Table Of ContentFLORA NEOTROPICA MONOGRAPH 83
MOREAEA, RTOCARPEAAE,N DD ORSTENIA
(MORACEAE)
With Introductions to the Family and Ficus and with Additions
and Corrections to Flora Neotropica Monograph 7
CORNELICS. BERG
~~~~~~~~~~~TIOPIC c Of CAN1CEl
FLORAt
NEOTROPICA(
' ^
IOP'C
CAPRICORN
Publishedfor
The Organizationf or FloraN eotropica
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Issued 31 October 2001
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Library of Congress Cataloging-in-Publication Data
Flora neotropica.- Monograph no. 1 -New York:P ublished
for Organizationf or Flora Neotropica by The New York
Botanical Garden, 1968-
v.: ill.; 26 cm.
Irregular.
Each issue has distinctivet itle.
Separatelyc atalogueda nd classified in LC before monographn o. 40.
ISSN 0071-5794 = Flora neotropica.
1. Botany - LatinA merica- Classification- Collected works.
2. Botany - Tropics - Classification - Collected works. 3. Botany
Classificatio-n Collectedw orks.I . Organizatiofno rF loraN eotropica.
II. New York Botanical Garden.
QK205.F58 581 .98'012-dcl9 85-647083
Libraryo f Congress [8508]
ISBN 0-89327-439-9
1 2 3 4 5 6 7 8 9 / 09 08 07 06 05 04 03 02 01
1V
MOREAE,A RTOCARPEAEA, ND DORSTENIA
(MORACEAE)
With Introductionst o the Family and Ficus and with Additions
and Correctionst o Flora NeotropicaM onograph7
CORNELISC . BERG
CONTENTS
Abstract/Resumen/Resum.o.. ..........................................................1
Introductiont o the Family Moraceae. .............................. .............................. 4
TaxonomicH istory ............................................................. 4
Morphology. ............................................................4
Pollination ............................................................ 12
Dispersal ............................................................. 14
Distributiona nd Ecology ................. ........................................... 15
Classificationa nd Diversification. .............................. ............................. 20
Use ............................................................. 21
ConservationS tatus ............................................................ 22
TaxonomicT reatment. ........................................................... 22
Key to the Tribes of the Moraceaei n the Neotropics ............................ .................. 22
Key to the Neotropical Generao f the Moraceae. ..................................................... 22
Moreae. ........................................................... 24
Artocarpeae.. .......................................................... 67
Dorstenieae. ........................................................... 143
Dorstenieaep .p. ("Brosimeae"a) ndC astilleae( "Olmedieae")A, dditionsa nd
Correctionst o FloraN eotropica Monograph7 ......................................................... 231
Ficeae ............................................................ 289
Acknowledgments.. .......................................................... 292
LiteratureC ited ............................................................. 292
Maps ............................................................ 300
NumericalL ist of Taxa ............................................................ 313
List of Exsiccatae ............................................................ 315
Index of Local Names ............................................................ 340
Index of Scientific Names ............................................................ 342
ABSTRACT
Berg, Cornelis C. (BotanicalI nstitute,U niversityo f Bergen/TheN orwegianA rboretum,
N-5259 Hjellestad,N orway). Moreae,A rtocarpeae,a ndD orstenia (Moraceae);w ith intro-
ductions to the family and Ficus and with additions and corrections to Flora Neotropica
Monograph7 . FloraNeotropicaM onograph8 3: iv+l-348. 2001.-The presentm onograph
includes the revisions of generao f the tribesM oreaea ndArtocarpeae,t he genus Dorstenia,
correctionsa nd additions to genera treatedi n Fl. Neotrop. Monogr. 7, "Brosimeae,"p res-
2 FLORAN EOTROPICA
entlyi ncludedin thet ribeD orstenieaea,n d" Olmedieaep,"r esentlyC astilleaeT. heya rep re-
cededb y ani ntroductiotno thef amilyM oraceaaen ds upplementewdi thb riefi ntroductions
to thet ribeF iceaea ndt heg enusF icus.T hee ntitietsr eatefdo rt hef irstt imef orF loraN eotropica
comprisen ine genera:B agassa (1 sp.), Batocarpus (3 spp.),C larisia (3 spp.),D orstenia
(46 spp.),M aclura( 2 spp.),M orus( 2 spp.),P oulsenia (1 sp.), Sorocea (14 spp.),a ndT rophis
(5 spp.)- 77 speciesi n total.E ightn ew speciesa ndt won ew subspeciesw ered iscovered
fort heseg enerad uringth ep reparatioonf t hem onographas, w ella s a secondy et-undescribed
speciesf orP oulsenia. Thisg roupo f generac omprisems anyw idespreada nd/orc ommon
speciesf orw hichn umeroucso llectionsh aveb eene xamineda ndl isted.H oweveri,n Dor-
stenia, manys peciesa rev eryl imitedi n theird istributioann dt hen umbero f specimense x-
aminedis oftenl imitedt o a few ors ometimeosn lyo ne.T heg roupo f then iner evisedg enera
is in manyr espectsh ighlyd iverse.F ort he groupo f generat reatedin Monograph7 the
additionsa ndc orrectionisn clude:r evisedo r extendedd escriptionosf severals peciesa nd
subspeciesc,h angedd escriptiondsu et o fusiono f speciesp reviouslyre cognizeda s distinct,
indicationasn dd ocumentatiofonr r angee xtensionss,o men ewc ombinationfsu, llt reatments
of taxad iscoveredsi ncet hep ublicatioonf Monograp7h ( 12 specieso rs ubspecies)s,u pple-
mentaryc ommentos n generaa nds pecies,a ndp artlyo r entirelyr evisedk eyst o speciesf or
severagl eneraF. ort hiss econdg roup7 0 species( int hen ineg eneraa) rer ecognizedat p resent.
Thei ntroductiotno thef amilyi ncludesc hapterds ealingw itht het axonomich istorym, or-
phologyp, ollinationa ndd ispersald, istributioann de cology,c lassificationd, iversification,
use,a ndc onservationT.h eg enusF icus (awaitingre visioni)s to somee xtenti ncludedin the
introductionin, particulairn thec hapteros rp aragraphdse alingw ithh abit,p ollinationa, nd
distributionm; oreovera, descriptioonf anda briefa ccounot f thisg enusa rep rovidedI.n t he
presenct ontributionth, en ewc ombinationCsl arisia Ruiz& Pavons ect.A canthinophyllum
(Allemao)C . C. Berg,P seudolmedia Treculs ect.O lmediopsis (KarstenC) . C. Berg,a nd
Naucleopsis ulei subsp.a mara (Ducke)C . C. Berga rem ade,a ndt hen ewt axonN aucleop-
sis ulei (WarburgD) uckes ubsp.s ubandina C. C. Bergi s described.
RESUMEN
Berg, CornelisC . (BotanicaIl nstituteU, niversityo f Bergen/ThNe orwegianA rbore-
tum,N -5259H jellestadN, orway)M. oreaeA, rtocarpeaea,n dD orstenia (Moraceae)w; ith
introductiontos thef amilya ndF icusa ndw itha dditionas ndc orrectiontso FloraN eotropica
Monograp7h. FloraN eotropicMa onograp8h3 :i v+1-348.2 001.- Lap resentme onografia
incluyel asr evisioneds e los generosd e last ribusM oreaey Artocarpeaeel, g eneroD orstenia,
correccioneys a dicioneas losg enerops resentadoesn l aF l.N eotropM. onogr7. , "Brosimeae,"
actualmenitnec luidaen l at ribuD orstenieaye " Olmedieaea,"ct ualmenCtea stilleaep,r ecedidas
poru nai ntroduccioan l a familiaM oraceaye complementadcaos nb revesi ntroduccioneas
la tribuF iceaey al generoF icus. Los grupost ratadops orp rimerav ez en FloraN eotropica
comprendennu eveg enerosB: agassa (1 spp.),B atocarpus (3 spp.),C larisia (3 spp.),D or-
stenia (46 spp.),M aclura (2 spp.),M orus (2 spp.),P oulsenia (1 spp.),S orocea (14 spp.)y
Trophis( 5 spp.)- 77 especiese n total.O chon uevase speciesy dos nuevass ubespecies
fuerond escubiertapsa rae stosg enerosd urantela preparaci6dne la monografiaa, si como
tambienu nas egundaes peciea, umn od escritad eP oulsenia.E steg rupod eg 6nerocs omprende
muchase speciesa mpliamendtei stribuidays/ o comunesd e lasc ualess e hane xaminadyo se
citanm uchacs oleccionesS. ine mbargoe,n D orstenia,m uchaess peciesti enenu nad istribucion
muy limitaday el nu'merdoe muestraes xaminadasse limitaa menudoa pocasy algunas
veces solamentea una.E l grupod e los nueveg enerosr evisadose s, en muchosa spectos,
extremadamendteiv erso.P arae l grupod e generosp resentadoesn la Monografia7 , las
adicioneys correccioneisn cluyend: escripcionerse visadaos aumentadadse variase species
y subespeciesc,a mbiose n lasd escripcionedse bidoa fusi6nd e especiesp reviamentter atadas
como diferentes,i ndicacionesy documentacionp arae xtensionese n los rangos,a lgunasc om-
RESUMO 3
binacionens uevast, ratamientocos mpletods et axad escubiertolus egod e la publicaciodne l a
Monografi7a (docee specieso subespecies)o,b servacionecso mplementariaacs ercad e los
generosy las especies,s e revisanp arciaol totalmentlea s claves,p aral as especiesy varios
g6nerosP. arae stes egundog rupos e reconocenen l a actualida7d0 especiese nn ueveg eneros.
Lai ntroduccioan l a familiai ncluyec apitulosa cercad e la historiata xon6micam, orfologia,
polinizaciony dispersi6nd, istribuciony ecologia,c lasificaci6nd, iversificaci6nu, sos y
conservacionE. l generoF icus (auinp or revisar)e sta incluidoh astac iertop untoe n la
introduccione,n particulaern capituloso parrafoqs ues e refierena l habito,p olinizaci6ny
distribuci6na;d emass, e proveeu nad escripci6ny unap equenirae laciond e esteg 6neroE. n
la presentec ontribuci6ns,e hacenl as siguientesc ombinacionensu evas:C larisiaR uiz&
Pav6n sect. Acanthinophyllum(A llemao) C. C. Berg, PseudolmediaT r6culs ect. Olmediop-
sis (KarstenC) . C. Berg,a ndN aucleopsisu lei (WarburgD) uckes ubsp.a mara( Ducke)
C. C. Bergy se describee l nuevot axonN aucleopsisu lei (WarburgD) uckes ubsp.s ub-
andinaC . C. Berg.
RESUMO
Berg, CornelisC . (BotanicaIl nstituteU, niversityo f Bergen/ThNe orwegianA rbore-
tum,N -5259H jellestadN, orway)M. oreaeA, rtocarpeaea,n dD orstenia( Moraceae)w; ith
introductiontos thef amilya ndF icusa ndw itha dditionas ndc orrectiontso FloraN eotropica
Monograp7h. FloraNeotropicMa onograp8h3 :i v+1-348.2 001.- Apresentemonografia
incluir evis6esd osg enerosd ast ribosM oreaee Artocarpeadeo, generoD orsteniac, orrecoes
e acrescimops arao s generost ratadons a Fl. NeotropM. onogr7. , "Brosimeaea"t ualmente
incluidan a triboD orstenieaee "Olmedieae,a"t ualmentCe astilleaeS. aot odasp recedidas
poru mai ntroducaop araa familiaM oraceaee complementadacso m brevei ntroducoes
paraa triboF iceaee parao generoF icus.A s entidadatsr atadaps elap rimeirav ez paraa
FloraN eotropicaa brangemn ove generosB: agassa( 1 sp.),B atocarpus(3 spp.),C larisia
(3 spp.), Dorstenia (46 spp.), Maclura (2 spp.), Morus (2 spp.), Poulsenia (1 sp.), Sorocea
(14 spp.)e Trophi(s5 spp.)- num totald e 77 especiesD. urantae preparaaod amonografia
foramd escobertops arae stesg eneroso iton ovase speciese duasn ovass ubespeciesa, ssim
comou mas egundae speciep araP oulseniaa, indan aod escritaE. steg rupod e generosc om-
preendem uitase species comunse /ou amplamented istribuidasp, ara as quais foram
examinadaes l istadans umerosacso le96esT. odaviae,m D orsteniam, uitase speciesp ossuem
distribuicaloi mitadae o nuimerdoe amostraes xaminadafso i reduzidoo u somenteu m.0
grupod os nove generosr evisadose , sob multiplosa spectosa, ltamentde iversificadoO. s
acrescimoes corre96eps arao grupod os generostr atadons a Monografi7a incluemd: escri-
qoesr evisadaso u ampliadasd e variase speciese subespeciesd, escric6esa lteradaps or
causad af usaod e especiesp reviamentree conhecidacso mod istintasi,n dicac6ees documen-
ta9aop araa extensaod e ocorenciaa, lgumasn ovasc ombina96estr, atamentcoo mpletod os
taxa descobertosd esde a publica9aod a Monografia7 (doze especies e subespecies),
comentirioss uplementarepsa rao s generose especiese chavesp araa s especiesd e varios
generosp arcialo u inteiramentree visados.P arae ste segundog rupos ao reconhecidas
atualment7e 0 especies( nos nove generos)A. introdu9apo araa familiai ncluic apitulos
coma bordagemda h istoriat axonormicma,o rfologiap, oliniza9aoe dispersaod, istribuicao
geograficae ecologia,c lassifica9aod, iversifica9aou,s o e conserva9ao0. generoF icus
(aguardandroe visao)e abordadoco m algumae xtensaon a introdu9aoe,m particulanr os
capituloso u paragrafoqs uet ratamso breo hibito,p olinizago e distribui9aoa;l emd isso,
saoa crescentadausm ad escrigioe umas intesed esseg eneroN. a presentec ontribui9asoa o
feitasa s novasc ombinaq6eCs larisias ect.A canthinophyllu(Aml lemaoC) . C. Berg,P seud-
olmediaT reculs ect. Olmediopsi(sK arstenC) . C. Berge Naucleopsis( WarburgD) ucke
subsp.a mara( Ducke)C . C. Berg,e e descritoo novo tdxonN aucleopsisu lei subsp.s ub-
andinaC . C. Berg.
4 FLORAN EOTROPICA
INTRODUCTION TO tinct fromb oth Moraceae( s.str.)a ndU rticaceae( s.str.),
Berg (1978a) proposedt o treatt he group of six genera
THE FAMILY MORACEAE
as a separatef amily,C ecropiaceaea, decision generally
The presentc ontributiont o FloraN eotropicac om-
accepted,s uch as by Cronquist( 1981), and confirmed
prises sections with generat reatedf or the firstt ime and
by Setoguchi et al. (1993) in a study on silicon-accu-
sections with corrections and additions of genera re-
mulating idioblasts in leaves of Cecropiaceae, which
vised in FloraN eotropica Monograph7 . This implies
also supports the position of Poikilospermum as a
thata ll neotropicalM oraceaee xcept for the representa-
membero f the family.F or the reducedM oraceae,C or-
tives of Ficus are included. However,i n ordert o make
ner (1962) proposeda subdivisioni nto six tribes:A rto-
the general survey of the family more complete, the
carpeae,B rosimeae,D orstenieae,F iceae, Moreae,a nd
genus Ficus is includedt o some extenti n the introduc-
Olmedieae.M oreover,h e reducedt he numbero f gen-
tion to the family.M oreover,a brief introductiont o this
era.T he conceptso f the classificationa nd delimitation
genus is provided.
of genera proposed by Corner were largely based on
knowledge of the Asian andA ustralasianm emberso f
TAXONOMIHCI STORY the family.C lassificationa ndd elimitationo f generah ave
been elaboratedb y Berg (1973, 1977a, 1977b, 1986,
In most classifications of the 19th century,s uch as
1988, 1989b) by including accumulatedk nowledge of
those of Baillon (1875a) and Bentham and Hooker
neotropical and African members of the family. The
(1880), the Moraceae were included in an "order"o r
results of these studies are discussed undert he tribes.
"family"w ith the subdivisions: Ulmaceae, Celtideae,
Accordingt o presentc oncepts,t he family Moraceae
Cannabineae,M oreae, Artocarpeae,C onocephaleae,
andU rticaceae.H owever,i n De Candolle'sP rodromus, comprises 37 genera and 1050-1100 species world-
volumes 16 (De Candolle, 1869; Weddell, 1869) and wide, with 19 generaa nd approximately2 70 species in
17 (Bureau, 1873; Planchon, 1873), the group had al- the Neotropics (Berg, 1998b).
readyb een split into five related" orders"(C annabineae,
Urticaceae,U lmaceae, Moraceae,a ndA rtocarpeae),a MORPHOLOGY
prelude to the subdivision of the order Urticales by
The Moraceae are highly diverse in the morphol-
Engler( 1889), in whicht he family Moraceaec omprises
ogy of bothv egetativea ndr eproductives tructuresT. he
the subfamilies Moroideae (with the tribes Fatoueae,
descriptiono f the diversity is largely focussed on the
Moreae,B roussonetieae,S trebleae,a nd Dorstenieae),
taxa treatedi n the presentc ontribution,b ut it includes
Artocarpoideae (with the tribes Euartocarpeae,
to some extenta lso the genus Ficus and Old Worldr ep-
Olmedieae,B rosimeae,a ndF iceae),C onocephaloideae,
resentativeso f the family.
andC annaboideaeT. he Cannaboideaae rem orer ecently
regarded as meriting the rank of family in the order
Urticales( cf. Cronquist,1 981), but the affinitiest o the Habit
otherf amilies of the Urticalesa re not quite clear;a po- In contrastt o its sisterf amily Urticaceae,t he Mora-
sitionc loser to the Myricaceaec ould be considereda nd ceae areb asically woody. However,n early 10%o f the
investigated. species are herbaceous. Most of the herbaceous spe-
In arevisionals tudy,C hew( 1963) reducedt heA sian cies belong to the speciose genus Dorstenia and only
genus ConocephalusB lume, the type of the subfamily two of them belong to the small paleotropical genus
Conocephaloideae,t o a subgenus of Poikilospermum Fatoua Gaudichaud.T he herbaceousm embers of the
Miquel, a genus which was accepted as a member of genus Dorstenia show a wide range of habits and life
the Urticaceae, e.g., by Engler (1889). This made the forms, as discussed in detail in the introductiont o this
weakness of the demarcationo f Moraceaea nd Urtica- genus in the presentc ontribution.
ceae apparent.C hew proposed to transfert he micro- About 30% of the species are basically hemiepi-
spermousg enera of the Conocephaloideae, Cecropia phytic and they belong to the genus Ficus. The ability
Loefling, CoussapoaA ublet, Musanga P. Brown, and to produce( aerial) adventitiousr oots is not only com-
Poikilospennum (s.l.), to the Urticaceae, leaving the mon in Ficus (hemiepiphytesa nds ome groupso f climb-
macrospermousg enera, MyrianthusP . Beauvois and ers,i ncludingr oot-climbersb) ut it is also foundi n other
PouroumaA ublet,i n the MoraceaeC. omer( 1962) made woody representativeso f the order:i t is characteristic
a morer adicald ecisionb y transferrintgh e Conocephal- for the Cecropiaceae and occurs in the (Old World)
oideae (including Poikilospermum)t o the Urticaceae, woody climberso f Urera Gaudichaud(U rticaceae)T. he
which made the Moraceaem ore homogeneousb ut the remaindero f the species (approx. 60%) are terrestrial
Urticaceae less so. As the Conocephaloideae are dis- trees, shrubs, climbers, or (a few) subshrubs.
INTRODUCTIONT O THE FAMILYM ORACEAE 5
Mosto f the specieso f thef irstc ategorya res mall the superstructuroef the fig tree andt he supporting
(= understoryt)o medium-sized(= canopy)t rees,b ut cylindero f rootsa rei n lineo r if thef ig treem anagetso
somec ana ttaina considerablhe eight( to 60 m), such sendd owna eriarl ootsa longa djacenttr eesa ndt here-
as specieso f Brosimum( subg. Ferolia), somes pecies withp roducesa broadc arryingb ase.O nlya few spe-
of Ficus sect.P harinacosycea,a ndM aquirac oriacea. cies arep owerfuel nought o replaceh ostt reesi n high
In thisg roupa numbero f architecturmal odelsa s de- forest,s uch as in the Neotropicss pecies like Ficus
scribedb y Hallea ndO ldeman(1 970)a ndH allee t al. gomelleira Kunth& Bouche, F nymphaeifoliaM iller,
(1978)a rer epresentedsu, cha st hem odelso f "Corner" andF schultesiDi ugandH. emiepiphytfiicg speciesa re
(discussedb elow), "Cook,"a nd" Roux."I t is note- alsoo ftenf oundo n rockys urfaces(a s hemiepilithes).
worthyth ats omet allo rm edium-sizetdr ees peciesc an Howevers, everals pecies,s ucha s the ? lianescenFt
also be shrubs,o r at least startf loweringa s shrubs. schippiiS tandleya, reo ftent errestriaTl.h es horts tem
Thef acultativfer utescenhta bitm ayb e foundi n parto f and the broad crown, general features of the
thes peciesr angea, si nH elicostylist omentosaM, aquira hemiepiphytFici cuss peciesa, llowd evelopmenats t er-
guianensis (subsp.c ostaricana), andt he OldW orld restriatlr eeso nlyi n openh abitats.
Antiaris toxicaria Leschenaulto,r almostt hroughout Ther epresentativoesf theo thern eotropicaslu bdi-
thes peciesr ange,a s in Perebea angustifolia. visiono f Ficus,a bout2 0 specieso f subg.P harmaco-
Numeroups aleotropicaFli cus speciesa res hrubs, syceas ect.P hamacosyceaa, re( normallyt)e rrestrial,
someo f themr heophyteAs. bout2 0%o f then on-Ficus oftene lementso f riverinea ndm ontanef orest,e stab-
woodys peciesa re( orc anb e) shrubs( or subshrubs). lishingi n ? disturbepdl aces,a traitt hatc anb e related
About 100 species are (or can be) climbers,t he to then eedf orl ightf org erminatioonf thes malls eeds.
majorityb eingr oot-climberosf Ficus sect.K alosyce
(MiquelC) ornear nds ect.R hizocladusE ndliche(rC or- Leaves
ner,1 965),t heo therso f thes tragglintgy pe,a s inM ac- Thel eavesa reu suallya ltematea ndt hena rranged
lura andF icus subsectP. alaeomorphe( King)C orner. in spiralso rd istichousT. hel eavesm ayb e arrangeidn
Int heg enusM aclura, thornsf acilitatec limbing. spiralso n the the maina xes but in two rows on the
laterabl ranchesa,s i n thes peciese xhibitingth em odels
Hemiepiphytism in Ficus of "Roux"a nd" Cook"( Hallee t al., 1978).I n some
Subgenus Urostigma (with about 280 species specieso f Dorsteniath el eavesa rer osulatde uet o short-
worldwidei)s characterizebdy theh emiepiphytilcif e eningo f the stem.O ppositele avesa reu ncommonin
form( also foundi n a subdivisiono f the Old World MoraceaeT. heya ref oundi n Bagassa,i n the paleo-
"subgenusS"y cidium).T hisl ife formi mpliest hatt he tropicagl enusB roussonetiVa entenaat,n di n a number
plantss tarta s epiphytesw, hichu suallys enda leading of paleotropicaFli cuss pecies.I n somep aleotropical
aeriarl ootd owna longt heb rancheasn d/orth et runko f Ficuss pecies,t hel eavesm ayb e (sub)verticillate.
theh ostt ree.W hent hisr ooth asr eachedth e soil and Thel aminais mostlyb asallya ttachedb,u ti t is peltate
canp riovidme orea mplen utrientst,h ep lantc anb egin in someD orstenias pecieso rr arely( as ana berration)
morer apidg rowtha nds oonero rl aterr eacht her epro- in someo thers pecies,s ucha s Castillae lastica.
ductivep haseT. hesem orer apidlyg rowingp lantss end Thel aminais simplea ndm ostlye ntireI. t is some-
downa dditionaale riarl ootsw hichc ana nastomosaen d timesm oreo rl essd eeplyin cisedp, innateloyr p almately
mayf orma networko f rootsa nde ventuallya cylinder (orp edately)e,i thero nlyi nj uveniles tageso f develop-
of rootsa roundth et runko f theh ostt ree.A t thats tage mento r alsow hena dult.T hei ncisionsc anr und own
thef ig treem ayo verpowe(ro rk ill)t heh ostt ree.S uch to the petiolea ndt he laminas egmentsc an even be
fig speciesa reo ftenr eferredto as stranglerosr stran- pseudopetiolulate( in Dorstenia bonijesu).
glingf igs, suggestingth atk illingo f theh ostt reeh ap- The venation is mostly pinnate, sometimes
pensb y strangulationc,u ttingo ff the sap streamso f (sub)palmate(,s ub)trinervateo,r in peltatel aminas
theh ostt reeb y preventingth eh ostt ree'st runkf rom (sub)radiatTe.h ev enationis basicallyb rochidodrom-
expandingI.t is morel ikelyt hati n mostc asesf ig trees ous,a lthougthh es ubmarginlaolo psc onnectintgh es ec-
overpowehr ostt reesb y starvationb,y a moree fficient ondaryv einsa res ometimefsa int.I n somes pecies( of
uptakeo f nutrientasn dw atera, ndo vershadowinhgo st Dorstenia)t,h el ateravl einsm ayr uns traighitn tot he
treesb y theirb roadc rownsK. illingo f theh ostt reei s margin(o ftene ndingin ah ydathode-likster uctureT).h e
oftena "suicidal"p rocess,a s the deadh ostt reem ay tertiaryv enationv ariesf roms calariform(t het ertiary
taket he fig treed owni n its fall. Thisn eedn otb e the veinsr unningtr ansversaen dp arallebl etweent hes ec-
case if the fig trees tartedit s life close to the soil, as ondaryv einsa t ? regulard istancest)o reticulateo, ri n
oftenh appenisn riverineh abitatos rs avannasI.n h igh severalF icuss peciesl argelyp arallewl itht he (main)
forest,f ig treesc an( occasionallyr)e mains tandingif secondarvye ins.
6 FLORA NEOTROPICA
Thel eafm arginis usuallye ntirei n sclerophyllous forc ertainfu ngia ndm ights ubstitutfeo rp ollena s at-
leavesa ndo ftend entates,e rratoer c renatien m esophyl- tractan(st eed iscussionu nderP ollinationF).u ngig row-
louso nes.I nt axaw ithe ntirel eafm arginst,h em argin ing on densep luricellulatrr ichomeas rea lsof oundi n
canb e dentatein j uveniles pecimensa, ndt hel aminais Pourouma(B erg,1 990b;B erge t al., 1990).
theno ftent hinner.
Thep etioleis normallsyh oritn leavesw ithe llipticto
Glandular Structures
lanceolatie coriaceoulsa minasw itha ne ntirem argin.
Ina dditionto the" glandulatrr"i chomeso,t hers truc-
Iti s oftenr elativelylo ngi n mesophylloulse avesa ndi n
turesh aveb eeno r canb e indicateda s "glandularI."n
thosew itha cordiformto suborbiculalarm inas, ucha s
(almost?a)l lF icuss pecieso nec anf indg landulawr axy
in Bagassa and species of Dorstenia and Ficus.
(ors hining)s potso n the lamina( andi n paleotropical
Scalel eavesa rep resenta s prophyllsa, s bractso n
taxaa lsoo n nodeso f leafyt wigs).O nt hel aminat hey
peduncless, ubtendinign florescenceosr a mongf low-
occuro n thel owers urfacea, tt heb aseo f them idrib(i n
ers,a st hep roximalle aveso n stemsa ndb ranchese,. g.,
theh emiepiphytimc onoeciousg roups,i ncludingt he
as scaleso f laterabl uds,s ucha s in Morus.I n several
neotropical subg. Urostigma sect. Americana), only
Ficus species,t he prophyllsf ormc alyptratec overs
in the axils of the basall ateralv eins (sucha s in the
enclosingt hey oungi nflorescences.
neotropical subg. Pharmacosycea sect. Pharma-
cosycea)a, nd/or(a lso)i n thea xilso f otherl ateravl eins
Stipules andm ajorf urcationos f the venation( in variousO ld
Thes tipulesa ref ullya mplexicauiln manyg enera Worlds ubdivisions)A. s they are (almost?)a lways
(ors ubgeneraI)n. o therg enerath eyv aryf romn otf ully presenitn onep laceo ra nothert,h eya rel ikelyt o have
amplexicautlo lateralT. hep airso f stipulesa ref reeo r somef unctiona, s suggestedb y Coiner( pers.c omm.)
fused.T heyv aryi n lengthf romm oret han 10 cm in in emittings ubstancetso attracpt ollinatorsT. hep osi-
someF icus speciest o less than 1 mm, as in several tiono f theg landsm ayh aved iagnosticv alue,a s it has
Dorstenias pecies.T heyc anb e (sub)persistenat,s in fort het won eotropicaslu bdivisionos f Ficus.
most Dorstenia species. Anothetry peo f "glandularst"r ucturies represented
Featureosf thes tipuless, ucha sp ersistencefu, sion, by smalleor rl arger" pocketsa"c cumulatinyge llowd ye,
dimensionss, hape,a ndi ndumentuma,r eo ftend iag- occurringi n Maclura, Batocarpus, and Clarisia, em-
nosticallyim portant. beddedin tepalsb, ractsa, nd/orc onnectiveos f anthers.
Int heseg eneray, ellow-coloresdu bstancecsa na lsob e
founde lsewherein the plant,a s in the barko f stems
Indumentum andr oots.T hey ellow substancem ayf unctiona s de-
Threet ypeso f trichomecsa nb e recognized:
terrenat gainsht erbivor(ys eed iscussionin theg eneric
1. Unicellulahr airso f variousk indsw ithr egardto
descriptionof Maclura).
length,f irmnessa, ndc olor.T heseh airsc an be unci-
nate.P resenceo r absenceo f uncinateh airsc anb e di-
Flowers
agnosticalliym portanatt v ariousle vels,r angingfr om
tribet o speciesU. ncinathe airsc anb ef oundo n allh airy The flowersa re smallt o very small andu sually
partso f thep lanti,n particulaitrs v egetativpe artsT. hey unisexualM. orphologicallbyi sexualb utf unctionally
tendt o be morea bundanitn j uveniles tagest hani n the unisexuafll owerso ccuri n a paleotropicaslu bdivision
adulto nes. of Ficus,a ndb isexuafl lowersm ayd evelopo ccasion-
ally (andt eratologicallyi)n taxan ormallyw ithu ni-
2. Pluricellularh airs,e longate,m oniliforma, nd
sexualf lowers.
oftenb rown(ish)S. uchh airsc anb e foundi n several
Thef lowersa rep edicellatoer s essile.P istillatfel ow-
Ficus species and in Poulsenia.T hey occurp romi-
ersa reo ftenc onnatea nd/ora dnateto ther eceptacloe f
nentlyi n someg enerao f theC ecropiacea(eC ecropia
thei nflorescence.
andP ourouma).
3. Pluricellulagrl obose-t o ellipsoid-capitatter i-
chomes( "glandulahra irs")T. hep resenceo r absence Perianth
andt he shapeo f theset richomehs aved iagnostics ig- Thep erianthis tepaloidI. n staminatfel owersi t is
nificancein manyg enerat;h es hapee .g. to distinguish often membranaceouasn d in pistillateo nes often i
thes ectionso f Perebeaa ndP seudolmediaan dt hep res- fleshy,m orep ronouncedlsyo in thef ruitings tate.
encet o distinguishth et wos ubgeneroaf Ficus.I ns ev- Them ost commonn umbero f tepalsi s four,l ess
eralS oroceat hese" glandulahra irs"a red enseo n the commonlfyi ve,t hreeo rt wo,a ndt heni n staminatfelo w-
perianthof pistillatefl owers.T heyp rovidea substrate ers commonlyw ith an equaln umbero f antitepalous
INTRODUCTIONT O THE FAMILYM ORACEAE 7
stamensT. etramerouasn d5 -meroufsl owersa rec om- Increaseo f the numbero f tepalsi s foundi n some
mont hroughoutth eo rderU rticalesS. mallern umbers generas, ucha sF icusa ndN aucleopsisI.t m ayo ccuri n
of tepalsa rec ommoni;n creaseo f numberiss lessc om- boths tamninaatned p istillatfel owerss, ucha si nNaucle-
mon( e.g., foundi n Naucleopsis). opsis, in whichg enusi t also leadst o the loss of the
Reductioonf thep eriantihn numbearn ds izeo f tepals normaal ndc learp ositiono f thet epals,i n particulairn
mayl eadt o vestigiapl erianthosr n akedf lowersw, hich thep istillatein florescenceTs.h ef actt hati nN aucleop-
occuri n severalg enera.I n the genusB rosimumth e sis thes taminatfelo wersa ree nclosedb y thep eripheral
wholer angef romw ell-develope4d- merousst aminate involucrablr actus ntila nthesicsa nb er elatedto a looser
flowerst hroughre ductionin numbear nds izeo ftepals arrangemenotf floralp arts.I n this genust hen umber
to nakedf lowers( witha single stamen)i s found.I n of stamenms ayb er educedw, hereatsh et epaln umbers
thisg enust hei ndividuafllo wersc ans tillb er ecognized. havei ncreased.
In other genera, such as Castilla and Pseudolmedia, Thet epalsc anb e freeo rp artlyt o entirelyc onnate.
the reductiono f the periantho f staminatefl owersi s In 4-merousfl owerst het epalsc anb e decussatew ith
linkedt o a processo f loss of then ormafl ixedr elative twoi nnera ndt woo utert epals.T hep airsa reo ftend if-
positiono f the floralp arts,r esultingin scatteredst a- ferenitn shapee, .g.,i nHelicostyliasn dS oroceaI.n t hese
mens on the receptaclea mongb ract-likes tructures,
generat he pairs of stamensm ay be more or less
apparentllyo oset epals,o rw ithouta nyr emnanot f the
anisomorphiocr, o nep aire vens taminodial.
perianth(i n theA fricang enus TrilepisiumTh ouars).
Thet epalso rt hef reep artso f thet epalsc anb e im-
Disorganizatioonf then ormafl lorals tructuries also,
bricateo r less often( sucha s in Trophisv) alvate.I m-
but less pronouncedlyf,o undi n staminatein flores-
bricatete pals( of flowersn ote nclosedb y bractsa nd/or
cences of Batocarpus and Clarisia.
ther eceptacloef thei nflorescencue)s uallyh avec iliolate
Inc asesi n whicht hep erianthlo sesi ts functiont o
marginsT. hem arginso f valvatet epalsa rep rovided
protecst tamenbs eforea nthesisp, rotectioins provided
withc urledh airst, hoseo f oppositem argines ntangled
otherwises,u cha sb yt hes hapeo f ther eceptaclde,e eply
and closingt he slits in between.I n Helicostylis,i n
cup-shapetdo (sub)urceolaoter ( inC astillab) ivalvate.
whicht het epalso f thep istillatef lowersa reo nly ba-
Bracts are often importanti n this respect, either
sallyc onnateo r entirelyfr ee( inc ontrastto mosto ther
interflorablr actso, ftenp eltate( sucha s in Brosimum),
genera)s, tripso f densec urledh airsa ref oundo n the
orbractas tt hep eripheroyf ther eceptaclea,s ( marginal)
inners urfaceo f thei nnert epalsa ndc loseo ff thei nte-
involucrabl racts( sucha s in the tribeC astilleae)o, r
rioro f thef lowerb, arrinagc cesst o theo vary( seeB erg,
ostiolarb ractsi n Ficus.T heo ccurrencoe f staminate
1972:8 0, 84, and9 1). Thed ifferencebs etweeni ndu-
flowersi n pits in the receptaclein some specieso f
mentumon imbricataen dv alvatem arginiss alsof ound
Brosimum(s ubg.F erolia)m aya lsob e relatedto pro-
for bracts( cf. Berg,1 990b).
tectiono f thes tamens.
Tepalso f staminatfelo werso ftenh avea ? distinctly
The staminatefl owerso f the generaH elicostylis,
cucullatea pex.T hisf eaturea ndt het wot ypeso f indu-
Maquiraa, ndP erebeao f thet ribeC astilleaeh, avw ell-
mentumo n them argincs anb e relatedto protectionof
developeda ndn ormallys tructurepde rianthsa, ndt he
stamenas gainsdt amageb y insectso ri nsectl arvae.
staminatien florescenceas reo penb eforea nthesisI. n
theo therg enerao f thet ribet hep erianthosf thes tami-
natef lowersa rem oreo r less disorganizedan d/orr e- Stamens
duceda ndt he staminatefl owersa ree nclosedb y the Thes tamenas reb asicallya ntitepalouasn dt hust heir
receptaclea nd/ort her elativelyl argea ndb roadi nner numberis equalt o thato f the tepals,o ftenf our,l ess
involucrabl ractsb eforea nthesisT. hei nneri nvolucral commonlyfi veo rt hreeT. hen umbeor f stamencsa nb e
bractos f staminatien florescenceosf specieso f Perebea reducedto one.T her eductionin numbemr ayc oincide
sect.N oyeraa rer elativelylo ng,n arrowin, curveda,n d witht her eductioonf thep erianthb,u ti t maya lsoo ccur
covert hes taminatfel owersw ithw ell-developepde ri- independentlsyu, cha s in Bagassa,o ri t mayc o-occur
anthsb, utt heym ayn oth avet hep rotectivfeu nctionth e witha n increaseo f tepals,s ucha s in Naucleopsis.
broadeneidn volucrabl ractsa pparentlhya ve. Twot ypeso f stamensc anb e distinguished:
Reductiono f the perianthh appensm oreo fteni n 1.T hosew ithf ilamenttsh ata rei nflexedi n theb ud
staminatfel owerst hani n pistillateo nes.N akedp istil- andb endo utwardssu ddenlya nde lasticallya t anthe-
latef lowersc anb e foundi n theA fricang enusT reculia sis,t hrowingo utt hep olleni n thea ir( "urticaceoustsa -
Decaisnea ndi n somep aleotropicaspl ecieso f Ficus.I t mens")T. hist ypeo f stameni s foundi n all generao f
is oftenu ncertainw hetherth ep erianthis stillp resent theM oreae(b utn oti n all species,s eeu nderM aclura);
in pistillatef lowersw hicha ree ntirelyf usedw itht he it is characteristoicf theU rticaceaeS. uddenr eleaseo f
receptacloe f thei nflorescenc(es ucha s in Brosimum). polleni s also reportedfo r some Ulmaceaei:n some
8 FLORAN EOTROPICA
specieso f CeltisL innaeu(sC uellar1,9 67;V azd eArruda Theo varyc anb e freeo rp artlyt o fullya dnateto the
& Sazima,1 988)a ndi nL ozanellaG reenma(nF ranco, tubula(rp arot f the)p erianthT. heo varym ayb e ? stipi-
1992);b utt hem echanismis somewhadti fferenats o nly tate,a featureo ftenc orrelatewd itht hed evelopmenotf
thef ilamenits inflexeda ndn ott hew holes tamenas i nt he theo varyi ntoa dehiscendt rupe( see discussionu nder
urticaceoutys pe.T heu rticaceoums odeo f explosive(o r Fruitsa ndI nfructescences).
ballisticp) ollenr eleaser equireas well-developepde ri-
anthto k eept hel ongi nflexedst amenusn derte nsionT. he
Inflorescences
pistillodec,o mmonlfyo undin f lowerws iths uchi nflexed
Thei nflorescenceisn Moracea(ea sw ella s in other
stamenss, eemsa lso to playa role in them echanism,
familieso f theU rticalesb) asicallyo ccuri n pairsi n the
by keepingt hei nflexeds tamensin ther ightp osition.
leaf axils. However,s olitaryi nflorescenceas re also
2. Thosew ithf ilamentss traightin the budo r in-
commons, ucha s inB rosimuma,n dt heya rec haracter-
flexedb eforea nthesiss, ucha s in Dorstenia,b utb e-
istic for Ficus subg. Pharmacosycea sect. Pharma-
comings traighgt raduallyA. s straighfti lamentso ften
cosycea.P istillatein florescenceas rem oreo ftens oli-
elongated uringa nthesist,h ee longationle adingt o in-
taryt hana res taminatoen es( eveni n thes ameg enus).
flexeds tamensc anb e regardeda s premature.
Pairedi nflorescenceas pparentlya risef romt he two
Thea nthervsa ryf rome xtrorsteo latrorsteo i ntrorse.
(prophyll-bearibnags)a nl odeso f a laterabl ranchw, hich
A peculiarc onstructioonf thea ntheris foundi nB rosi-
mostlyd oesn otd evelopf urthe(rc f.E ichler1, 878)T. he
muma licastrums ubsp. alicastrum.T he antheri s pelt-
prophyllss ometimess ubtendin floresenceso, nly ini-
atea ndt het hecaea ref used.T hisc onstructiocna nb e
tiallyo rp ersistentlyI.n severals pecieso f Ficuss ubg.
relatedto promotioonf anemophil(ys eeu nderP ollina-
Urostigma, theya rew ell-develope(dso metimetso a bout
tion,b elow).
2 cm long) andf used,f orminga calyptratceo vere n-
closingt het woy oungi nflorescence(sf igs).
Ins everagl enera(e .g.,i n thoseo f thet ribeC astilleae
Pistillodes
Pistillodeasr ec haracterisftoicr s omeg roupos f taxa, andi n Ficus),s hort-shoot(ss purs)m ayd evelopi n the
sucha s thet ribeM oreaew ithi ts urticaceousst amens, leaf axils andp roducem oret hant wo inflorescences
and in some subdivisionso f Ficus, includings ubg. (almosts)i multaneousolyr successivelyb,i sexuaol nes,
staminatoen es,o rb othp istillatea nds taminatoen es.I t
Pharmacosycea sect. Pharmacosycea and the paleo-
tropicaSl ycidiumgr oupT. heyr egularloyc curi n some is notc ertainw hethers uchl eafless( buts tipule-bear-
specieso f Dorstenia,a nda rer areo r absenti n most ing) branchletds evelopf romt he samem eristemsa s
otherg enerao r species.H oweveri, n Helianthostylis the( abortivea)x illarby ranchlebtse aringo nlyt hep aired
spruceit hey arep rominentlpy resentw iths tylesa nd or solitary" axillaryi"n florescencesO. n theses hort-
stigmast o 2 cm long, andt heym ightp lay an insect- shootst,h ei nflorescencmesa ya lsoo ccurin p airsT. hese
attractinrgo lei n pollination. short-shootms ayc ontinuet o producein florescences
aftert hes ubtendinlge afhasf allent, husp roducintgh em
on the lesserb rancheso r even on the mainb ranches
Pistil andt het runkT. heses hort-shootasr er arelyb ranched
Theo varyc ontainsa singleo vule,w itha subapical andt heys howr egularp attemsin theiro ccurrenceI.n -
positiont;h es eedi s occasionallayt tachedal mostla ter- florescencems aya lsob e bome( ofteni n pairs)o n an-
ally( Castillat unu,c f. Berg,1 972).T hep ositiono f the othert ypeo f leaflessb ranchleot,f tenw ithl ongerin ter-
ovule is the only "technicalc"h aractewr hichd istin- nodesa nd/oro ftenb ranchedT. hesel eaflessf lowering
guishesM oraceafer omt her elatedfa miliesU rticaceae branchletosf tend o not showr egularp atternisn their
andC ecropiaceaei;n these familiest he ovule has a presencea nda reu suallyf oundo n them ainb ranches
(sub)basapl ositionw, hichs eemst o be correlatewd ith or on the trunk,e ven at the base of the trunk.S uch
the presenceo f single stigma.I n the Moraceaeth ere branchletso ccur in Clarisiar acemosa( and in the
areb asicallytw os tigmasb, utt hes tigmac onnectewd ith paleotropicisn somes pecieso f Artocarpuasn di n nu-
thes ideo f theo varyn otb earingth eo vulem ayb e re- merouss pecieso f Ficus).
ducedi n lengtho rf inallya bsents, ucha sf oundi n some Caulifloryis rarei n then eotropicaMl oraceaer,e -
species of Dorstenia,i n the genusM aclura,i n the strictedto theg enusC larisiaa, ndi t occasionallfyo und
majoritoyf thes pecieso f Ficus,a ndi n thes mallp aleo- in Sorocea.S omen eotropicaFl icuss peciesa rer ami-
tropicagl enusF atoua. florous( to subcauliflorous)C. aulifloryi s far more
Thes tigmasv aryf romf iliformt o band-shapetdo commonin thep aleotropicaMl oraceaaen di s foundi n
tongue-shape(do ri n somep aleotropicaglr oupso f Fi- theA fricang enusT reculiai,n theA sian-Australasian
cus to clavateo rt o infundibuliform). genusA rtocarpusa, ndi n particulairn Ficus, which
