isotypes
C, CGE,F, G, NY, US, W).
Distribution.Along riverbanksin periodicallyflooded forests.
GUYANA.Schomburgk54 (Type). BRAZIL.Amazonas:ChagasINPA9 71 fr (INPA).
Ceara:Schery417 fl (GH,MO),Ducke216 fl, fr (NY, US).
This varietycorrespondsto Bentham'sTrigoniamacrocarpa.I do not consider
that it merits specific status, as morphologicallyit is very close to varvillosa,from
which it differs in the shape of the leaf, and the size and pubescenceof the fruit.
19. TrigoniabolivianaWarming,(Trigoniaceae)Mart.Fl. Bras.13(2): 134. 1875.
TrigoniasimplexWarmingvarpilosulaKuntze,Rev. Gen. 3(2): 18. 1898. Type. Kuntze

sn Bolivia,Yapagani,fl (holotype NY; isotype US).
Shrub, the branchesterete, slightly striate, lenticellate, tomentellous when

young, becomingglabrouswith age. Stipulesconnate at base, triangularto subulate,
slightly tomentellous;petioles 5.0-8.0 mm long, 0.4-0.8 mm thick, strigose;lamina
elliptic to obovate-elliptic,4.0-7.0 cm long, 2.0-3.5 cm wide, chartaceous,the
marginsentire, the apex acuminate, the base obtuse to subrotund;venation
eucamptodromous,glabrousabove, slightly villous beneath, the midribplane above,
prominulousbeneath, secondaryveins 7-8(-9) pairs, plane above, prominulous
beneath, tertiary venation not visible;intercostalpubescence sparselystrigulose
above,lanatebeneath.Inflorescencesterminaland axillaryracemes,4.0-8.0 cm long.
Flowersin groups(cincinni) of 1-2; peduncles0.1-1.0 mm long, villous, the bracts
linear, 3.0-5.0 mm long, curved upwards,villous; peduncles 1.0-2.0 mm long,
villous, the bracteoleslinear, 1.0-2.0 mm long; sepals oblong or ovate, 3.04.0 mm
long, 1.0-1.5 mm wide, strigose;standard4.6-5.0 mm long, 2.5-2.8 mm wide, the
pouch extendingalong 1/4 of the length, barbateat the throat, the wings spathulate,
3.0-4.0 mm long, 1.2-1.5 mm wide, barbateat the base, the keel petals 3.0-3.5 mm
long, 3.5-4.0 mm wide, the pouch coveringmost of the petal;stamens8-9, with 6-7
fertile, staminodes 2, the filaments 2.0-2.2 mm long, connate along 2/3 of the
length, the antherssubglobose,ca 0.3 mm in diameter;glands2,2-3-lobed,irregular,
ca 0.4 mm per side; style slender, 1.9-2.1 mm long, glabrous,the stigmatrilobate,
ca 0.2 mm in diameter;ovarysubglobose,ca 0.4 mm in diameter,villous, the ovules
numerous.Fruit oblong, 2.0-2.5 cm long, the valvesca 1.0 cm per side, cornateat
the apex, the horn ca 4.0 mmlong; exocarp rugose,glabrouswhen mature;endo-
58 Flora Neotropica
carpwoody, reddish-brown.Seeds 3-4(-5) per locule, subglobose,ca 2.0 mm long,

barbate-villous,the trichomesca 5.0 mm long.
Type. Cuming214, Bolivia,fl (holotype W;isotype F).
Distribution.Known only from Bolivia,thus far only from the Departmentof
SantaCruz.
BOLIVIA.SantaCruz;Bridgessn fl (CGE,G); Herzog1417 fl, fr (G, U); Steinbach
8132fl, fr (F, GH,NY,U, US).
This species is very close to Trigoniaeriosperma,especiallyto subspsimplex,
with which it has been associatedin the past. As with other species associatedwith
T. eriosperma,it is possible that with more data it will be impossibleto maintain
them as separatespecies. I am recognizingT. bolivianaas a separatespecies based
on the differencesin the morphology of the fruit which has cornate valves, a
characterfound in no other Trigonia.In addition to the fruit character,T. boliviana
differs from T. eriospermain the longer bracts and bracteoles, and the shorter
peduncles.
20. TrigoniabracteataLleras,sp nov
Frutex, ramulisjuvenilibusstrigosis,glabrescentibus,lenticellatis. Stipulae

connatae,triangulares,6.0-8.0 mm longae, 4.0-5.0 mm latae, chartaceae,strigulosae.
Folia opposita, petiolo 12.0-14.0 mm longo, 1.0-2.0 mm crasso,strigoso;laminae
ellipticaevel obovatae,12.0-18.0 cm longae, 4.5-9.0 cm latae, subcoriaceae,margine
integrae, apice acuto vel acuminatovariante,basi obtusa, supraleviter strigosae,
infra strigosae;costa media supraplanainfra prominens,strigosa,costis secundariis
6-8-jugis.Inflorescentiaein paniculisterminalibusvel axillaribusdispositae,8.0-17.0
cm longae. Flores saepe cadentes,in cincinnos 1-4-floraes;axibus cincinnorumad
3.0 mm longis, ca 0.8 mm crassis,strigosis;pedunculi0.1-1.0 mm longi, 0.7-0.9 mm
crassi,strigosi, bracteis persistentis,triangularis,2.0-5.0(-7.0) mm longis, 0.8-2.0
mm latis, marginesaepe glanduloso-papillatis, strigosis;pedicelli 2.0-3.0 mm longi,
0.4-0.5 mm crassi, strigosi, bracteolis persistentibus,subulatisvel triangularibus,
1.5-3.0 mm longis, 0.5-1.0 mm latis, margineut in bracteis,strigosis;sepalaovata
vel oblonga, 3.5-4.5 mm longa, 1.5-2.2 mm lata, lanatavel strigosa;vexillum 3.54.0
mm longum, 2.8-3.2 mm latum, usque ad 1/3-1/4 longitudinalitersaccatum,apice
revolutum,intus barbatum,alae late spathulatae,3.2-3.5 mm longae, 2.0-2.3 mm
latae, concavae,glabrae;carinaepetalasaccata,2.4-2.6 mm longa, ca. 1.0 mm lata;
stamina8(-10?), sterilia2(-4?), fertilia6, filamentisad mediumconnatis, 1.6-1.8
mm longis, antherisoblongis vel ellipticis, 0.5-1.0 mm longis, 0.3-0.4 mm latis,
glandulae2, bilobae, 0.5-0.7 mm latae, a fronte visaeleviterlabiatae,intus villosae;
stylus erectus, 1.8-2.0 mm longus, villosus,stigmatecirculari,ca 0.5 mm diametro,
albo; ovariumsubglobosum,0.8-1.0 mm diametro,3-loculare,villosum, ovulisin
quoque loculo numerosis.Fructusjuvenilisoblongus,dense villoso-tomentosus.
Type; Steyermark& Rabe 96588, Venezuela,Barinas,35 km SWof Santa
Barbara,fl, imm fr (holotype U; isotypes US, VEN).
Distribution.Known only from type gathering.
This species can easily be distinguishedby its largebractsand bracteoles,that
persistafter the flowershave fallen. In some cases, rosettes of bractsand bracteoles
can be seen on the floweringaxes, even when there are no longer any flowers,hence
the specific epithet for the species.
Superfically,Trigoniabracteataappearssimilarto T. virens;however,it differs

in enough significant charactersto be treated as distinct. Trigoniabracteatahas
staminodeswhile T. virenshas only fertile stamens;T. virenshas a unilocularovary,
whereasin T. bracteatait is 3-locular.The lack of fruitingmaterialmakesit diffi-
cult to establishthe relationshipsof these two species to other species of the genus.
21. TrigoniarytidocarpaCasaretto,Nov. Stirp. Decand. 76. 1845.
TrigoniaglaziovianaWarming,(Trigoniaceae)Mart.Fl. Bras.13(2): 129. 1875. Type.

Glaziou733 Brazil,Rio de Janeiro,fl (holotype C;isotypes BR, P).
Subscandentshrub(fide Casaretto),the branchesterete, lenticellate,toment-

ellous, becoming glabrouswith age. Stipules subulate, bifid, to 2.0 mm long,
tomentellous; petioles 5.0-10.0(-13.0) mm long, 0.8-1.5 mm thick, glabrousor
nearly so; laminaoblong-ellipticto elliptic, sometimesbroadlyelliptic, 5.0-11.0 cm
long, 2.5-5.0(-7.0) cm wide, the marginsentire to very slightly revolute,the apex
abruptlyacuminate,the base acute to obtuse;venationeucamptodromous,glabrous
above, very slightly lanate beneath, the midribdepressedto plane above, prominent
beneath, secondaryveins 6-8(-9) pairs,tertiaryvenation reticulate,visiblebeneath;
intercostalpubescenceabsent above, sparselyarachnoidto lanate beneath. Inflores-
cences terminal and subterminalaxillary panicles, 5.0-15.0 cm long. Flowers in
groups (cincinni) of 1-4, usually grouped dichotomously in racemoseultimate
inflorescences;pedunclesand pedicels of variablelength, 0.4-1.5 mm long, dimin-
ishingtowardsthe apex of the inflorescence,tomentellous, the bractsand bracteoles
of equalsize, triangular,1.0-2.5 mm long, tomentellous,sometimessheathingthe
pedunclesand pedicels;sepals ovate or oblong, 2.5-3.0 mm long, 1.0-1.5 mm wide,
tomentellous, sometimeslaciniateat the margins;standard3.4-3.8 mm long, the
pouch extending to 34 of the length, barbateat the throat, the wings narrowly
spathulate(to almost linear),2.8-3.0 mm long, 0.4-0.5 mm wide, barbateat the
base, the keel petals 2.5-2.8 mm long, 0.9-1.5 mm wide, the pouch along the upper
portion; stamens 9-10, with 6 fertile, staminodes34, the filaments 1.5-2.0 mm
long, connate for 2/3 of the length, the anthers0.2-0.3 mm long, ca 0.2 mm wide;
glands2, 2-3-lobed, irregular,ca 0.2 mm per side, barbateor glabrous;style 1.5-1.8
mm long, villous, the stigmatrilobate,ca 0.2 mm in diameter;ovarysubglobose,ca
0.5 mm in diameter,the lateralseptae not fused at the center, the ovules numerous.
Fruit 2.8-3.0 cm long, broadly oblong;the exocarp rugose,yellow-velutinouswhen
young, becomingglabrouswhen mature;endocarpvery slightly tomentellous. Seeds
ovate, ca 0.4 mm in diameter,the trichomesca 5.0 mm long.
Type. Casaretto1956, Brazil,Rio de Janeiro,fl fr (holotype TO;isotype G).

Distribution.A species probablyendemic only to coastal forests in Rio de
Janeiro,Guanabaraand northernSao Paulo.
BRAZIL. Rio de Janeiro: Glaziou 2506 fl (BR, C, P); Glaziou 3670 st (BR, C, P);
Glaziou 12499 fl (BR, C, G, P); l.uschnath sn imm fr (BR); Riedel 660 fl, fr (G);Schott 5980 fr
(F, US, W). Guanabara; Glaziou 10729 fl (IPEAN, K); Glaziou 6877 fl (C, F, P); Luschnath sn
fr (BR). Sao Paulo: Martius sn fl (M).
Trigoniarytidocarpacan be separatedfrom T. eriospermaby its larger,more

complex inflorescences,its generallylargerleaves, and by its unique rugose,crested
fruit, as well as severalother minor characters.
60 Flora Neotropica
The relationshipsof this species are closer to Trigoniaeriospermathan to any

other. Warminginterpreted T. crotonoides (T. eriospermavar eriosperma)as an
intermediatebetween this species and T. simplex (T. eriospermavar simplex);
althoughin a sense this may be true, I find that T. rytidocarpais distinct enough to
merit specific status, but consider T. simplex a variety of T. eriosperma.
This specieshas been commonly known as Trigoniaglazioviana,name that is
now being rejectedfor the earlierT. rytidocarpa.
22. TrigoniahypoleucaGrisebach,Linnaea22:30.1849;Warming,(Trigoniaceae)
Mart.Fl. Bras. 13(2): 140. 1875; Macbride,Publ. Field Mus.Hist. Bot.
11(2): 69. 1931; Stafleu,(Trigoniaceae)PulleFl. Surin.3(2): 175. 1951.
Trigonia hypoleuca var pubescens Warming, (Trigoniaceae) Mart. Fl. Bras. 13(2): 140.
1875. Type. Wullschlaegel8161 Suriname, fl, fr (holotype BR, isotypes IPEAN,
NY, VEN).
Trigonia xanthopila Garke, Linnaea 22: 51. 1849. Kegel 1177 Suriname, fl (holotype on
2 sheets as indicated by Garke, GOET).
Shrubor scandentshrub,the branchesterete, densely lenticellate, the young

ones strigose, becoming glabrouswith age. Stipules subulate, 2.0-3.0 mm long,
caducous;petioles 5.0-12.0 mm long, ca 1.0 mm thick, rugulose,almost glabrous;
laminaoblong elliptic or obovate, sometimeslaterallyunequal,8.0-18.0(-20.0) cm
long, 4.0-10.0 cm wide, subcoriaceous,the marginsentire, the apex acute to
acuminate,the base cuneate to obtuse;venationeucamptodromous,strigose-pubes-
cent, the midribplane above, prominulousbeneath, the secondarynerves 7-8 pairs;
intercostal pubescence absent above, lanate beneath. Inflorescencesterminaland
subterminalaxillarypanicles,to 30.0 cm long. Flowersin groupsof 14; peduncles
and pedicels of equal length, 2.0-4.5 mm long, strigose,the bractsand bracteoles
linear, 1.0-2.5 mm long, densely strigose;sepalsovate to oblong, 4.8-6.0 mm long,
1.5-3.0 mm wide, strigose along exposed portions, lanate on protected areas;
standard5.5-8.5 mm long, 4.0-5.0 mm wide, the pouch extending along lh of the
length, the upperportion revolute,barbateat the throat, the wings broadlyspathul-
ate, 5.5-8.5 mm long, 2.8-5.0 mm wide, barbateat the base, the keel petals 5.0-7.0
mm long, 2.2-3.5 mm wide, the pouch extending along /2 of the length, barbateat
the base;stamens 10-11, 6-7 fertile, 34 staminodes,the staminodeswith terminal
globose knobs, the filaments 1.5-1.8 mm long, free along the upper %,the anthers
obovate, 0.5-0.7 mm long, 0.3-0.4 mm wide; glands2, 2-3-lobate,the lobes 2.0-3.0
mm high, the upper Mformed by subulate,pointed, strigoselaciniae;style clavate,
1.5-1.8 mm long, glabrous;the stigmaca 0.3 mm in diameter;ovarysubglobose,
1.0-1.5 mm in diameter,barbate-pilose,1-locular,the ovules numerous.Fruit
5.0-7.0 cm long, the valves2.5-3.0 cm wide; exocarp coriaceous,the nervespromi-
nulous, glabrous;endocarp ending flush with the exocarp, cartilaginous.Seeds
6.0-8.0 mm long, elliptic, flattened, echinate-pubescent.
Type. Schomburgk315, Guyana,fl (holotype GOET;isotype K).
Distribution.This species is found mainly along forested, periodicallyflooded
riverbanks in Guyana, Suriname, and French Guiana and northeastern Brazil.
GUYANA. Schomburgk 224 fl (CGE, G, K, P, W); Gleason 516 fl (G, K, NY, US);
Gleason 549 fr (GH, NY, US);Jenman 1155 fl (K, NY);Jenman 1296 fl (K). SURINAME.
B. W. 2161 fl (U); Florschiitz & Maas 2741 fl (NY, U, US); Splitgerber 1136 fl (L);Mennega
371 fl (A, C, U). FRENCH GUIANA' Herb. Sagot s.n. "Ile Portal Bar." fl (F, G, GH, NY, US);
Service des Eaux & Forets 4204 fl (U). BRAZIL. Ducke 8955 fl, fr (MG); Prance et al 22982
fl (COL, F, INPA, MG, NY, US).
Trigoniahypoleuca is probablycloser to Trigoniavirensthan to any other

species. The presenceof unilocularovariesin both species suggestssimilartrendsin
evolution. Trigoniahypoleuca can be distinguishedfrom Trigoniavirens by the
lanate pubescenceon the leaves, the presenceof laciniaeon the glands,and the
presenceof staminodes,as well as other minor characters.
Warming'srecognitionof varietiesin this species appearsuntenableat present;
no clear distinctionscan be made within the species that would merit formalrecog-
nition.
Grisebach'scitation of the type as "Schomburgk313" is erroneous.He was
misled by a not altogetherclear 5 that could easily be interpretedas a 3; the situa-
tion is clarifiedon examinationof the Kew isotype on which the collection number
is unmistakeablySchomburgk315.
The materialused by Garketo typify T. xanthopila, and labelled as Kegel
1177, is accordingto the label data, constituted by two differentcollections: one
made on the SaramaccaRiverin May 1846, and an earlierone gatheredon the
CassepocreekRiverin November1845; it is impossibleto determinewhetherone of
the two sheets deposited at Goettingenis a mixed collection, or if both are mixed
collections, or if one sheet correspondsto each locality. The specimenswere
treatedby Garkeas one specimenon two sheets, and I considerthat for all practical
purposesit would be convenientto treat the holotype as such. This name is obvi-
ously a later synonym of T. hypoleuca.
23. Trigoniaechiteifolia Rusby, Bull. N. Y. Bot. Gard.4: 324. 1907; Macbride,

Publ. Field Nat. Hist. 13(3); 96. 1950.
Shrub, the brancheslenticellate, strigose pubescent, turningglabrouswith

age. Stipulessubulate,6.0-8.0 mm long, ca 2.0 mm wide, strigose-lanate,caducous;
petioles 6.0-10.0 mm long, 0.8-1.0 mm thick, strigoseto glabrous;laminainequilat-
erallyovate to obovateor oblong, 5.0-12.0 cm long, 2.0-6.0 cm wide, subcoriaceous,
the marginsentire to slightly revolute, the apex acuminate,the base obtuse to
slightly cordate;venationeucamptodromous,strigosepubescent,the midribplane
above, prominulousbeneath, the secondarynerves 10-12 pairs;intercostal areas
arachnoid-pubescentor glabrousabove, lanate beneath. Inflorescencesterminaland
subterminalaxillarypanicles,to 10.0 cm long. Flowersin groupsof 14 (usually 3);
pedunclesandpedicelsof equal length, 0.5-1.0 mm long, 0.4-0.5 mm thick, strigose;
the bractssubulateto linear,2.5-3.5 mm long, ca 0.2 mm wide, strigose,the bract-
eoles linear, 1.5-2.0 mm long, strigose;sepals ovate to deltoid, 4.0-5.0(-6.0) mm
long, 1.5-2.5 mm wide, strigose exterally, the interior lanate; standard4.5-5.0
mm long, 4.0-5.0 mm wide, the pouch extending to 2/5 of the length, the upper
portion erect or revolute, the apex rounded,barbate-pubescentat the inside of the
throat, the wingsspathulate,4.0-4.5 mm long, 1.5-2.0 mm wide, barbateat the base,
the keel petals 3.8-4.0 mm long, 3.0-4.0 mm wide, the pouch extending along 2/3
of the length; stamens8-10, fertile ones 5-7, staminodes2-3, the filaments 1.8-2.0
mm long, connate to the middle, the anthersobovate, 0.5-0.8 mm long, 0.3-0.4
mm wide; glands2, 3-lobed, the lobes deltoid, ca 0.2 mm per side, strigose;style
1.8-2.0 mm long, barbate,the stigmatrilobate,ca 0.2 mm in diameter;ovarysub-
globose, the locules not always totally fused at the center, barbate-villous,the
ovules numerous.Fruit not seen.
Type. Bang2812. Bolivia,La Paz, fl (holotype NY; isotypes F, GH, MICH,
MO,NY, US).
62 Flora Neotropica
Distribution.Known only from 3 collections.

BOLIVIA.La Paz:Krukoff10466 fl (A, F, NY). BRAZIL.Rond8nia:Black& Cordeiro
52-14678 fl (IPEAN).
The relationshipsof this species are impossibleto determineat present,due
to the lack of fruitingmaterial.
24. TrigoniasericeaHumboldt,Bonpland& Kunth, (Hippocrateaceae)Nov. Gen.

et Sp. P1.5: (4). 1821.
TrigonianajadumWarming,(Trigoniaceae)Mart.Fl. Bras. 13(2): 126. 1875. Type.

Martiussn, Brazil,Rio Solim6es,fl (holotype M;isotype M).
Shrubor treelet,the branchesterete, lenticellate. Stipulesnarrowlytriangular,

membranaceous,20.0-25.0 mm long, ca 5.0 mm wide, glabrous,caducous;petioles
9.0-15.0(-20.0) mm long, 1.0-2.0 mm thick, slightly strigose;lamina elliptic to
broadlyelliptic, 6.0-19.0 cm long, 4.0-11.0 cm wide, subcoriaceous,smooth, the
marginsentire, the apex acuminate,the base obtuse; venationeucamptodromous,
the midribplane above, prominulousbeneath, glabrousabove, tomentose beneath,
the secondarynerves 8-11 pairs, plane above, prominulousbeneath; intercostal
pubescence absent above, white-lanatebeneath. Inflorescencesloose terminaland
subterminalaxillarypanicles,to 10 cm long. Flowersin groupsof 3-10, (commonly
6-7); peduncles2.5-4.0 mm long, 0.5-0.7 mm thick, tomentose, the bractsovate,
1.5-2.0 mm long, 0.3-0.5 mm wide, tomentose;the pedicels 0.9-1.2 mm long, ca
0.6 mm thick, tomentose;sepals ovate, 3.5-5.0 mm long, 1.5-2.5 mm wide, strigose;
standard4.0-5.0 mm long, 3.5-4.0 mm wide, the pouch extending 1of the length,
the upperportion erect or revolute,the apex irregular,barbatepubescentat the
inside of the throat, the wings spathulate,3.5-4.0 mm long, 2.0-3.0 mm wide, the
apex irregular,barbateat the inside of the base, the keel petals 2.5-3.5 mm long,
3.0-3.5 mm wide, the pouch extending along 2/3 of the length, the apex irregular;
stamens 10, 6 fertile, 4 staminodes,the filaments 1.4-1.8 mm long, free for /2 the
length, the antherselliptic to oblong, 0.6-0.8 mm long, 0.3-0.4 mm wide, adhering
3 on each side of the style and completely surroundingit; glands2, 2-3-lobate,ca
1.0 mm high, 1.5 mm long, with a knob-likeprojectionat the apex of each lobe,
glabrous;style 1.5-1.7 mm long, slightly pilose or glabrous,the stigmatrilobate,ca
0.2 mm in diameter,ca 0.2 mm high; ovarysubglobose, 1.0-1.2 mm in diameter,
barbate-pubescent,the ovules numerous.Fruit 3.0-7.0 cm long, the valves0.5-1.5
cm wide; exocarp thin, fleshy, velutinous-tomentose;endocarpcartilaginous.Seeds
numerous,subglobose,ca 2.0 mm in diameter,barbate-pubescent,the trichomes
ca 10.0 mm long.
Type. Ihumboldt1859, Colombia,"Andesde Quindie,"fl (holotype P nv;
isotype P).
Distribution.Knownonly fromperiodicallyflooded rainforestsin the Amazon
basin of Colombia,Venezuela,Peruand Brazil.
COLOMBIA. Antioquia:Romero C. 2423 fr (COL).Caldas:Bro. Daniel2317 fl (US).
Valle:Hutchinson& Wright3293 fl (COL,F, US); Urihe1550 fr (COL,US). VENEZUELA.
Bolivar:Steyermark74649 fl (F, NY, VEN). PERU.Loreto:Killip& Smith 29542 fl, fr (F,
NY, US);Schunke50 fl (A, F, NY, US). BRAZIL.Amazonas:Lleras,Stewardet al P16897 fl
(INPA, MG, NY).
Trigonia sericea is probably related to T. prancei, from which it differs in the
simplerinflorescence,largerflower parts(in general),and the pubescentleaves, as
well as other minor characters.
2. HumbertodendronLeandriin Heim, Compt. Rend. Acad. Paris.229: 847.

1949; Perrier& Leandri(Trigoniaceae)Humbert,Fl. de Madagascar.
108(bis): 1-4. 1955.
Smalltree, branchesterete. Leavessimple, opposite, petiolate, with connate

stipules. Inflorescencesaxillary triflorate cymes. Flowers zygomorphic,sharinga
common peduncle;bracts2, eglandulate;sepals 5, quincuncial,unequal;petals 5,
contorted, unequal, papilionaceous;fertile stamens 6, staminodes absent, the
filaments connate at base, free above, the anthersbasifixed, bilocular,introrse,
dehiscingalong two lateralslits; pollen triporate,deltoid in equatorialview. Gland
one, strongly adpressedto the ovary. Ovarysuperior,3-winged,3-sulcate,trilocular,
the locules incompletely closed, lacking a central column; ovules 1 per locule,
attached to the interiorends of the lateralseptae. Fruit (fide Leandri)a 3-winged
samara;seed lacking endosperm;embryo with an inferiorradicle.
Type species.HumbertodendronsaboureauiLeandri.
Distribution.Monotypicendemic genus known only from the easterncoastal
forests in the MalagasyRepublic.
1. HumbertodendronsaboureauiLeandriin Heim, Compt. Rend. Acad. Paris.

229: 847. 1949; Perrier& Leandri,(Trigoniaceae)Humbert,Fl. de
Madagascar.108(bis): 2. 1955.
Tree, branchesterete, lenticellate, glabrous.Stipules oblong, 2.5-3.0 mm long,

strigose;petioles 4.5-6.0 mm long, ca 1.0 mm thick, strigose;lamina elliptic to
obovate, 2.5-5.0 cm long, 1.0-3.0 cm wide, chartaceous,the marginsentire, the
apex obtuse to emarginate,the base acute to obtuse;venationbrochidodromous,
glabrous,the midribprominulousabove, prominentbeneath, secondaryveins 4-6
pairs. Flowers in trifloralaxillary cymes; peduncles 6.0-18.0 mm long, strigose;
pedicels 4.0-7.0 mm long, strigose,the bractsand bracteolessubulate, 1.5-2.0 mm
long, strigose;sepals ovate to oblong, 4.0-5.0 mm long, 2.0-2.5 mm wide, strigose;
standard3.5-5.0 mm long, 3.0-4.0 mm wide, the pouch extending 2/3 of the length,
revoluteat the apex, slightly strigoseexternally,barbateat the throat, the wings
sublinearto obovate, 3.5-4.5 mm long, 0.5-1.5 mm wide, slightly strigose,the keel
petals deltoid to ovate, 3.5-4.5 mm long, 2.5-3.5 mm wide, strigose externally;
stamensca 4.5 mm long, the filamentsconnate to the middle, 1.5-3.5 mm long, the
anthersovate to elliptic, 1.0-1.4 mm long, ca 0.5 mm wide;gland reniform,ca 1.0
mm long, 1.5 mm wide, strigose or pilose; style 1.5-3.0 mm long, glabrous,the
stigmatriangular,ca 0.4 mm wide; ovary ovate to subpyramidal,strigoseor pilose,
1.0-1.5 mm high, 0.9-1.3 mm per side. Fruit (fide Leandri)2.0-2.5 cm long, 1.5-1.8
mm per side; wingsnarrowlysemi-elliptic.Seeds glabrous,narrowlyovate.
Type. Ramarokoto1522 RN, Malagasy,Ambodivila,near Ambila,fl (Holo-
type P; photographNY).
Distribution. Easterncoastal forests of Malagasy.Floweringin February,
fruitingin October(?).
MALAGASY. Ambila: Louvel & Perrier 14896 fl (P); Tamatave: Louvel & Perrier 14896
bis fl (P); Service Forestier 44 fl, fr (P).
Local names.Hazombaroranalahy
(Ambila),Fandrianakanga(Tamatave).
3. TrigoniastrumMiquel,Fl. Ind. Bat. Suppl. 394. 1862; Bentham& Hooker,
Gen. P1. 1: 139. 1862; Baillon, Hist. PI. 5: 91. 1873; Clhodat,Bull.
64 Flora Neotropica
Herb.Boiss. 3: 1895; Barth,Bull. Herb.Boiss. 4: 481. 1896; Engler&

Prantl,Nat. Pflanzenfam.209. 1897; Van Steenis, Fl. Malesiana1(4):
58. 1948; Ng, Tree Fl. Malaya.1: 449. 1972.
Isopteris Wall.Cat. 7261. 1831, nom. nud.
Smallto medium tree, branchesterete, lenticellate. Leavessimple, alternate,

petiolate, the marginsand acumensometimesglandular;stipulescaducous.Inflores-
cences terminaland subterminalaxillarypanicles.Flowerszygomorphic,the bracts
glandularon the margins;sepals 5, quincuncial,unequal;petals 5, contorted,
unequal,papilionaceous;fertilestamens6, staminodesabsent;the filamentsconnate
at base, free above, the anthersbasifixed, bilocular,introrse, dehiscingalong a
centralslit; pollen 3-4 porate;glands 1-2; ovary3-locular,lackinga centralcolumn;
ovules 2 per locule, pendulous,attachedto the interiorends of the lateralseptae.
Fruit a 3-wingedsamara;seeds one per locule, lackingendosperm.
Type species. Trigoniastrumhypoleucum Miquel.
Distribution.Monotypicendemic genus known from tropicalrainforestsin
the MalayPeninsula,Sumatraand Borneo.
1. TrigoniastrumhypoleucumMiquel,Fl. Ind. Bat. Suppl. 394. 1862; Chodat

Bull. Herb.Boiss. 3: 136. 1895; Barth,Bull. Herb.Boiss. 4: 481. 1896;
Van Steenis, Fl. Malesiana1(4): 58. 1948; Ng, Tree Fl. Malaya1:
449. 1972.
IsopterispenangianaWallichex Benn.,Hookerf. Fl. Br. Ind. 1: 208. 1872.

Trigoniastrum hypoleucum var oliganthum Airy Shaw, Kew Bull. 1940: 253. 1940.
Type. Native collector, Richards 1921. Malaysia, Sarawak, fl (holotype A; isotype
A).
Trigoniastrum hypoleucum var viride Airy Shaw, Kew Bull. 1940: 253. 1940. Type.
Elmer21302, Malaysia,Sabah,Tawao,fl, fr (holotype A; isotypes BR, F, GH,
M, MO,U, US).
Tree, to 30 m. Petioles 4.0-8.0 mm long, 2.0-4.0 mm wide, sulcate, sometimes

sparselystrigose,the stipulescaducous(nv); laminaoblong, 7.0-20.0 cm long, 2.5-
6.0 cm wide, glabrousabove, thinly lanate beneath, the marginsentire, with small
impressedglands,the apex acuminate,the acumencuneate, to 30.0 mm, glandular
thickened, the midribplane to depressedabove, prominulousbeneath, slightly
strigose,the secondarynerves4-6 pairs,brochidodromous.Inflorescences15.0-40.0
cm long, the bracts foliolate on lower panicles, becoming fusiform towards the
upper ones, the glands circular,fleshy. Flowers (fide Van Steenis) white; sepals
2.0-3.5 mm long, 1.0-2.0 mm wide, ovate, the apex acute, strigose;standard4.0-5.0
mm long, the pouch extending to ? the length, 2.0-3.0 mm wide, slightly pilose
externally, the wings 3.0-4.0 mm long. 1.0-2.0 mm wide, spathulate,the keel petals
3.5-5.0 mm long, oblique to oblong;filaments 1.0-2.0 mm long, connate for 2/3
of the length, the anthers0.5-1.0 mm long, ovate;glandsca 0.7 mm wide, 0.6 mm
high, reniform,puberulous;style 1.0-1.5 mm long, sparselystrigose,the stigmaca
0.2 mm diameter;puberulous;ovary0.5-0.8 mm in diameter,subglobose,strigose
to lanate pubescent. Fruit 3.0-5.0 cm long, the locules semi-elliptic,extending to
2.5 cm, the wings to 25.0 mm long, 10.0 mm wide, extending upwards, oblong, the
innerside straight, roundedon the externalmargin,nervationprominulous.Seeds
obovate, flattened, ca 20.0 mm long, 5.0 mm wide, velutinous.Seedlings(fide Van

Steenis) with epigealcotyledons, first pair of leaves opposite.
Type. Teysmann,Herb. Bogor.# 4548, Sumatra,"MangalaLamp" fl, fr
(lectotype U; isotypes GH, US).
Distribution.Forests of Indonesia,Malaysia,Singaporeand Brunei.
INDONESIA.Sumatra:Forbes sn fr (A); Forbes 3069 fl, fr (A, GH, MO,NY, US);
Forbes 3187 fr (A, NY, US); Soepadmo136 fl (A); Teysmannsub. n. Herb.Bogor. 4411
fl, fr (Syntype U). MALAYSIA:Penang:Dr. King'scoll. 1680 fl (A). Perak:Dr. King'scoll.
(Soping)6002 fr (US);Dr. King'scoll. 6768 fl (U, US);Dr. King'scoll. (Thaiping)8467 fl (F,
US); Wray3253 fr (US). Kelantan:Henderson2482A fl (A). Selangor:Kochummen94447
fl (A). Trengganu:Cockburn8259 fr (A). Sarawak:Ashton 22620 fl (A);Haron21332 fl (A);
Hou 410 fl, fr (A); Jacobs6264 fr (US);Richards2324 fl (A); Richards2448 fl (A); Wee-lek
621 fl (A); Sabah:Gibot33034 fl (A); Sinanggul30548 fl, fr (A); Singh21020 fl (A, US);
Tikun39342 fl, fr (A); Wood16258 fr (A). SINGAPORE. Ngadiman34965 fl (A); Ngadiman
36148 fl (A). BRUNEI.S. W.sub. n. Kepong80176 st (A).
Commonname. SE Borneo:Kikir:mangkudor;Asanan:tinga catu: Malaya:
marasali,mata pasak, suginara.
Trigoniastrumhypoleucumis the only species in the genus. Airy Shaw, 1940,
establishedtwo varietiesunder this species: varoliganthum,and varviridebasedon
leaf characters.I can find no justification to maintainvarietiesunderthis species,
the leavesareveryvariable.The typification of the species has been somewhatprob-
lematic, as Miquel'streatmentis obscureas to the type. He indicatedit as "Mangala
prope Lampong."SeveralTeysmannspecimensseen on loan are annotated as
"MangalaLamp,"andprobablyrepresentsyntypes. Othertreatmentsin the literature
have ignoredthis problemby not citing the type. I have annotatedthe Utrecht
specimenlabeled as Herb.Bogor.#4548, as the lectotype.
66 Flora Neotropica
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68 FloraNeotropica
LISTOFTAXA
NUMERICAL
1. Trigonia 15. T. niveaCambessedes

1. T. coppenamensisStafleu a. varnivea
2. T. rotundifoliaLleras b. varfasciculata(Grisebach)
3. T. pranceiMacbride Lleras
4. T. virensMacbride c. varpubescens
5. T. macranthaWarming (Cambessedes)Lleras
6. T. laevisAublet 16. T. killipiiMacbride
a. varlaevis 17. T. paniculataWarming
b. varmicrocarpaSagot 18. T. villosaAublet
7. T. spruceanaBenthamex Warming a. varvillosa
8. T. reticulataLleras b. varmacrocarpa(Bentham)
9. T. candelabraLleras Lleras
10. T. rugosaBentham 19. T. bolivianaWarming
11. T. eriosperma(Lamarck)Fromm& 20. T. bracteataLleras
Santos 21. T. rytidocarpaCasaretto
a. subsperiosperma 22. T. hypoleucaGrisebach
b. subspsimplex(Warming) 23. T. echiteifoliaRusby
Lleras 24. T. sericeaH.B.K.
12. T. floccosa Rusby 2. Humbertodendron
13. T. costanensisSteyermark& 1. H. saboureauiLeandri
Badillo 3. Trigoniastrum
14. T. subcymosaBentham 1. T. hypoleucumMiquel
List of Exsiccatae 69
LISTOF EXSICCATAE
The figuresin parenthesesrefer to the genusnumber(first) and to the species

number(second). The numberscorrespondto those givenin the numericallist of
taxa.
Ackermann,12 (1-lla); sn (1-15a) Calder6n,S., 1976 (1-10); 2124 (1-10)

Alemao,602 (1-15a) Casaretto,585 (1-lla); 1535 (1-lla);
Aparicioet al, 59 (l-15a) 1740 (1-1la); 1956 (1-22)
Anderson,W. 9230 (1-lla) Cavalante,P. B., 552 (1-7)
Aristequieta,L. et al, 3988 (1-15a); Chagas,J. et al, INPA5349 (1-6b)
5554 (1-15a) Chaves,213 (1-10)
Ashton, P. S., 18880 (3-1) Claussen,P., 19 (1-22); 25 (1-15a);1096
Asplund,E., 14281 (1-4) (1-lla)
Aublet,J. B. C. F., sn (1-6a) Coelho, L., INPA 1139 (1-7); INPA3722
Aviles,S., 961 (1-10) (1-6b)
Bacle, 1834 (1-15c) Croat,T., 16581 (1-10)
Baker,C. F. et al 2554 (1-10) Cuming,H., 214 (1-19)
Baldwin,J. T., 2771 (1-6b) Debeaux,48 (1-15a)
Bang,M., 2191 (1-12); 2812 (1-23) den Held, 1335 (1-22)
Barreto,M., 1467 (1-15a);7154 (1-15a); Dialer,618 (1-15b)
7156 (1-15a);8301 (1-15a); Dlurtzsn (1-15a)
8302 (1-15a) Dodge,C. W.et al, 6219 (1-10); 623b
Barth,P2 (1-17) (1-10)
Beccari,O., 3161 (3-1) Domingues,239 (1-18a)
Belem,R. P. et al, 1170 (1-15a); Donselaar,2812 (1-8); 2891 (1-8)
1185 (1-lla); 3632 (1-15a);3652 Duarte,A. P. etal,4011 (1l-a); 4529
(1-15a) (1-17);4801 (1-17)
Bernoulli,C. G. et al, 3139 (1-llc) Duchafraing,1850 (1-10)
Boilley, 2638 (1-10) Ducke,A., 179 (1-7); 1208 (1-6b); 1282
Black,G. A. et al, 47-932 (1-6b);47-989 (1-6b); 1318 (1-18a);2453 (1-15a);
(1-6b);47-1039 (1-6b);47-2091 2453 (1-15a);2900 (1-6b);6811
(1-6b);47-7957 (1-15a);52-14618 (14); 6814 (l-15a); 7228 (1-6b);
(1-6b);52-14678 (1-23) 8955 (1-22); 11586 (1-7)
Blanchet,J. S., 2029 (1-15a);2921 (1-15b); Dugand,A. et al, 919 (1-10); 921 (1-10)
3593 (1-lla). Dusen,K. H., 108 (1-18a);8696
Blanco,C., 426(1-6b);526 (1-6b) (1-15a); 12162 (1-15a);16520
Bonpland,A., sn (1-24) (l-15a); sn (1-1la)
Boon, 1069 (1-22) Eiten, G., 1731 (1-15a)
Bowie,J. et al, 10 (1-15a);sn (1-17) Elmer,21302 (3-1)
Brade,A. C., 10618 (1-15a);11215 (1-lla); Fanshawe,D. B., 2470 (1-6b);2470 (1-6b);
11329 (1-17) 6303 (1-6b)
Brenes,A. M., 12736 (1-10); 15576 (1-10) Fiebrig,K., 5133 (1-15a)
15593 (1-10) Florschiitz,P. A. et al, 1120 (1-22); 2741
Breteler,F. J. 4953 (1-6b) (1-22); 2755 (1-22)
Bridges,sn (1-19) Forbes,H. O., sn (3-1); 3039 (3-1); 3187
Bro. Elias,258 (1-10); 357 (1-10); 1105 (3-1)
(1-llc); 1202 (1-llc); 1651 (1-11lc) ForestDepartment,BritishGuiana
Broadway,W. E., 383 (1-18a);539 (1-18a); (Guyana),F 300 (1-22); F 1777
662 (1-18a) (l-6b); F 2973 (1-6b);F 799z (1-8)
Buchtien,O., 1631 (l-6b) Friedrichstahl,E. P. 924 (1-10)
Bunting,G. S. et al, 4067 (1-7) Fr6es,R. L. et al 20368 (1-18a);21293a
Burchell,W. J., 1405 (1-15a);4029 (1-17); (1-7); 22077 (1-7); 22588 (1-7);
4608 (1-17) 22946 (1-15c); 23503 (1-6b);24606
Burger,W.et al, 7841 (1-10) (1-15a);25235 (1-7); 27405 (1-18a);
B. W. (BoschwezenSuriname),631 (1-6b); 29067 (1-7); 29559 (1-7); 29747
2163 (1-22); 3289 (1-6b);6485 (1-15c); 31305 (1-6b); 32897 (1-6b);
(1-6b) 32914 (1-6b)
70 FloraNeotropica
Gabriel,1802 (1-18a); 1802 (1-6a) Hou, D., 410 (3-1)

Gandoger,A. M., sn (1-6b) Houllet,sn (l-lla); sn (1-17)
Gaillard,164 (1-7) Huber,J. E., MG1048 (1-18a)
Gardner,G., 334 (1-15a);943 (1-15a); Humbolt,A. von et al, 1870 (l-15a)
4461 (1-15c);5387 (1-22); sn (1-15a) Hutchinson,P. C. et al 3293 (1-24)
Garnier,F. A., 1075 (1-10); 1283 (1-10) Irwin,H. S. et al, 679 (1-15a);2195 ( 1-17);
4230 (1-10) 2713 (1-17); 2068 (1-15a);
Gaudichaud,C., 259 (1-15c);978 (1-1la) 20852a (1-15a);21071 (1-15a);
Geay, 1889 (1-6b); 1890 (1-6b) 28792 (1-15a);28981 (1-15a);
Gentle,P. H., 525 (1-1 c) 30860 (1-15a);30875 (1-15b);
Gevieski,A., 62 (1-15c) 31006 (1-15a) 31139 (1-15a);
Glassman,1723 (1-10) 31170 (1-15b);32565 (1-15b)
Glaziou,A. F. M., 733 (1-22); 2114 (1-lla); Jacobs,M., 5264 (3-1)
2505 (1-15a);2506 (1-22); 2938 Jenman,G. S., 1083 (1-18a); 1155 (1-22);
(1-17); 2939 (1-18a);3670 (1-22); 1296 (1-22); 6736 (1-18a)
3881 (1-1 a); 3882 (1-17);5793 Jenner,sn (1-17)
(1-17);5994 (1-22); 6485 (1-17); Jimendz,M. A., 1575 (1-10)
6877 (1-22);8670 (1-15a);9417 Kapler,A., 134 (1-22); 2008 (1-22)
(1-22); 9417 (1-lla); 10394 (1-15a); Kepong,F 8259 (3-1)
10728 (1-17); 10729 (1-22); 10730 (1-15a); Kichl,SP 3766 (1-15a)
12499 (1-22); 13480 (1-lla); Killip,E. P. et al, 14396 (1-1lc); 23846
13811 (1-1la); 14688 (1-22); 14688a (1-16); 28136 (1-5); 29539 (14)
(1-7); 14689 (1-lla); 14690 (1-15a); King'scoll. 1680 (3-1); 6002 (3-1); 6410
sn (1-22); sn (1-18a). (3-1); 6729 (3-1); 6768 (3-1); 7709
Gleason,H. A. et al, 549 (1-22);576 (1-22); (3-1); 8467 (3-1)
576 (1-22); 877 (1-22) Klein,R. M., 547 (1-15c); 735 (1-15c)
Gomez, 1836 (1-lla) Kluge,H. G., 1004 (1-24); 2954 (1-5);
Guedes,M., 588 (1-15a);483 (1-15a) 3028 (1-5)
Guedes,T. N., 134 (1-6b);483 (l-15a); Kochummen,94447 (3-1); 97743 (3-1)
588 (1-15a) Krukoff,B. A., 6149 (14); 6285 (14);
Guillemin,181 (1-lla); 245 (1-15a);654 8339 (14); 8424 (14); 10338
(1-17); 696 (1-18b) (1-6b); 10466 (1-23); 10956 (1-6b)
Guillot,sn (1-15a) Kuntze,C. E. O., VI 92 (1-19)
Halle,847 (1-18a) Labroy,M., sn (1-7)
Haron,O., 21332 (3-1) Lalande,sn (1-15c)
Hassler,5362 (1-15a);5588 (1-15a);8416 Lambert,sn (1-6b)
(1-15a) Langsdorff,sn (1-17); 1821 (1-lla);
Hatschbach,G. et al 3722 (1-15c);6782 Lanjouw,J. et al 323 ( 1-22); 349 (1-6b);
(l-15c); 6861 (1-15a); 12617 1168 (1-22); 1221 (1-22); 2416
(l-15c); 26974 (1-15c); 14278 (l-6b)
(1-15a); 15841 (1-15a); 19042 Leblond,J. B., 35 (1-6b); 36 (1-18a)
(115a) Lemde,A. M.V., sn (1-18a)
Haught,O., 1800 (14); 1907 (1-4); 2211 Leprieur,F. R., 237 (1-18a);238 (1-6a);
(1-4);4186 (1-10);4227 (1-10); 1833 (1-18a); 1834 (1-18a); 1838
4866 (1-10) (1-18a); 1839 (1-18a); 1840 (1-18a);
Hayes,S., 720 (l-lc) sn (1-18a);sn (l-6b)
Henderson,24824 (3-1) Level,J. S., 101 (1-7)
Herb.C. M. Lemann,sn (1-15a) Levy,P., 192 (1-10); 1073 (1-10)
Herb.Exp ColonialMinisterede la Marine, Lleras,E. et al, P 17115 (1-3)
sn (1-22) LimaD. A., 49230 (1-15a)
Herb.Maire,sn (1-6b) Lindeman,J. C. et al, 4372 (1-15a);4408
Herb.Moquim- Tandon,sn (1-18a) (l-15a); 4496 (1-15a);5785 (1-6b)
Herb.Musel- Paris,sn (1-18a) Lindley,sn (1-lla); 749 (1-lla) 749 (1-lla)
Herb.Poiret,sn (1-18a) Lisb6a,A., MG2388 (-15Sa);MG2433
Herb.Richard,sn (1-18a);sn (1-6b) (1-15a)
Herb.SchwackeIII, 281 (1-7) Lofgr6n,A., 632 (1-15a);718 (1-15a)
Herb.Zuccarini,sn (1-17) Luetzelburg,P. von 20058 (1-22); 20394
Herzog,1417 (1-19) (1-15c); sn (l-15a)
Hohne,F. G., 24890 (1-18a);28821 Lund,P. W., sn (1-15a)
(1-17); sn (l-15a) Lundell,C. L. 7021 (1-llc)
List of Exsiccatae 71
Luschnath,B., sn (1-17); sn (1-22); sn Poiteau, 1826 (1-18a);sn (1-6b)

(1-lla) Prance,G. T. et al, 2657 (1-7);4945 (1-7);
Maas,P. et al P12822 (-6b) 5195 (1-16); 6711 (1-16); 14529
Me Rae, 1678 (1-15) (1-16); 15200 (1-7); 22523 (1-22);
Macbride,J. F., 5513 (1-16) 22982 (1-4); 58579 (1-15c);58711
Magalhaes,G. M., 765 (1-15c); 5343 (1-18a)
(1-15b); 17247 (1-15a) Pursell,R. A., 8911 (1-15a)
Maguire,B. et al, 24857 (1-1); 24858 Quelchet al, 222 (1-15a)
(l-6b); 30834 (1-7);41134 (1-15a); Quiros,C. M., 734 (1-10); 848 (1-10)
43907 (1-18a);56795 (1-16b) Regnell,A. F., 281 (1-7); 1009 (1-15a)
Maingaus,135 (3-1) Reitz, P. R. et al, 1110 (1-15c); 1748
Marinho,7(1-18a) (1-15c); 2016 (1-15c); 2216 (1-15c);
Martin,J., 100(1-18a);sn (1-6b) 2374 (l-15c); 2549 (1-15c); 3016
Martius,K. F. P. von, 122(1-lla); 123 (1-15c)
(1-lla); 176 (1-18a);625 (1-15a); Renner,sn (1-15a)
989 (1-17); sn (1-lla); sn (1-15a);sn Richardset al, 1921 (3-1); 2324 (3-1);
(1-18a);sn (1-22) 2448 (3-1)
Matuda,E., 2517 (1-10); 16548 (1-10); Riedel,L., 10 (1-lla); 12 (1-lla);660
17630 (1-llc) (1-22); 661 (1-lla); 819 (1-15a);
Maxon,W. R. et al, 7675 (1-10); 7686 1198 (1-15a); 1228 (1-17); sn
(1-10) (l-15a); sn (1-1la); sn (1-22)
Meijer,43892 (3-1) Rodrigues,J. S. et al, 44 (1-7)
Meli'on, M., 134 (1-6b); 136 (1-6b); 141 Rodrigues,J. V. 233 (1-10); 391 (1-10);
(1-6b);230 (1-6b); 1865 (1-18a); 2078 (1-10)
an (1-6b) Rodrigues,W., 268 (1-6b);726 (1-7); 9009
Mennega,A. M.W., 371 (1-22) (1-6b)
Mexia,I., 4120 (1-lla); 4516 (1-17);4521 Rohr,J. P. B. von, 29 (1-18a);30 (1-6b)
(1-lla); 4705 (1-17);4768a (1-lla) Romero,C. R., 1124 (1-10); 2423 (1-24)
Miers,J., sn (1-15a);sn (1-1la) Rose, J. N. et al, 1220 (1-6b);2449 (1-6b);
Mikan,J. G., 62 (1-18a);5983 (1-15c) 2450 (1-12); 2596 (1-22)
Miranda,R., 8 (l-15a) Ryan, J., 62 (1-l1a); 390 (1-6b);sn (1-15c)
Molinaet al, 119 (1-10); 131 (1-10); 1066 Sagot,H., sn (1-22)
(1-10); 3613 (1-10); 7494 (1-10); Sagot,P., 36 (1-6b); 1217 (1-18a);sn
12477 (1-10); 14206 (1-10); 22436 (1-6b)
(1-10) Sampaio,A., 6744 (1-15a);7163 (1-15a)
Mosen,H., sn (1-15a) Sandwith,N. Y., 563 (1-6b); 1592 (1-6b)
Nadeaud,J., sn (1-15a);sn (1-lla) Santoro,412 (1-15a);567 (1-15a)
Ngadiman,S., 34713 (3-1); 34965 (3-1); Sarawak,ForestDept., 22620 (3-1); 24909
36148 (3-1) (3-1)
Novaes,J. de C., 1108 (1-15a) Schenck,H. 1111 (1-15c); 1589 (1-lla);
Oliveira,E., 3608 (1-6b);4025 (1-6b) 3061 (1-1la);4250,(1-15a)
Oliveira,J. E., sn (1-15a) Schomburgk,Rob. et al, 56 (1-14);56
Srsted,3721 (1-10); 3722 (1-10); 3723 (1-22); 63 (1-14); 249 (1-14); 313
(1-10) (1-22); 373 (1-14); 951 (1-6b);
Pabst,G., 10252 (1-15a) 953 (1-6b);sn (1-22)
Peckolt, 91 (1-lla); 129 (1-11lla);526 Schott, H. W., 1677 (1-17); 1678 (1-15a);
(1-lla) 1679 (1-1la); 1680 (1-1la); 5977
Perdonnet,649 (1-17) (1-17); 5979 (1-17); 5980 (1-22);
Pereira,E. et al, 1246 (1-1la);4256 (1-17); 5982 (1-15c);5984 (1-lla)
4258 (1-15a);9835 (1-lla) Schiich,404 (1-15a);5985 (1-lla); sn
Perrottet,G. S., 259 (1-18a);260 (1-18a); (1-15a);sn (1-17); sn (1-18a);sn
261 (1-18a);262 (1-6b);sn (1-18a) (1-1la).
Persaud,A. C., 179 (1-6b) Schunke,J. M., 50 (1-24); 2299 (1-16)
Pickel,B., 133 (1-15a);sn (1-15a) Schwacke,C. A., 3330 (1-lla)
Pires,J. M. et al, 654 (1-7); 2481 (1-15c); Schwarz,G. J., 12224 (1-15a)
3314 (l-15a);4570 (1-18a);4734 Scoteschini,R. F., sn (3-1); 20142 (3-1)
(1-6b); 14331 (1-18a);52283 Selo, sn (1-15a);sn (1-15c); sn (1-1la);
(1-18a);52522 (1-18a) 660 (1-17)
Pohl, J. E., 6031 (1-15a);sn (l-15a); sn Sellow, F., sn (1-18a)
(1-1 a) Servicedes Aux, 4204 (1-22)
72 Flora Neotropica
Shotsky, 1832 (1-lla) Stork,H. E., 3334 (1-10)

Shultes,R. E., 24511 (1-7) Teixeira,1610 (1-15a)
Silva,M. et al, 20 (1-18a);94 (1-18a); Tessmann,G., 5230 (14)
200 (1-18a);439 (1-6b); 950 (1-7); Teysmann,440 (3-1);4411 (3-1);4548
2313 (1-6b) (3-1)
Silva,N. T., et al, 351 (14); 439 (1-6b); Tikun,S., 39342 (3-10)
1168 (1-6b); 1229 (1-6b); 3016 Tillett, S. S., et al, 45524 (1-8)
(1-18a);3050 (1-18a);3585 (1-15a) Tonduz,A., 13486a (1-10); 13846 (1-10)
Sinanggul,H. T., ForestDepartment Toro, R. A., 1155 (1-4)
Sandakan30548 (3-1) Traill,J. W. H., 36a (1-7)
Sinclair,1843 (1-10) Trintaet al, 511 (1-15a). 821 (1-15a);
Sing, 21020 (3-1) 994 (1-18a)
Smith,A. C., 2752 (1-6b) Troll,522 (1-19)
Smith,F. D., et al, 32 (1-15a);5795 Tutin,T. G., 172 (1-6b)
(1-15c); 15441 (1-2) Ule, E., 1029 (1-15c);6074 (1-7); 6140
Smith,H. H., 884 (1-10); 886 (1-10); (1-14); 9519 (1-15a);9520 (1-6b);
887 (1-10); 888 (1-10) 9640 (1-6b)
Snedaker,C-24 (1-11c); C-25 (1-1 c) Uribe,U. L., 1550 (1-24)
Sobrinho,J. S., 2040 (1-15b) Valerio,J., 619 (1-10)
Socpadmo,136 (3-1) Van Donselaar,2894 (1-6b)
Sonntag,29 (1-llc) Van Emden,W.C., sn (1-6b)
Soubirou,sn (1-6a) Vauthier,88 (1-lla); 139 (11-17); 162
Splitgerber,F. L., 1136 (1-22) (1-22); 168 (1-17); 552 (1-lla)
Spruce,R., 150 (1-7); 2416 (1-7); 3871 Ventenat,sn (1-15a)
(1-5);4944 (1-6b) Versteeg,G. M., 121 (1-22); 153 (1-22)
St. Hilaire,A., C-102 (1- la); 125 (1-18a); Voisin, sn (1-6b)
194 (1-lla);226 (1-15a) B1-385 Wachenheim,H., 479 (1-6b)
(1-lla); 1234 (1-15c); 1830 Webster,G. L. et al, 1240 (1-10); 12479
(1-15c); 2205 (1-15c); sn (1-1la) (1-10)
Stahel,G. et al, 631 (1-6b) Weddell,H. A., 202 (1-15a);484 (1-lla)
Standley,P. C. et al, 3068 (1-10); 3411 Wel-Lek,C., 621 (3-1)
(1-10); 11268 (1-10);20594 (1-10); Widgren,J. F., 726 (1-1a); 727 (1-15a);
21340 (1-10); 28073 (1-10); 40045 748 (1-15a);sn (1-15a)
(1-10);44908 (1-10);40949 (1-10); Williams,L. O. et al, 6684 (1-15a)
46678 (1-10); 62202 (1-10); 73760 7472(1-15a); 7478 (1-15a);11262
(1-10); 73850 (1-10); 74348 (1-10); 11879 (1-10); 13221 (1-10);
(1-10); 74402 (1-10); 75137 26630 (1-10)
(1-10); 75476 (1-10); 78584 (1-llc); Williams,LI., 2395 (14); 3357 (1-24)
78860 (1-10); 79279 (1-10); 79307 Williams,R. S., 252 (1-10)
(1-10); 87416 (1-1lc); 87755 Wood,G. H. S., 16258 (3-1)
(1-10); 88330 (1-10); 88718 (1-10) NWray, L. Jr., 179 (3-1); 3253 (3-1)
Steinbach,J., 8132 (1-19) Wullschlaegel, H. R., 8161 (1-22)
Stern,W.L. et al, 163 (1-10); 224 (1-10); Wurdack,J. J. et al, 39679 (1-6b);
953 (1-10) 43194 (1-7);43283 (1-7);43747
Steyermark,J. A. et al, 77 (1-8); 30199 (1-7)
(1-10); 30244 (1-10); 74649 (1-24);
87550 (1-6b);87616 (1-6b);88095
(l6b); 96588 (1-20)
Index 73
INDEX
Note: Pagenumbersin bold-faceindicate primarypage references;page

numbersin italics indicate synonyms;the * indicatespageswith illustrations.
Crotoneriosperma1, 45 varmicrocarpa12, 39
Euphronia1, 2, 11, 14, 20, 21, 24, 25 macrantha9, 11, 18, 19, 30, 31, 37
Herrania2 macrocarpa41, 57
Hoeffnagelia1, 29 membranacea9, 47
Humbertiodendron 3 micrantha46
Humbertodendron 1, 3, 4, 6, 12-14, 16, 18, microcarpa39
20, 29, 63 mollis55, 56
saboureaui4, 15*, 17*, 63 najadum62
Isopteris64 nivea9, 16, 18, 19, 31, 32, 50
penangiana2, 64 f paniculata50
Lightia2 varfasciculata9, 13, 51, 52
Mainea2, 29 varnivea9, 13, 51
racemosa45 varpubescens13, 51, 52
Malpighia1, 19 ovalifolia50, 52
Nuttallia1, 29 panamensis44
Paullinia16, 20 paniculata8, 9, 11, 19, 31, 47, 54, 55
Stephanopodium20 parviflora39, 40
Trigonia1-4, 6-14, 16, 18, 19, 25, 26, 28, prancei9, 10, 14, 16, 17*, 19, 30,
29-62 33, 35*, 62
sect Cincinnatae2 pubescens53
sect Cymosae2 racemosa46
sect Racemosae2 rasa4 7
bicolor39, 40 reticulata9, 12, 16, 19, 30, 41
boliviana9, 10, 19, 31, 57, 58 rigida44
bracteata9, 10, 19, 31, 41, 58, 59 rotundifolia9, 12, 16, 19, 30, 33, 34*
candelabra9, 10, 19, 30, 41, 42*, 43* rugosa6, 9, 10, 12, 16, 17*, 19, 30,
candida50, 52 44,45
cepo 55, 56 rytidocarpa9, 12, 19, 31, 59, 60
coppenamensis9, 12, 19, 29, 30, 32 schottiana54, 55
costanensis9, 11, 19, 31,48 sericea9, 11, 16, 19, 32, 36, 49, 62
crassiflora36, 37 simplex47, 60
crotonoides45, 60 varpilosula57
varelliptica46 spruceana3, 5, 9, 11, 14, 16, 19, 30,
varincana45 32,40
varoblongifolia46 subcymosa9, 10, 19, 31, 33, 49, 50
echiteifolia9, 11, 16, 19, 32, 54, 61 thyrsifera44
eriosperma14, 19, 30, 31, 45, 55, villosa1,9, 16, 17*, 19, 29, 31, 50,
58-60 55
subsperiosperma9, 10, 46, varcuneata55
48, 60 varmacrocarpa9, 11,41, 56,
subspmembranacea9, 10, 46, 57
47,48 varoblonga55
subspsimplex9, 10, 46, 47, varobtusata55
48, 58, 60 varvillosa9, 11, 56
euryphylla44, 45 virens9, 11, 16, 18, 19, 30, 31, 36,
fasciculata52 59,61
floccosa9, 10, 19, 30, 48 xanthopila60, 61
floribunda44, 45 Trigoniaceae28
glazioviana59, 60 tribeLigthiae28
hypoleuca9, 11, 16, 19, 32, 44, 60, tribeTrigoniastrae28
61 tribeTrigoniae28
varpubescens60 Trigoniastrum1-4, 6, 11-14, 16, 18, 29, 63
kaieteurensis38, 39 hypoleucum4, 64, 65
killipii9, 11, 19, 31,53 varoliganthum64
laevis1,9, 16, 30, 38, 47 varviride64
varlaevis 12, 39

Flora Neotrópica - Trigoniaceae

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Flora Neotrópica - Trigoniaceae

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Organization for Flora Neotropica

Organization for Flora Neotropica

Issued 28 April 1978

The New York Botanical Garden

Libraryof CongressCatalogCardNumber 77-91706

Lleras, Eduardo, 1945-

(Flora neotropica; monograph no. 19)

The presentstudy includes all of the known Trigoniaceae,both neotropical

In his Histoire des Plantes (1775), Aublet describedthe genus Trigoniain

Based in part on a dissertation submitted to the Graduate Faculty in Biology in partial

those who have undertakensystematictreatmentsof the Trigoniaceae.Vellozo, in

The Trigoniaceaeare found in the Neotropics and in the Paleotropicsof

due to severalcauses. First, gallery forests are usually botanically diverse,while

Dichapetalaceaeand Vochysiaceae, all presumablymigratedbetween Africa and

(1971), Haffer(1969) and Vanzolini(1970), using palynologicaland climatological

been able to observepollinationin the field, the label on one herbariumspecimen

differentelements cited here when the savannaswere reclaimedby the rainforests

FIG 3. Known distributionof species of Trigonia.0 T. rugosa; v T. eriospermasubsp

FIG4. Knowndistributionof speciesof Trigonia. 0 T. prancei; T. boliviana; T.floccosa;

FIG 5. Known distribution of species of Trigonia. N T. macrantha: 0 T. killipii; * T. virens;

The anatomy of the Trigoniaceaewas studied thoroughlyby Barth(1896),

FIG 6. Known distribution of species of Trigonia. * T. reticulata; [ T. coppenamensis:

In the presentstudy five randomlyselected specimensof each of the more

The terminologyand formatused here to describethe pollen of the Trigo-

EVOLUTIONOF CHARACTERSWITHINTHE FAMILY

Althoughit is difficult at presentto discerngeneralizedtrends of evolution

FIG 9. Pollen of Trigoniaceae.A-C,ScanningElectronMicrographs,X 436. D-F, with light

This section on Taxonomic Historypresentsthe taxonomic positions previ-

FIG 10. Schematicrepresentationof similaritiesand differenceswithin Trigonia.

Aublet (1775) Decandria Monogynia

Hallier(1921) with Dichapetalaceae& Chrysobalanaceae

As discussedin the section on TaxonomicHistory,Euphroniawas placedin

Trigoniaceae Dichapetalaceae Malpighiaceae Sapindace

position of leaves opposite or alternate alternate opposite (u) alternate,

type of leaves simple simple simple simple or

stipules + + + (-) + (R)

floral symmetry zygomorphic actino (-zygo) actinomorphic actino, zygo

arrangement of sepals imbricate imbricate imbricate imbricate or

number of petals 5 (papil.) 5 5 3-5

arrangement of petals convolute imbricate convolute imbricate

number of stamens (total) 5 - 13 5 10 Ca 8

filaments connate free or connate connate (u) free

number of locules on anthers 2 2 2 2

type of dehiscence of anther introrse introrse introrse introrse

Trigoniaceae Dichapetalaceae Malpighiaceae Sapindac

position of ovary superior superior- superior superior

number of locules 3(14) 2-3 3 3(14)

type of style simple simple divided simple or di

number of ovules per locule 1 - many 2 1 1-2 (many)

orientation of ovules epitropous epitropous epitropous epitropous

placentation on intruded placentas axile-pendulous ascending- axile or par

type of fruit capsules or samaras drupes samaras or various

embryo straight straight straight or plicate or tw

Trigoniaceae Euphronia Vochysiaceae

Pollen3-5 porate Pollentricolporate Pollentricolporate

Petals5 Petals3 Petals 1-5

Stamensall connatein one Stamensin 2 or 3 groups Stamensin 1-severalgroups

StaminodesO-several Staminode1 Staminodesseveral

Disc glandspresent Disc glandsabsent Disc glandsabsent

Ovarylackinga central Ovarywith a central Ovarywith a central

Placentationon innerends Placentationaxile Placentationaxile

Fibersnot libriform Fiberslibriform Fiberslibriform

Parenchymaapotracheal Parenchymaparatracheal Parenchymaparatracheal

Foliarbundlesimmediately Foliarbundlesextending Foliarbundlesextending