Organization for Flora Neotropica

Cecropia
Author(s): Cornelis C. Berg, Pilar Franco Rosselli, Diane W. Davidson
Source: Flora Neotropica, Vol. 94, Cecropia (May 9, 2005), pp. 1-230
Published by: New York Botanical Garden Press on behalf of Organization for Flora Neotropica
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 FLORA NEOTROPICA MONOGRAPH 94

CECROPIA
CORNELISC. BERG AND PILARFRANCOROSSELLI
WITHA CHAPTERWRITTENBY
DIANE W. DAVIDSON

rRF)Pg( OF CANCER

FLORAL
NEOTROPICA

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Organization for Flora Neotropica
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ISSN 0071-5794 = Floraneotropica.

1. Botany - Latin America - Classification - Collected works.

2. Botany - Tropics - Classification - Collected works. 3. Botany
Classification- Collected works. I. Organizationfor FloraNeotropica.
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 CECROPIA
CORNELIS C. BERG AND PILAR FRANCO ROSSELLI

WITH A CHAPTERWRITTENBY DIANE W. DAVIDSON

CONTENTS
Abstract/Resumen/Resumo.....................................................2
Foreword......................................................3
Introduction.....................................................4
TaxonomicHistory .....................................................4
Morphology..................................................... S5
Habit......................................................5
Sap and Wax.....................................................7
Trichomes......................................................7
Colors......................................................8
Leaves......................................................8
Developmentof Leaves......................................................9
Trichilia..................................................... 13
Stipules..................................................... 14
Inflorescences..................................................... 14
Spikes of Staminate Inflorescences ..................................................... 16
StaminateFlowers ..................................................... 16
Key to the Types of Stamen..................................................... 19
Pollen..................................................... 19
Spikes of Pistillate Inflorescences................ ..................................... 20
Pistillate Flowers..................................................... 20
Fruitsand Seeds ..................................................... 20
Natureand Variationof MorphologicalCharacters.................................................... 20
Anatomy ..................................................... 21
Protectionof Flowers and Inflorescences..................... ................................ 22
Phenology..................................................... 22
Pollination..................................................... 22
Dispersal ..................................................... 23
Survey of Myrmecophytismin the Genus ....................... .............................. 23
OrganismsOtherthan Ants Associated with Cecropia................................................... 23
Ecology..................................................... 24
Distribution..................................................... 25
Representation in the Major Phytogeographic Subdivisions ...................................... 26
SystematicPosition ..................................................... 28
TaxonomicDifferentiation..................................................... 29
Uses ..................................................... 31
VernacularNames ..................................................... 31
Conservation..................................................... 32
SystematicTreatment..................................................... 32
Keys to the Species ..................................................... 32
Key to the Species of CentralAmerica, Mexico, and the West Indies ............ ..........32
Key to the Species of Colombia ............... ...................................... 33
Key to the Species of Venezuela..................................................... 37
2 FLORANEOTROPICA

Key to the Species of the Guianas..................................................... 38

Key to the Species of Ecuador....................................................... 38
Key to the Species of Peru ....................................................... 41
Key to the Species of AmazonianBrazil....................................................... 44
Key to the Species of Bolivia ............. .......................................... 45
Key to the Species of Extra-AmazonianBrazil, Paraguay,and Argentina.......... ......46
Key to the Species of the Cecropiapeltata-group..................................................... 46
Excluded Name ....................................................... 188
Doubtful Names ....................................................... 188
Nomina Nuda ....................................................... 188
Acknowledgments....................................................... 188
LiteratureCited....................................................... 189
NumericalList of Taxa ....................................................... 196
List of Exsiccatae ....................................................... 196
Cecropia and Its Biotic Defenses (by D. W. Davidson)................................................ 214
Benefits to Cecropia-ants....................................................... 214
Benefits to Cecropia ....................................................... 215
InterspecificVariationin Defense Investment....................................................... 217
Ants Symbiotic with Cecropia ............... ........................................ 220
Coevolutionof Ants and Cecropia.................... ................................... 223
EvolutionaryColonization,Host Shifts and HabitatShifts ................... ..................225
Index of Scientific Names ....................................................... 227
Index of VernacularNames ....................................................... 229

ABSTRACT
Berg, Cornelis C. (The Norwegian Arboretum/BotanicalInstitute,Universityof Ber-
gen, N-5259 Hjellestad, Norway; e-mail: cornelis.berg@bot.uib.no),Pilar Franco Ros-
selli (t 3 Feb 2000; UniversidadNacional de Colombia, Institutode Ciencias Naturales,
Santafe de Bogota, Colombia), and Diane W. Davidson (Universityof Utah, 257 South,
1400 East, Salt Lake City, Utah 84112-0840, U.S.A.; e-mail: davidson@biology.utah
.edu). Cecropia.Flora NeotropicaMonograph94: 1-230. 2005. Cecropia,with 61 species
presentlyrecognized,is the largestgenus of the Cecropiaceae.It is a genus of pioneertrees
in the more or less humid parts of the Neotropics. The majorityof the species are myr-
mecophytic. The genus is characterizedby some unusual traits: spathes fully enclosing
the flower-bearingpartsof the inflorescencesuntil anthesis,patchesof dense indumentum
(trichilia) producingMuillerian(food) bodies at the base of the petiole, and anthersbe-
coming detachedat anthesis.The ecological role of the genus and its associationwith ants
have been the subject of numerousstudies, most of them cited in the presentmonograph.
Although the taxonomy has been sorted out reasonablywell, the coverage by collections
and field notes are still insufficientin many cases to fully understandmorphologicaland
ecological variationand plasticity in relation to taxonomic delimitations,and in several
cases, to preparecomplete descriptions.The introductorychapterdeals with taxonomic
history, morphology,ecology, plant-animalrelations, pollination, dispersal,distribution,
classification,diversification,use, vernacularnames, and conservation.Myrmecophilyin
Cecropia is treatedin a separatechapter(by Davidson). In the present contribution,the
new combinationCecropiaschreberianaMiquel subsp.antillarum(Snethlage)C. C. Berg
& P. Francois made and numerouslectotypes or neotypes are designatedas follows.
Neotypes for: Cecropia concolor Willdenow, C. herthae Diels, C. hololeuca Miquel,
C. insignis Liebmann,C. litoralis Snethlage,C. maximaSnethlage,C.palmataWilldenow,
C. pastasana Diels, and C. velutinellaDiels.
Lectotypes for: Cecropiaacutifolia Trecul, C. adenopusMiquel, C. amphichloraStan-
dley & L. 0. Williams, C. angulata I. W. Bailey, C. cyrtostachyaMiquel, C. ferreyrae
Cuatreccasas,C.ficifolia Snethlage, C. glaziovii Snethlage,C. inchuensisCuatrecasas,C.
FOREWORD 3

megastachyaCuatrecasas,C. pachystachyaTrecul, C. palmatisecta Cuatrecasas,C. por-

venirensisCuatrecasas,C. saxatilis Snethlage, C. scabra Martius,C. strigilosa Cuatreca-
sas, C. strigosa Trecul,and C. ulei Snethlage.

RESUMEN
Cecropiacon 61 especies reconocidasen la actualidad,es el genero mas diversificado
de la familia Cecropiaceae.Sus especies son arboles pioneros en localidades huimedasy
muy hdmedasdel Neotr6pico.La mayorfade las especies son mimerc6filasy se caracter-
izan por rasgos extraordinarios:espatas que envuelven completamentelas partes de la
inflorescenciahasta la anthesis, parchesde indumentodenso (trichilia)que producenlos
cuerpos de Muller en la base del peciolo, los cuales sirven como fuente de alimentoa las
hormigas, y anterasque se separanen la antesis. El papel ecol6gico del genero y la aso-
ciaci6n con las hormigashan sido fuente de nuemerososestudios, la mayoriade los cuales
se resefian en la presente monograffa.Aunque la taxonomfa esta razonablementebien
clarificada,el cubrimentogeograficode las colecciones y las observacionesde campo son
insuficientesen numerososcasos paracomprenderlas variacionesmorfologicasy ecol6-
gicas y la plasticidad,y en otros casos parapreparardecripcionescompletas.El capitulo
inicial se relacionacon la historiataxonomica,la morfologia,la relaci6nplanta-animal,la
polinizaci6n, dispersion,distribuci6n,clasificaci6n, diversificaci6n,usos, nombrescomu-
nes y conservaci6n.
Cecropia y sus hormigas son tratadasen un capitulo aparte.En esta contribuci6nig-
ualmente se propone la nueva combinaci6n Cecropia schreberianaMiquel subsp. antil-
larum (Snethlage)C. C. Berg & P. Francoy se designannumerososlectotipos y neotipos.

RESUMO
Cecropia, com 61 especies atualementereconhecidas,e o maior genero das Cecropi-
aceae. Apresentaavores pioneirasnas partesumidas ou semi-hutmidasda regiao Neotrop-
ical. A maioria das especies e mirmec6fita.0 genero apresentaalgumas caracteristicas
impares:espatas cobrindo integralmenteas partesflorais das inflorescenciasate a antese,
partescom denso indumento(triquilios)produtoresdos corpu'sculosde Muller(alimentos)
na base do peciolo, e anteras tomando-se destacadas na antese. 0 papel ecol6gico do
genero e a associac,o com a formigasja'foram assuntoparanumerosaspesquisas,muitas
das quais referidasna presentemonografia.Emboraa taxonomiaseja razoavelmentecon-
hecida, os dados existentes nas coleqc,es e notas de campo sao ainda insuficientes em
muitos casos para serem esclarecidas a morfologia, variac6es ecol6gicas e plasticidade
com respeito as delimita,6es taxonomicas,e em varios casos, na preparac,ode descric,oes
completas. 0 capitulo introdut6rioabordao hist6rico taxonomico, morfologia, ecologia,
relac,o planta-animal,polinizac,o, dispersao, distribuicvo,classificaqao,diversificaqao,
usos, nomes popularese conservaqao.A mirmecofiliaem Cecropiae tratadanum capitulo
a parte.Na presentecontribuicboconstaa nova combinaqaoCecropiaschreberianaMiquel
subsp. antillarum (Snethlage) C. C. Berg & P. Franco assim do designados numerosos
lect6tipos e ne6tipos.

FOREWORD of the monographcame into a final stage, Pilar died

Pilar Franco Rosselli had agreed to participatein
the preprationof a monographof Cecropia in con-
junction with the preparationof a treatmentof the
genus for Colombia,which would have been a partof
her doctoral thesis to be presentedat the University
of Utrecht (the Netherlands).Before the preparation
(03.02.2000) in an accidentwhile makinga collection
of a species of Cecropia.Ourdiscussions on the rec-
ognition and delimitationof some taxa had not fin-
ished, in particularthe inclusion of C. polyphlebia in
C. angustifolia, the way to treat some taxa of the C.
peltata-group, and the publication of a new species
4 FLORA NEOTROPICA

(referredto in the commentunderC. membranacea). TAXONOMICHISTORY

The first authoris solely responsiblefor the final de-
The first accounts on Cecropia are from Marc-
cisions on these mattersand for the final version of grave (1648) and Piso (1658), the latterwith an illus-
the introduction. tration(of "Ambalba")in which the characteristicfea-
tures of the genus were depicted (Piso, 1658: 147).
The genus name Cecropia was created by Loefling
INTRODUCTION
(1758) and is conservedagainstCoilotapalusBrowne
"JederNaturforscher derim tropischenAmerikaje (1756). The name Ambaiba, (also) used by Barrere
gewesen,wirdsichCecropien ( ... ) wohlerinneren; (1741) and Adanson (1763), was reintroducedby
dieselbengehorenzu densonderbarsten Biumenje-
Kuntze(1891) who made severalnew combinations.
nerLander,. . ."(Schimper, 1888:26).-"Everynat-
uralist who has been in tropicalAmericawill Linnaeus (1759) described Cecropia peltata, a
remember Cecropia,belongingto themostconspic- name applied to many species for a long time. Two
uoustreesof thosecountries". additional species were published by Willdenow
(1806), of which C. palmata was also applied to var-
The Neotropicalgenus Cecropiais the largestge- ious species. Some additionalspecies were described
nus of the Cecropiaceae, with 61 species presently by Bertoloni (1840), Martius (1841), and Liebmann
recognized. It is the most importantgroup of pioneer (1851). The first extensive collections of Cecropia,
trees in more or less humid regions of the Neotropics many specimens of several species, were made by
and, therefore,very commonin these regions.In spite Ruiz and Pavon and distributedto many herbaria,of-
of its common and conspicuous occurrence and its ten in mixed sets. This mixing of specimens of more
importantecological role, it has been studied little than one species duringdistributionor mountingis a
taxonomicallyandthe studiesavailablewere not very problemwith many older (andeven more recent)col-
useful. lections of Cecropia. The Ruiz and Pavon material
There are severalreasons for the slow progressof was used for descriptionsof severalspecies by Trecul
taxonomic knowledge. One reason was (and still is) in 1847 and later in the same year by Klotzsch. The
the scarcity of collections and the poor quality of type materialof the species describedby Klotzsch in
many of them. Most plant collectors avoid Cecropia Berlin was destroyed.Due to the mixing of elements
because the plant parts are too large to easily make of the Ruiz and Pav6n collections, it is not easy to
good herbariumspecimens, most species are inhab- relate duplicatesto the Klotzsch names. The species
ited by aggressiveants, andmost species areelements decribed by Trecul and Klotzsch were included in a
of secondarygrowth. Because the species are super- survey of the genus by Miquel (1853) in Flora Bras-
ficially so similar,even conscientious collectors nor- iliensis, who added several new ones and recognized
mally collected eitheronly staminateor pistillatema- 37 species for the Neotropics,some of them based on
terial of one species and left uncollectedthe othersex material in cultivation in Hortus Schonbrunnin Vi-
and the other five to seven or so species often occur- enna. For a long time, Miquel's was the only com-
ring at a locality. The relative paucity of collections prehensive taxonomic treatmentfor the neotropical
made (and still makes) it difficultto preparedescrip- region.A numberof species were describedby Hems-
tions of all details (such as those of stamens)for sev- ley (1883), Richter (1897), Donnell Smith (1899),
eral species and to understand the extent of the Rusby (1907, 1910), Huber(1910), Robinson(1912),
morphological variation; to evaluate the nature of Pittier (1917), and Bailey (1922) before Snethlage
morphological differences; and to comprehend the startedto work on the genus and published a good
ecological variation,the limits of the ranges of dis- numberof new species (1923, 1924), many of them
tribution, and the reality of disjunctions. It is often based on the collections of Ule, who consistentlytried
still difficult to decide whether differences found in to collect sets of both staminateand pistillate mate-
species describedfromdifferentregions(oreven from rial. In 1923, Snethlage also proposed a subdivision
the same region) justify recognition at the species for the genus. Several additionalspecies were pub-
level. The numberof collections of severalrecognized lished by Burret (1924), Mildbread (1925, 1933),
taxa is so small in relation to the variationthat their Standley (1929, 1940a, 1940b), Macbride (1937),
circumscriptionsare incomplete. Because important Diels (1941), Standley & Steyermark(1944), and
diagnostic characterswere often not included in de- Standley & Williams (1952). In the period 1932-
scriptions, even in the extensive ones provided by 1970, Cuatrecasasmade extensive collections of Ce-
Cuatrecasas(1945, 1956), identificationof material cropia in Colombia, providingthe basis for the pub-
has often been impossible or at least difficult. lication of numerous Colombian species (1944,
MORPHOLOGY
1945). Collections made in later years, also from
other parts of South America, provided the material
for the establishmentof a total of 68 additionalspe-
cies and several variety names (Cuatrecasas,1949,
1951, 1956, 1959, 1967, 1971, 1976, 1982). Cuatre-
casas planned to revise Cecropia and the related
Coussapoa Aublet and PouroumaAublet (Cuatreca-
sas, 1945). Unfortunately,he could not realize his
plans. Many of the names establishedby Cuatrecasas
are synonymizedin the presenttreatment.One reason
is that he paid little attentionto previouslypublished
species, possibly as he could not get hold of type ma-
terial from European herbariabecause of war and
postwar circumstances when he started to publish.
The other reason is that he did not or could not un-
derstand the variation, plasticity, distribution, and
ecological amplitudesof most Cecropiaspecies, treat-
ing them as "normal"species, clear-cutin their mor-
phology, distribution, and ecology. More recently,
poor understandingof the natureof variationand dis-
tributionpatternsin Cecropia caused prematurees-
tablishmentof a numberspecies (Berg & FrancoRos-
selli, 1993, 1996).
In 1972 the senior authorof this treatmentstarted
to work on the genus, initially on the taxa occurring
in AmazonianBrazil, later extendingthe study to in-
clude extra-AmazonianBrazil, the Guianas, Vene-
zuela, and Peru (Berg, 1972, 1977a, 1978b, 1980,
1981, 1985, 1992, 1998, 2000, 2002; Berg & Carauta,
1986). More recently, these studies included collab-
orationwith FrancoRosselli in Ecuadorandthe entire
Neotropics (Berg & FrancoRosselli, 1993, 1996).
Regional treatmentsof the genus include those for
Peru (Macbride,1937), Panama(Woodson& Schery,
1960), Costa Rica (Burger, 1977), Venezuela(Velas-
quez, 1971), the species of Kcosniipataand Manuval-
leys in Peru(Galiano-Sanchez,1976), andthe stateof
Rio de Janeiroin Brazil (Duarte, 1959).
The Cecropia-Aztecaant associationreceived at-
tention in several papers in the 19th century and the
beginning of the 20th century (Muller, 1876; Schim-
per, 1888, 1898; Buscalioni & Huber, 1900; Rettig,
1904; Ule, 1900;Ihering, 1907;Fiebrig, 1909;Bailey,
1922; Wheeler & Bequaert, 1929; Wheeler, 1942).
Numerousmore recentstudieson the myrmecophytic
traits have been published by Janzen (1969, 1973),
Janzen& McKey (1977), Andrade(1984b), Andrade
& Carauta (1979, 1982), Longino (1989a, 1989b,
1991a, 1991b), Davidson et al. (1988, 1991), and
Davidson & McKey (1993). Otheraspects of the bi-
ology of Cecropiathathave been studiedincludepop-
ulation dynamics (Alvarez-Buyalla & Martinez-
Ramos, 1992); the role as pioneertreein treefallgaps,
5
forest edges, landslides, etc. (Brokaw, 1985, 1986,
1987, 1998; Guariguata,1990; Didham & Lawton,
1999; Sposito & Santos,2001a, 2001b); light regimes
(Davidson & Fisher, 1991; Folgarait & Davidson,
1994); dispersal (Charles-Dominique, 1986); seed
banks (Holthuijzen & Boerboom, 1982; Vazquez-
Yanes & Smith, 1982; Alvarez-Buylla & Martinez,
1990; Garay-Arroyo& Alvarez-Buylla, 1997); con-
trol of seed germination(Vazquez-Yanes& Smith,
1982; Vazquez-Yanes & Orozco-Segovia, 1986);
seedling and saplingdevelopmentandphotosynthesis
(Reekie & Bazzaz, 1989; Strauss-Debenedetti& Baz-
zaz, 1991; Poorter& Oberbaurer,1993); tannincon-
tents (Coley, 1986); isozyme variation(Garay-Arroyo
& Alvarez-Buylla, 1997); associations with various
groups of beetles (Andrade, 1984a; Jolivet, 1987,
1989, 1990b); exploitationby leaf-cuttingants (Vas-
concelos & Casimiro, 1987); and presence of lianas,
hemiepiphytes,and hemiparasites(Putz, 1982; Clark
& Clark, 1990; Jolivet, 1990a).
MORPHOLOGY
HABIT
Cecropia trees are few-branched,usually with a
candelabrum-likebranchingsystem and with the ar-
chitectural Model of "Rauh" (Halle & Oldeman,
1970; Halle et al., 1978). In some species of small
trees the branchingis reduced,so thatthe tree is usu-
ally or mostly monocaul (as in C. megastachyaand
C. ulei). A tendency to monocauly is also found in
some types of C. obtusifolia in western Colombia
(Valle). In many species (e.g., C. concolor and C. fi-
cifolia), flowering often startsin the monocaul state.
Branch development is often initiated in seedlings,
even in the axils of the firstformed(opposite)leaves;
prophylls are formed, and often the developmentof
the first leaf begins but is arrested(if the seedling is
not decapitated).In the axils of the leaves formeddur-
ing later development,the axillary branchprimordia
do not producemore thanone or two prophyllsand a
bud.
In Cecropiagarciae and C. hispidissima,branch-
ing often occurs at a height of 0.5-1 m; the branches
departat acute angles and the tree has a bushy habit.
More commonly, the branchesdepartat more obtuse
angles (at least the lower ones mostly at ca. 450), and
the crown becomes more or less distinctly umbrella-
shaped.However,in some species, as C. putumayonis
and C. utcubambana,as well as forms of C. obtusi-
folia, the crown is usually small and more or less glo-
bose.
A remarkablevariation in branchingis found in
6 FLORANEOTROPICA
Cecropiaalbicans. In this species, the branchestend
to be short,departingat wide angles, or the treesome-
times remains unbranched.In a populationin Huan-
uco, Peru,individualsrarelyproducelateralbranches
departingat wide angles, but often producebranches
departingat acute angles in one or two whorls.
Most Cecropia species form small to medium-
sized trees, 5-15(-20) m tall; in others, like C. di-
stachya, C. herthae, C. insignis, and C. sciadophylla,
trees become taller,even to 40 m tall. In others(e.g.,
C. ulei) the height rarely surpasses5 m. The size of
the trees shows regional variation in some species,
such as C. pachystachya, which is representedby a
small type in restingas of easternBrazil (Andrade&
Carauta,1981), and C. concolor nearManaus.
A conspicuous variationin habit is found in the
co-occurring Cecropia concolor and C. polystachya
in SantaCruz, Bolivia, at the (climatic) limit of their
distribution. Here, trees are robust with relatively
short,thick stems and brancheswith shortinternodes
and relatively dense and wide umbrella-shaped
crowns. This tree shapeis typical of montanespecies,
such as C. telenitida or C. plicata. Individualswith
this tree shape are found within short distances of
others with the shape normallyfound in the two spe-
cies. This variation in habit occurs occasionally in
otherspecies, e.g., a largetreeof C.putumayoniswith
a broad umbrella-shapedcrown (near Villavicencio,
Meta, Colombia), apparentlyin the peripheryof its
range. Individuals of C. obtusifolia are atypically
large-canopy trees in Veracruz, Mexico, and are
therewith extending longevity (Alvarez-Buylla &
Martinez-Ramos,1992) and those of C. membrancea
often are large-canopytrees outside theirtypical hab-
itat (see p. 119).
The differencesin tree shape and size appearto be
related to longevity (see Sposito & Santos, 2001a,
2001b); the large or robust specimens (rarein some
species, but common in others) are less ephemeral
than normal in the genus (see Vester, 1997 for Ce-
cropia distachya, C. ficifolia, and C. sciadophylla).
Accordingto Duarte(1959), trees of C. hololeuca can
become more than 100 years old, whereasthose of C.
glaziovii have a lifespan of 30-40 years. The mono-
caul treelets of C. ulei live for only a few years. This
suggests that the tree shape as found in montanespe-
cies could be (at least partly)ecologically defined(by
climatic conditions,affecting growthrate)ratherthan
genetically. From the variation in habit described
above, it is clear that various factors play a role in
shaping the trees of many Cecropia species.
All Cecropiaceaecan produceadventitiousroots.
In Cecropia,they become stilt-roots,which are com-
mon and manifest, especially in large trees and in
trees growing along rivers or in marshyplaces.
Although Cecropia trees often occur in sites rich
in vines, they are rarelyovergrownby them (cf. Clark
& Clark, 1990), even individualsthat are not inhab-
ited by Azteca ants (cf. Putz, 1984; Putz & Holbrook,
1988). Vines can often be observed on the stem and
the lower partsof the lower branches,but not higher
up. This also applies to the non-myrmecophyticAf-
rican sister-genus Musanga R. Brown with a habit
similarto thatof most Cecropiaspecies. The angle of
ca. 450 at which (at least) the lower branchesdepart
from the trunkin these genera may affect the growth
of vines on the tree. In (still) unbranchedspecimens,
the rapidcontinuousgrowth and the continuousloss
of the lower leaves (and handholds for vines) may
reduce possibilities to become overgrown.
In most species, the internodes are hollow and
contain sparse,usually whitish pith. Such internodes
providenesting space for ants. However,in some spe-
cies the intemodes are usually or often filled with
brownpitheithercompletely(as Cecropiabullataand
C. gabrielis) or partly(as C. schreberianaand C. te-
lealba). In some other species, abundantbrownpith
is found occasionally (as in C. strigosa) or consis-
tentlyin partsof the species range(in C. angustifolia).
Abundantpith is found mostly in montanespecies, C.
schreberianabeing a distinctexception.The presence
of abundantpith could be relatedto a low growthrate.
In young plants, the diameter of the stem often
increases considerably upward, either gradually or
more or less abruptly.The walls of the inflatedinter-
nodes are thin. In some species, Cecropia albicans
and C. elongata, this phenomenon can also be en-
countered in adult trees. Stems with inflated inter-
nodes are also found in C. angustifolia in Ecuador.
Most species have distinctprostomata, thin spots
in the upperpart of the wall of the interuodes.They
are located above the insertion of the petiole, as a
small depression, often at the end of a longitudinal
groove thatstartsjust above the middle of the petiole.
The development of the groove and prostoma (or
"diaphragma")is describedand discussed by Schim-
per (1888).
In trees occupied by ants, the prostomata are
mostly opened and thus present as small holes. By
perforatingprostomata,ant queens enterthe stem and
establishcolonies. In non-myrmecophyticspecies, the
prostomatacan be eitherobvious, as in Cecropiascia-
dophylla (see Bailey, 1922: 385) or are more or less
indistinctto (virtually)absent,as in C. hololeuca (see
Schimper, 1888: fig. 7; Schimper, 1898: fig. 82) and
severalAndeanspecies in which the outerwoody wall
MORPHOLOGY
of the internodes of leafy twigs (with short inter-
nodes) is relatively thick; even if prostomatadepres-
sions can be detected, they cannot be penetratedby
ants.
Whetheroccurrenceof pith in the internodesand
the absence of (distinctor functional)prostomatacan
be regardedas primitivestates or as consequencesof
growth rate, which tends to be less in montanethan
in lowland species, is not clear, but these featuresaf-
fect inhabitationby ants.
The length of the internodes varies. In young
plants, the internodesare long, but in many species
the internodesbecome short (0.5-2 cm long) in adult
trees, especially in the distal partsof the branches.In
other species, the internodesremainlong (>2 cm) in
adult trees, especially those of the continuously fast
growing, "weedy"species (see below). Shorteningof
the internodes is correlated with harder internode
walls.
On the stem and main branches,the scars of the
stipules are usually conspicuous and in some species
(e.g., Cecropia annulata, C. engleriana, and C. lito-
ralis) presentas prominentannularridges.
SAP AND WAX
Most parts,like young branches,petioles, and pe-
duncles, exude a watery, often somewhat mucilagi-
nous sap, when cut. Exposureto the air turnsthe sap
black (and can cause persistentdark stains on cloth-
ing). Ducts and cells containingthe mucilaginoussap
are common in Cecropiaceae(cf. Renner, 1907). In
some montane non-myrmecophytic species (e.g.,
Cecropia telenitida), mucilage may fill the space in
the terminalbuds and in the upper internodes.This
preventsinhabitationby ants but also preventsor re-
duces occupation and damage by herbivorousinsect
larvae.
In several species, leafy twigs are covered by a
waxy layer making them bluish. Such a layer is al-
ways presentin Cecropiaandina; in some other spe-
cies it is often, occasionally, or (in C. albicans) re-
gionally present. This layer is found only on
(sub)glabroustwigs. However,leafy twigs with waxy
surfacesor dense (villous) indumentumcan be found
in individualsof the same species (e.g., C. telenitida).
Tanninis often abundant(Renner,1907), as in the
rachises of the spikes of the inflorescence,the apices
of the perianthsof pistillate flowers, and in the peri-
carps.
TRICHOMES
The abundanceand composition of the indumen-
tum are importantfeaturesin Cecropia,morphologi-
7
cally and partly also functionally. Six types of tri-
chomes can be recognized.
1. Unicellular hairs which are relatively thick,
straight,curved, uncinate, or crinkled. In some spe-
cies these hairs can be setose, and, e.g., in Cecropia
hispidissima and C. megastachya placed in sockets
and resembling the urticating trichomes of Urtica-
ceae. Such hairs easily breakat the sockets, causing
rough surfaces.They easily penetrateand irritatethe
skin. Uncinatehairs are very common. Theirabsence
is a differentiatingcharacter.
2. Unicellular hairs which are very thin and in-
terwoven, usually white but sometimes brownish.
This arachnoid or cobwebby indumentumis found
on several parts of the plant. Dense covers of arach-
noid indumentum,causing white or whitish surfaces,
are common on the spathe, the stipules, the petiole,
and the upper and lower surfaces of the lamina. On
the lower surface,this indumentumcan be very short
and confined to the areoles. Arachnoidindumentum
of longe hairs is always present on the outer surface
of the perianthof the pistillate flower, shorterhairs
are often found inside. If arachnoidindumentumoc-
curs on the outer surfaceof the perianthof staminate
flowers,then it is usually short.Hairsas long as those
on the perianthof the pistillate flower are found in
some species, e.g., Cecropiaglaziovii and C.palmata.
3. Pluricellular trichomes. They are eitherelon-
gate and usually more or less clearly moniliformor
shortenedand more or less globular.They are mostly
pale to darkbrownor sometimes reddishor purplish.
They arecommonly found on young parts.In dry ma-
terial, the indumentumof pluricellulartrichomes is
usually manifest as a powdery layer.
4. Cystolith hairs. These protrudingcystoliths
are found at the uppersurfaceof the laminaandcause
a rough surface.
5. Pearl glands (or "pearl bodies"). These pluri-
cellulartrichomesare ca. 1-2 mm long, (sub)clavate,
and whitish opaque. According to Rickson (1976),
they containa few glycogen plastidsandlipids. These
trichomesarepresentin all (or most) species. Because
they are apparentlysoon harvestedby ants (cf. An-
drade, 1984a), they are not often observedin the field.
They are abundantlypresenton young partsof plants
grown in greenhouses.Pearl glands are also found in
CoussapoaandMyrianthusP. Beauvois (othergenera
of the Cecropiaceae).
6. Mullerian bodies. These trichomesoccuronly
in patches (or cushions) of dense indumentum(tri-
chilia, see below). They are ellipsoid to oblongoid,
mostly ca. 1-2 mm long, and whitish; in Cecropia
chlorostachyathey are pinkishand in C. hispidissima
8 FLORANEOTROPICA
pink and ca. 3 mm long. Their inner cells are filled
with glycogen plastids, and in the apex of the tri-
chome the epidermis is provided with a stoma
(Schimper,1888;Renner,1907;Rickson, 1976). They
abscise when full-grownand are "pushed"upwardto
the surface of the trichilia by the surroundingdense
indumentum,consisting largely of pluricellulartri-
chomes. Moreover,this indumentumkeeps the Mill-
lerian bodies at the surfaceof the trichiliaand delays
droppingfrom the trichilium (Schimper, 1888). The
bodies are rich in lipid and also contain protein and
glycogen (Marshall & Rickson, 1973; Rickson,
1973). These trichomes can be regardedas modified
pearl glands. A detailed account of the development
of trichilia and Miillerian bodies was provided by
Rickson (1976). The Mullerian bodies are the main
food source for ants associatedwith Cecropiabut are
also eaten or collected by other animals:beetles (cf.
Andrade, 1984a), bees, and small birds. In material
grown in greenhouses (where Mullerian bodies are
not removed by animals), the trichilia are often soon
covered by fungi. Andrade (1982) suggested that
Muillerianbodies could have a protective function
equivalentto thatof extrafloralnectaries.Food bodies
are also found in the swollen and saccate base of the
petiole of Pouroumaformicarum and P myrmeco-
phila (Benson, 1985; Berg et al., 1990). Whether
these food bodies have the same structureand con-
tents as Miillerianbodies is unknown.
The indumentumvaries considerablywithin spe-
cies. This is most conspicuousfor the white arachnoid
indumentum.Dense (villous or velutinous)indumen-
tum on leafy twigs-as found in Cecropia angusti-
folia, C. albicans, C. bullata, C. telenitida, and C.
velutinella-may affect the inhabitation by ants,
hindering locomotion. In some of these species (C.
albicans and C. telenitida), both this type of dense
indumentumand very smooth, waxy twig surfaces
occur.Scabroussurfacesarecausedby cystolithhairs,
short and rigid unicellularhairs, or bases of bristle-
like hairs.
COLORS
Red-coloringsubstancesare quite commonin var-
ious plantparts,in particularthe stipules, the spathes,
and the main veins of the lamina. The concentration
of these substances varies, even within species, col-
oring the parts pale pink to dark red, or even dark
purple to almost blackish. The coloring substances
may be distributedequally or in patterns,e.g., lon-
gitudinal stripes. Colors may fade with age. In some
species (e.g., Cecropia membranacea)red-coloring
substancesare rareand all partsare usually green. In
several species, both red morphs (with red-colored
stipules, etc.) and green morphs (with green-colored
stipules, etc.) occur. Both morphs can be found side
by side throughoutthe range of the species (as in C.
heterochroma)or in separateparts of its distribution
ranges (as in C. angustifolia). Proportionsof red and
green morphs vary within populations. Leaves of
young plants of many species are often silvery mac-
ulate.
Colors of plant parts of Cecropia can also be de-
termined or influenced by dense indumentum.The
white arachnoidindumentumcan be so dense as to
mask underlying reddish colors such that spathes,
stipules, or leaf surfacesappearwhitish. In some spe-
cies-in particularseveral montane Andean species
(C. albicans, C. maxima, C. telealba, and C. teleni-
tida), but also in the Brazilian C. hololeuca, which
occurs at low elevations-the layer of arachnoidin-
dumentumon the uppersurfaceof the laminais usu-
ally so dense that the trees can then be detected at
great distances by their white leaves. More or less
white and green surfacesmay occur in the same spe-
cies. In C. pastasana, the lamina is often distinctly
white above in the southernpartof its rangebut only
slightly so in the northernpart. In C. reticulata,the
laminais usually green but sometimeswhitish above.
In C. maxima, the lamina is normally white in pop-
ulations occurringat 1500-2400 m, but in a popula-
tion at ca. 1000 m (in Narifno,Colombia) the leaves
are green above.
The brown to purplish(or blackish) pluricellular
trichomescan be so dense that they impartthese col-
ors to the plantparts.In Cecropiaangustifolia,brown
trichomes on leafy twigs may turn black and make
younger partsof the stems blackish.
In some species, young plant partsare coveredby
a bluish waxy cover;always in the stems of C. andina
and sometimesin C. albicans and C. pastasana. Len-
ticels, in particularlarge white and pink ones, may
add to the color range,particularlyon leafy twigs.
LEAVES
In all Cecropia species, the leaves of adult speci-
mens are large and peltate, almost circularin circum-
ference. The lamina is excentrically attachedto the
petiole. The venationis radiateand the laminais usu-
ally radially incised in between the radiatingmain
veins. Incisions can run down to the petiole, and the
leaf segments can then be petiolulate (in C. sciado-
phylla). In some species (e.g., C. putumayonisand C.
subintegra),the lower partof the lamina is hardlyor
not incised. The number of lobes or leaf segments
vary from 5 to more than 20. Withinthe species, the
MORPHOLOGY 9

number of segments of the lamina mostly varies (Figs. 1.3, 1.6). Even later leaves have inequilateral
within certainlimits. laminas of which the correspondinglobes of both
In some species (e.g., Cecropiapachystachyaand sides arenot equally long. Lobationis correlatedwith
C. polystachya),the upperlamina segments are often a changein venation.Initiallyone or two lateralveins,
or usually lobed; the lobes are usually more frequent usuallynot the basal ones, become stronger(Figs. 1.5,
and pronouncedin subjuvenile stages than in adult 1.13). However,the formationof lobes is soon asso-
specimens. ciated with the basal pair of lateral veins. The tran-
The main patternof venation is ratheruniformin sition from the 3-lobate state to the 5-lobate one goes
the genus. However, the details of the venation may with the formation of strong veins departing from
vary somewhat to considerably within many of the the base of the strong primary basal lateral veins
species. The variationis partlycaused by shifting of (Fig. 1.8). The base of the lamina of the first formed
featuresof the subjuvenilestate to those of the adult leaves is acute to rounded (Figs. 1.1, 1.2, 1.9), de-
state (or vice versa). In spite of the variation,the ve- pending on the species. Then the base becomes cor-
nationprovidesimportantdiagnosticcharacters,such date and successively peltate (Figs. 1.7, 1.9). Pelta-
as the numberof secondaries of the main radiating tion may precede lobation or may happen after the
veins, the branchingof the secondaries,and the loop- firstpairor two of lobes have been formed.In species
connections of the secondaries in the margin or with laminas with the radial incisions down to or
(sometimes closely) near the margin:in the descrip- nearly down to the petiole, as Cecropiasciadophylla
tions and keys this is indicatedas marginallyand sub- or C. concolor, the lamina becomes peltate when the
marginallyloop-connected.The venationbetweenthe sixth lobe is formed, opposite the midlobe (or mid-
secondariesis scalariformwith parallelveinlets. segment) (Figs. 1.14, 1.15). In all species the sixth
The lamina can be coriaceous (to subcoriaceous) lobe or segment is formed opposite the midsegment.
or chartaceous(to subcoriaceous).Coriaceous lami- The fact that there are often two veins runningclose
nas are often smooth, and chartaceous ones often togetheroppositethe primarymidrib,or thatthe mid-
more or less scabrous.Segments of coriaceous lami- rib of the sixth lobe is often furcate,suggests thatthe
nas are usually plicate in Cecropiaplicata or some- midrib of the sixth lobe (and other lobes opposite
times in C. insignis, C. strigosa, and C. telealba. In the midsegment) is formed by fusion of two veins
some species the uppersurfaceis usuallymore or less (Fig. 2).
bullate in C. bullata and often so in C. andina. The next step is the formation of a lamina with
The length of the petiole (in adult trees) is often seven lobes (Figs. 1.16, 1.17). The midribs of these
proportionalto (and often approximatelyas long as) lobes departfrom the bases of the primarybasal lat-
the length (and width) of the lamina. However, in eral veins. The increase of the number of lobes or
some species, e.g., Cecropia marginalis and C. vir- segments is caused by the successive formation of
gusa, the petiole is about half as long as the lamina, branchesfrom the base of the (original)pair of basal
with a maximumlength of ca. 40 cm in adult speci- veins in basipetal direction: thus from 3 -X 5 -> 6 -*
mens. 7 -> 8 -> 9 -X -lO10 1 and so forth, up to 24 (in
In the majorityof the species the laminais folded Cecropia velutinella).The even numbersof lobes are
in the bud and extendedfrom the petiole; in Cecropia causedby the presenceof a segmentoppositethe mid-
angustifolia and C. montana the lamina is often re- segment. The attachmentof the laminato the petiole
flexed aroundthe relativelylong petiole in the bud. moves from the base towardthe centerof the lamina,
The circulararea where the radiatingveins meet but remainingmore or less pronouncedlyexcentric.
at the uppersurface of the lamina is in some species These developments also affect other features of
descriptionsindicatedas the "umbilicus." the venation, with an often considerableincrease in
the numberof lateralveins (Figs. 1.2, 1.3, 1.13, 1.14).
DEVELOPMENTOF THE LEAVES The events of the developmentas describedabove
The cotyledons are small and somewhat cartilag- are (at least in part)repeatedon the firstnodes of the
inous, and lack chlorophyll and venation. The first branches.
trophophyllsare opposite. The two sets of opposite Before branching,the leaves on the stem are often
leaves are succeeded by leaves arrangedin spirals. larger,are more deeply incised, and often have more
The firstformedleaves arepinnatelyveined andentire segments and more lateralveins in the free partof the
andhave a basally attachedlaminaand a shortpetiole midsegment;in certain species the free parts of the
(Figs. 1.1, 1.2, 1.9). Laterleaves have a longerpetiole segmentsare more pronouncedlylobatethanthose on
and the lamina lobed on one side or both sides the branches.Such subjuveniletraitscan be retained
10 FLORA NEOTROPICA

~~~2 }

6 7 810

l9 C 1010~~~~~~~~~~~1
FIG. 1. Early stages of the development of the leaves of Cecropia, schematic and not on scale. C. sciadophylla,
1 2 -> 3 -* 6 -* 11 -> 15; C. membranacea,4 -* 5; C. putumayonis,7; C. subintegra,8 -> 12; C. concolor, 9 -* 10;
C. angustifolia 13 -* 14; C. multisecta, 16 -> 17.
MORPHOLOGY 11

1z 13

1F6 171
FIG. 1. Continued
12 FLORA NEOTROPICA

1 2.

FIG. 2. Development of the venation at the base of the lamina of Cecropia subintegra.Stages 1 2 - 3; whole
development of the main venation, schematic, 4.
MORPHOLOGY
in leaves on the branchesto some extent. A remark-
able situation is found in Cecropia subintegra, in
which the leaves of the main stem are deeply incised
with numerous segments, whereas the leaves of the
branches are subentirewith relatively few radiating
main veins.
Laminas with incisions down to the petiole or
nearly so reach the basic shape of the lamina rather
quickly in the process of leaf development.In these
species, furtherdevelopmentimplies the increase of
the numberof segments, usually withoutshowing in-
termediatestages. This type of lamina is rathercon-
stant in its features. Another type of lamina mostly
showing stability in its featuresis the one with rela-
tively few (ca. 7) radiatingmain veins anda shallowly
incised or (sub)entirebase. Species with intermediate
traitsof the laminaoften show considerablevariation
with regardto the numberof segments and the extent
of the incisions.
The varioustraitsof the developmentof the lamina
are presentedin Figures 1 and 2.
TRICHILIA
In most species of Cecropia, one or two patches
of dense indumentum are found abaxially at the
broadenedbase of the petiole of the adult leaf. They
consist of various types of trichomes: whitish (to
brownish) unicellularand more or less stiff hairs of
various lengths, dense and relatively long brown
pluricellular trichomes, and Mullerian bodies (see
above). These characteristic concentrations of tri-
chomes are called trichilia. In literturethe trichilium
has been called "pulvinus,""giba,"and,in the Spanish
descriptionsby Cuatrecasas(1945, 1956), "cojin"or
"almohadillabasilar."
The Mtillerianbodies are usually initially hidden
by relativelylong and very dense brownpluricellular
hairs, probablyto preventprecocious harvesting(by
animals other than Azteca ants inhabitingthe tree).
When "mature,"the Muillerianbodies become de-
tached from their bases and they are pushed upward
to the surfaceof the trichiliaby the surroundingdense
indumentumand are "released"in the course of some
weeks. A single trichiliummay produce 2500-8000
Muillerianbodies (Janzen,1973) and/orca. 10 g Mill-
lerian body tissue a week (Rickson, 1973) in C. pel-
tata.
The unicellularhairs intermixedwith the brown
indumentumof pluricellularhairs vary in density and
length: from about as long as to somewhatlonger to
conspicuously longer than the brown pluricellular
trichomes.The trichiliamay vary more or less in size
13
and shape. In some species (as Cecropiapastasana
and C. reticulata), the trichilia cover the scrotiform
swollen bases of the petioles. In some other species
(as C. herthae), the trichilia cover shallow depres-
sions at the base of the petiole.
The trichiliaof Cecropiavirgusaare hiddenin lat-
eral (depressions)pockets with a slit-shapedopening
at the base of the petiole. The pockets develop as
grooves departingfrom the base of the petiole and
become longer anddeeperbefore trichiliaareformed.
In Cecropiahispidissima,unlikethe otherspecies,
the pluricellularhairs are short and do not cover the
relativelylarge(ca. 3 mm long) Miillerianbodies. The
trichiliaare often not clearly definedpatchesand dif-
fer in size. They may even extend to the base of the
stipules or may be almost absent. This species is in-
habited by ants of the genus Pachycondyla,which
harvest the Miillerian bodies when the trichilia are
still enclosed by the stipules.These antspenetratethe
node septum below the bud cover to enter the space
containingthe Mullerianbodies.
The majorityof the species have a single trichil-
ium, median and extending laterally.Cecropia mar-
ginalis and C. strigosa usually or often have two lat-
eral trichilia; they are fuly developed. Separate
trichilia are often found in species in which trichilia
are often or sometimes absent. In these species tri-
chilia occur in a "primordial"state, i.e, as two small
patches with only unicellularhairs, as small patches
(of equalor unequalsize) also with pluricellularhairs,
or as largerpatchesalso containingMullerianbodies,
and finally large fused patches. This appearsto indi-
cate that fused trichilia representthe advancedstate.
In species with consistentlyfully developedfused tri-
chilia, the first formed trichiliaon the sapling are al-
readyfused. In these the firsttrichilia(initiallyusually
without Mullerianbodies) appearon the leaves (5th-
10th formed) with long petioles and peltate laminas.
However, in some species (C. latiloba and C. mem-
branacea, both of periodically inundatedhabitats),
trichiliamay appearmuch laterin the developmentof
the trees, at aboutthe 40th-5Othleaf formedafterthe
cotyledons(Berg, 1978b), as observedin materialcul-
tivatedin a greenhousefrom seeds obtainedfrom an
area near Manaus,Amazonas,Brazil. But, at least in
aberranttypes of C. membranacea,trichilia can al-
readybe presentin a very early stage of development.
This variationcould be related to differences in the
degree of inundation.
Trichilia are always completely absent in some
species: Cecropia hololeuca, C. pittieri, C. sciado-
phylla, and C. tacuna.In severalspecies, trichiliamay
14
be absent or present, and if present,in various states
of development. In C. schreberiana, trichilia (often
reducedand two per petiole base) are sometimespres-
ent but, according to Rickson (1977), without Mill-
lerian bodies (or with perhaps abnormal ones; cf.
Wheeler, 1942). In other species, absence, presence,
and differentdegrees of developmentof the trichilia
appear to be accidental, while in still others these
states are characteristicsof populations or certain
partsof the distributionrange. Clearcorrelationwith
age or habitat are wanting. Nor does absence, pres-
ence, or state of the trichilia appearto be correlated
with the amount of pith in the internodes and,
therewith,the suitabilityfor inhabitationby ants. In
one of the morphologicaltypes of C. angustifolia,the
internodes are filled with pith but the trichilia are
mostly fully developed, yet in C. gabrielis, also in
which the intemodes are filled with pith, trichiliamay
be absent, more or less reduced, or fully developed.
These different stages of developmentalso occur in
C. telenitida, in which, however, the intemodes are
clearly hollow. In C. albicans, trichilia are present
only in some parts of the species' range and are as-
sociated with differences in indumentum and tree
shape.
Absence of trichilia or presence in various states
is usually associatedwith species of montaneor sub-
montane habitatsin the Andes. Trichilia are always
absent in the lowland species Cecropia pittieri, C.
hololeuca, and C. sciadophylla, and mostly in C.
schreberiana.
In most other Cecropia species, the majority of
which occur in lowland habitats,trichilia are always
presentand well-developed. Cecropiapeltata is a re-
markable exception, for the trichilia are well-
developed throughoutits wide range of distribution
except in Jamaica,where trees with and without tri-
chilia can be encountered.In Africa, Asia, and the
Pacific, where this species is introduced,trichiliaare
absent or poorly developed.
The types of trichomesfound in trichiliaare com-
mon in Cecropiaceae, but their concentrationinto
more or less clearly definedpatches and the transfor-
mation of pearl glands into Mullerianbodies is char-
acteristic only of Cecropia. The trichiliumis one of
the requisitesfor the association with ants. The pres-
ence of ants can be related to preventionof damage
by fungi and insects (cf. Bailey, 1922: 388). The hol-
low leafy stems with thin internodewalls and the ap-
ical meristem are susceptible to damage. Their pro-
tection, particularlyin early stages of development,
i.e., before branchingtakes place, promotesthe rapid
and continous growthof the pioneer tree.
FLORANEOTROPICA
STIPULES
The stipules in Cecropiaare always fully amplex-
icaul and fused. They vary in length from about 5 to
50 cm, often dependingon the diameterof stems and
leafy twigs in species. However, some species with
relativelyslender stems and brancheshave long stip-
ules and then often terminalbuds with a curvedapex
(e.g., C. marginalisand C. virgusa). The stipules are
usually caducous, but in some species (e.g., C. chlo-
rostachya and C. palmata) they remain temporarily
attachedto the stem, and in other species (e.g., C.
angustifolia and C. engleriana) they are sometimes
or often subpersistent,or they may be only subper-
sistent in juvenile or subjuvenilestages (e.g., C. gar-
ciae and C. velutinella).In some species (C. albicans
and C. strigosa), the terminalbuds are more or less
inflated. In slow-growing species the stipules often
enclose (and protect)the inflorescencesuntil nearan-
thesis; and in C. hololeuca, in which the inflores-
cences lack spathes, they are enclosed until anthesis.
In general, the durationof the protection of young
leaves and inflorescences is greater in species with
longer stipules.
INFLORESCENCES
There are usually two inflorescencesper leaf axil
with the lateralbud between them (Fig. 3). One of the
two inflorescencesmay be abortive.
In many species, each of the inflorescencesis sub-
tendedby a basal bract,which is probablya prophyll.
These bractsare sometimes insertedon the peduncle
(e.g., in Cecropiaalbicans). They vary in lengthfrom
ca. 0.5 to 8 cm and usually have the same textureand
indumentumas the stipules. They are mostly cadu-
cous and, therefore,not present in herbariummate-
rial. There are sometimes bracts at the bases of the
spikes;they are small butoccasionallygrow to 2.5 cm
long.
The inflorescenceconsists of a common peduncle
bearing a number of spikes (or spadices) initially
completely enclosed by a spathe(Fig. 3). At anthesis,
the spatheopens and drops.Cecropiahololeuca is the
only species without a spathe. Here, the spikes are
subtendedby one or two free bracts, to 8 cm long.
They do not enclose the spikes. In this species, the
spikes are enveloped by the terminal bud cover
(formedby the stipules) until anthesis.
The spathe is usually somewhat longer than the
longest spikes just before anthesis. Sometimes the
spatheis shorterand then the spikes are curved,crin-
kled, or coiled, but if straightthen piercing the apex
of the spathe.The spatheopens adaxiallybeforedrop-
ping. In staminate inflorescences of Cecropia stri-
MORPHOLOGY
1 Z
FIG. 3. Structureof the inflorescence of Cecropia,
schematic.1. Pair of inflorescenceswith the axillary bud in
betweenand subtendedby basal bracts,the spikes fully en-
veloped by a spathe, reduced/modifiedflowers in between
the (stipes of the) spikes; 2. Spike with flowers and hairs, at
the base of the spike a modified flower.
peduncle
primary
FIG. 4. Basic structureof the inflorescence of Cecropiaceae, as still largely present in Myrianthus,schematic.
(Redrawnfrom Bull. Jard.Bot. Etat 46: 475. 1976.)
15
gosa, the spathe may remain hanging on the one or
two longest spikes. The spathesare similarto the stip-
ules in texture,color, and indumentum.
The spikes are sessile or stipitate;the stipe (or the
pedicel or peduncle of some authors)is the basal part
of the rachis.The flowers aredensely set on the rachis
and not intermixedwith interfloralbracts.The rachis
is often hairy,mostly with shortstiff hairs,sometimes
also with soft crinkled hairs. If perianthsare basally
fused, then stiff short hairs, similar to those on the
rachis,are found on the perianthsjust above the fused
parts.
The number of spikes of the pistillate inflores-
cence is usually four but may be one (Cecropia ga-
brielis) or as many as 20 (C. garciae and C. hispidis-
sima).
The staminate inflorescences usually have more
spikes, even up to 50 spikes in, e.g., Cecropia mar-
ginalis and C. membranacea),or up to 100 in C. her-
thae. In some species, e.g., C. palmata and C. putu-
mayonis,four spikes arecommon. Inflorescenceswith
less than four spikes are occasionally found.
The numberof spikes of both staminateand pis-
tillate inflorescences usually varies more or less
within a species, more in staminate inflorescences
than in pistillateones. The firstformedinflorescences
on the plant and the branches tend to be relatively
small and often have fewer and shorterspikes thanthe
succeeding ones. Poorly developed trees also tend to
n/
n-i
n-2~
branches
dichotomous
secondary ramification
16
have shorterand fewer spikes than vigorously grow-
ing trees. Independentof the variationrelated to the
stage or state of developmentof trees, the length of
the peduncle, the length and the number of spikes,
and/or the length of the stipes, may also vary from
one partof the species rangeof distributionto another.
In Cecropia obtusifolia, specimens with conspicu-
ously differentspikes are found side by side.
The diameterof the spikes may vary considerably,
especially the staminateones, partlyin relationto the
numberof spikes per inflorescence.Usually the more
spikes present,the thinnerthey are. Spikes, in partic-
ular pistillate ones, are sometimes (dichotomously)
branched.The spikes of the pistillate inflorescences
of Cecropiagoudotianaare two by two basally fused.
In both staminateandpistillateinflorescences,sol-
itary flowers are presentat or among the bases of the
spikes (Fig. 3). They are more conspicuous in pistil-
late inflorescencesthanin staminateones. These flow-
ers are often more or less reduced and modified, the
pistillate ones often with more or less clearly cup-
shaped perianth containing some fluid (nectar?,cf.
Andrade, 1984c). The position of these flowers is of-
ten irregular,however, with a tendency to occur in
between two spikes. Moreover,minuteswollen bracts
may occur, often accompanyingthe solitary flowers.
The length of the peduncle, stipe, and spike varies
considerablywithin the genus, and may vary in spe-
cies as well. Dependingon the length and the rigidity
of the peduncle, the stipe, and the rachis, the stami-
nate inflorescence may have an erect peduncle and
erect spikes, an erect peduncleand pendulousspikes,
or a pendulouspeduncleand (thus)pendulousspikes.
In the pistillate inflorescences, the peduncle is pen-
dulous and with pendulous spikes, or it is erect with
erect spikes. In the lattercase the peduncleand spikes
may still be erect when fruiting or they may curve
downwardwhen fruiting.The length of peduncleand
spikes can, to some extent, be related to pollination
(shedding of pollen) and dispersal (the group of ani-
mals involved). Staminate inflorescences with long
peduncles and/or long spikes can be more easily
moved by the wind, promotingthe release of pollen.
Long and pendulous pistillate inflorescencesin fruit
are associated with dispersal by bats. Long to very
long peduncles, in particularin pistillate inflores-
cences, aremore commonin lowlandthanin montane
species.
The structureof the inflorescencein Cecropiamay
be explained from a presumablybasic type of inflo-
rescence of Cecropiaceaeas found in the African ge-
nus Myrianthus(de Ruiter, 1976). This basic type is
cymose and preservedin staminateinflorescencesof
FLORANEOTROPICA
some Myrianthusspecies (Fig. 4). It has a peduncle
with aboutfour primarybranchesat the apex. Further
ramificationof these branches is usually repeatedly
dichotomous.In a morereducedstate,as can be found
in some Myrianthusspecies and in many Coussapoa
species (Akkermans& Berg, 1982; Berg et al., 1990),
the ramificationmay be(come) dichotomousfromthe
peduncle on. In the basic type of inflorescence, the
flowers are concentratedon the distal branchlets(or
only on the ultimate branchlets), thus forming
branchedspikes (or unbranchedones). Solitaryflow-
ers can be found at the top of the branches,i.e., at the
bases of the dichotomies.
One could explain the structureof the inflores-
cence of Cecropia by assuming a shorteningof the
proximalbranchesof the basic type of inflorescence
and the elongation(and reductionof branching)of its
ultimate flower-bearing parts. The most common
numberof spikes (four), more or less distinctpairsof
(pistillate) spikes, occasional (dichotomous) ramifi-
cation of spikes, and the most common position of
solitary flowers can all be regardedas indicationsfor
a basically dichotomousramification.
In many Cecropiaceae the inflorescences are
enclosed in the terminal bud cover (formed by the
stipules) until anthesis or just prior to anthesis. The
developmentof a spathe fully enclosing the flower-
bearing parts created a substitutionfor the original
way of protection.This could be relatedto continuous
rapidgrowthof the stems, too rapidto allow full de-
velopment of inflorescences within the bud covers,
and to the need of more or less continuousproduction
of flowers and diaspores. Cecropia hololeuca is the
only species in which the inflorescencesare still pro-
tected in the original way. The independenceof the
developmentof inflorescencesinside the terminalbud
cover allows long peduncles and/orspikes.
SPIKES OF STAMINATE INFLORESCENCES
The rachis of the spike is terete or, more often,
more or less angularand mostly triangularin cross
section. It is mostly hairy,usuallyonly with shortstiff
hairs, sometimes also with arachnoidindumentum.
The rachiscontainsmostly tannin,often concentrated
in its outercell layers.
The spikes are mostly pale yellow to whitish at
anthesis. However, they are green in Cecropia chlo-
rostachyaand occasionally darkyellow to orange or
even almost scarletin C. polystachya.
STAMINATE FLOWERS
The flowers are sessile or (shortly) pedicellate.
They are free in all species, except for Cecropiapur-
MORPHOLOGY
purascens, in which the upperparts of the perianths
are connate. In C. heterochroma,the more or less
fleshy apices of the perianths are interlocking and
close off the slits among the flowers.In the otherspe-
cies the perianthapices are free from each other.The
length of the flowers varies, partlyin connectionwith
their position on the rachis. They are relativelyshort
at the apex and the base, and at sides of an angular
rachis. The flowers on the angles tend to have longer
perianthsand have pedicels or longer pedicels than
those on the sides. The flowers are 2-merous, very
rarely 3-merous. The perianthis tubular,althoughin
some species, as C. marginalis, the tubularpart is
short. In most species the tepals are almost entirely
connate, leaving only a narrowslit-shaped aperture.
But in C. marginalis the tepals are partly or alsmost
entirely free. The apical part of the perianthis often
more or less thickened.It is hollow in severalspecies
and often also contains (some) tannin.In contrastto
the perianthof the pistillate flower, the apex of the
perianthis mostly smoothand the partbelow the apex
is often glabrous, or if hairy, then mostly with stiff
short hairs or, if arachnoidindumentumis present,
then it is mostly sparseand short.However,in a small
numberof species (as in those with flowers of the C.
sciadophylla type, see below) the arachnoid indu-
mentumbelow the apex is similar to that of the pis-
tillate flower.
The staminateflowersof Cecropiashow morevar-
iation than the pistillate flowers. This variation is
largely related to the stamens. The anthers are in-
trorse,and the filamentsare flat or more or less thick-
ened. In most species the filamentsdiffer in length in
the bud and then elongate one afterthe other,pushing
the anthersthroughthe slit-shapedaperture(Fig. 5. 1).
The anthersmay remainattachedto the filament,but
more often they become detached and subsequently
reattachedto the marginsof the apertureof the pern-
anthor remainconnectedwith the filamentin one way
or another.The mechanicalaspect of the detachment
of the anthersis not quite clear. It can be explained
by contractionof the filamentafter it has pushed the
antherthroughthe aperture,which is narrowerthan
the anther.The filamentsare usuallydistinctlyshorter
than the tube of the perianthafter detachmentof the
anthers.
Several types of staminateflowers are found. But
the state of the material makes it often difficult to
determinefor each species the type of flower, either
because the material in a dried state often does not
show clearly the features of the stamens and anthers
or because the flowers are not in the right state of
development.One needs materialat the right phases
17
of development conserved in liquid. Moreover,two
types of staminateflower may occur in the same spe-
cies, sometimes in the same area, sometimes geo-
graphicallyseparated.The following types of stami-
nate flowers can be recognized.
1. Cecropiapeltata-type(Fig. 5.1-4). The anthers
bear mostly ca. 0.1 mm subulate appendagesat the
bases on the thecae in between the locules. The an-
thersareca. 0.5-1 mm long. The perianthsaretubular
and mostly ca. 1-2 mm long, glabrous or sparsely
hairy outside below the apex. The apex is plane to
slightly convex. The filaments are relatively broad,
flat or slightly thickened. The anthers become de-
tachedfromthe filamentandreattachedto the margins
of the slit-shapedapertureby the (sticky) tips of the
appendages.The appendagesbecome thread-like,and
thus the anthers loosely attached to the flower and
movable by wind, so that pollen can be shed. The
antherwhich is pushed first throughthe apertureis
often removed by the second anther. This type is
found in the majorityof the species.
2. Cecropia latiloba-type(Fig. 5.10-12). The an-
thers are appendiculateor not. At anthesis, they are
detachedfrom the middle of the filamentbut remain
laterally loosely connected to the filament by a pair
of filiform connections between the uppermarginof
the filamentand the V-shapedbase of the connective
of the anther.The perianthis tubularwith a planeapex
and an aperturethat opens somewhat more widely
than in the C. peltata type. The perianthis relatively
short, ca. 1-1.5 mm long, and bears short stiff hairs
outside below the apex. The anthersare 0.6-1.2 mm
long. The filamentsareoften somewhatnarrowerthan
those of the C.peltata-type.The anthersareso loosely
attachedthat they can be moved by the wind. After
the antherbecomes fully detached, the filiform con-
nections remainon the uppermarginof the filaments.
It is not clearhow the antherbecomes finallydetached
from the filament.This type of flower is found in C.
albicans, C. latiloba, and C. utcubambana.
3. Cecropia sciadophylla-type (Fig. 5.5-6). The
anthersare relatively long, 1-2 mm, and are appen-
diculate or not. After detachmentof the antherthey
remain (temporarily)loosely attached by stretched
spiralthickeningsof the tracheidsof the (median)vas-
cular bundle of the filament. The perianth is often
relatively long, 1.5-4 mm, and it bears more or less
dense arachnoidindumentumbelow the apex outside,
as typical for the perianthof the pistillateflower.The
apex is distinctlyconvex to plane with an narrowap-
erture.This type of staminateflower is found in C.
glaziovii, C. palmata, C. saxicola, and C. sciado-
phylla. However, in the former two species, the C.
18 FLORA NEOTROPICA

LI~~~~~~~LL

5 6 8 9

10 12 13 lL4
FIG. 5. Staminateflowers of Cecropia; various types, more or less schematic. C. peltata type. 1. Perianthand two
stamens, the filaments unequal in length. 2. Stamen before anthesis, the thecae appendiculate.3. Anther detached and
reattachedto the margin of the apertureof the perianth with the appendages.4. Filament and detached anther with the
appendagedfiliform.C. sciadophylla type. 5. Perianthand detachedanther,remainingconnectedto the filamentby stretched
spiral-thickeningsof tracheids.6. Stamen with the antherconnected to the filamentby spiral-thickenings.C. heterochroma
type. 7. Perianthand stamens of which the anthersremainattachedat anthesis by the membranousapex of the filament.8.
Antherstill attachedto the filament.9. Antherdetachedafteranthesis,the marginsof the apicalpartof the filamentremaining
as threadsat the marginof the lower partof the filament.C. latiloba type. 10. Perianthand detachedantherat anthesis. 11.
Anther remaining attached to the filament by filiform connections between the connective and the upper margin of the
filament. 12. Anther fully detachedafter anthesis. C. membranacea type. 13. Perianthand stamens remainingattachedto
the thick filaments. 14. Stamen.
MORPHOLOGY 19

peltata-type of stamens is also found. The type de-
scribed is found in thick spikes with relatively long
flowers and anthers, the other type in more slender
spikes with shorterflowers and anthers.
4. Cecropia heterochroma-type(Fig. 5.7-9). The
anthersare pushed throughthe aperture(which is of-
ten slightly wider thanin the C. peltata-type)but they
remainattachedto the filamentby its thin (membran-
aceous) upperpart.The anthersare appendiculateor
not. The perianthis of the normallength,with a plane
or slightly convex apex, and it is hairy or glabrous
below the apex outside. The filaments are thick and
before anthesis differentin length. After anthesisthe
filaments fill the space inside the perianth and the
apices of the filamentsreach or even exceed the ap-
erture.The anthersmay subsequentlyremainattached
to the filamentonly by two filiform connectionsrun-
ning from the upper margin of the filament to the
connective of the anthers, like in flowers of the C.
latiloba-type. This type of staminateflower is found
in C. angustifolia, C. heterochroma,C. polystachya,
and C. tacuna.
5. Cecropia membranacea-type (Fig. 5.13-14).
The anthersremainattachedto the filamentat anthe-
sis. The perianthis short, with a wide aperture,and
it is glabrous.The anthersare short,0.5-0.7 mm long,
and are not appendiculate.The filamentsarerelatively
narrow,(rather)thick, (almost) equally long before
anthesis, and exceed the apertureafter anthesis. The
spikes are very slender and pendulous and thus are
easily moved by wind to get pollen releasedfrom the
anthers.This type of staminateflower is found in C.
marginalis, C. membranacea,and C. strigosa.
One could regard the C. membranacea-typeof
flower as primitive. However, this is not correlated
with primitivecharactersin vegetativeparts.Not only
for this type but also for the others,correlationswith
vegetativecharactersarenot evident.Similarityin the
type of staminateflower appearsnot always to be an
indicationof relationship.
The partial detachment of anthers, such that
they remain loosely attached to the filament by
stretchedspiral thickenings, was discovered and de-
scribed by Allemao (1860). The complete detach-
ment of antherswhich become secondarily attached
to the flower by appendagesof the thecae was dis-
covered and described by Berg (1977b), who then
also regardedthe case of attachmentof the anthers
by spiral thickeningsas new and undescribed.How-
ever, he misinterpretedthe loose attachmentof the
anthers by filiform connections between filaments
and anthersas anthersremainingattachedto the up-
per margin of the filamentby the stretchedappend-
ages of the thecae.
Complete or partialdetachmentof the anthers,by
which they become loosely attachedto the flower,are
clearlyadaptationsto wind pollination.In cases where
the anthersremainfirmlyattachedto the filaments,as
in Cecropiamembranacea,the spikes arevery slender
and pendulousand are easily moved by the wind. In
othercases, too, slenderand/orlong pendulousspikes
may promoterelease of pollen.
The staminateinflorescences(flowers)of many(or
all?) species emit a (faint) sweetish smell; for some
species, e.g., Cecropiastrigosa, it is rathercharacter-
istic.

Key to the types of stamen

1. Antherscompletely detachedfrom the filamentsand reattachedto the marginsof the apertureof the
perianthby sticky appendagesof the anther................................................................... 1. C. peltata-type
1. Anthersremainingconnected to the filaments.
2. Anthersremainingloosely connected to the filaments.
3. Anthersremainingconnected by stretchedspiralthickeningsof tracheids.......... ............ 3. C. sciadophylla-type
3. Anthersremainingconnectedby two filiformconnectionsbetween the connective and upper
marginof the filament.................................................................. 2. C. latiloba-type
2. Anthersremainingfully connected to the filaments.
4. Anthersremainingattachedby a membranaceousapex of the filament:stamensdifferentin length
before; perianthand antherof normallength; appertureof the perianthslit-shaped..... 4. C. heterochroma-type
4. Anthersremainingattachedwithout a membranaceousapex of the filament:stamensabout equal
in length before anthesis;perianthand anthershorterthan normal;apertureof the perianth
wide ........................................................... 5. C. membranacea-type

POLLEN (1974). Cecropia pollen found in sediments play a

The pollen is dry. Clouds of pollen are released role in palaeoecological studies revealingchanges in
from the spikes when they are moved or touched.Pol- vegetation and climate (Wijmstra,1967; Liu & Col-
len of some species has been decribed by Barth invaux, 1985, 1988; Bush & Colinvaux, 1988; Col-
20
invaux et al., 1988a, 1988b; Bush et al., 1990; Col-
invaux, 1993; Wijninga, 1996) and in relation to
agriculturalhistory (Bush et al., 1989).
SPIKES OF PISTILLATEINFLORESCENCES
The spikes are mostly terete,less commonly more
or less angularthan in the staminateones. The rachis
is mostly hairy, with only stiff straighthairs, some-
times also with (or rarelyonly with) arachnoidindu-
mentum.Tanninis mostly present,often concentrated
in the outercell layers;but in severalspecies, such as
many of the Cecropia peltata-group, the outermost
layers are free from tannin. The spikes are normally
whitish until the spathe drops, but they soon turn
green.
PISTILLATEFLOWERS
Most of the variation in the pistillate flowers is
presentedin Figure 6. The flowers are sessile, mostly
free, but in several species of the Cecropiapeltata-
group, such as C. litoralis and C. peltata, they are or
may be basally connate. If so, the stiff hairs com-
monly arising from the rachis are situated on junc-
tions of the adjacentperianths.The perianthis tubu-
lar; the tepals are fused entirely. The length of the
perianth is shorter at the base and the apex of the
spike. The perianthsare usually about 1 mm longer
in the fruitingstate than at anthesis.The upperpartis
more or less stronglythickenedand the apical partis
mostly more or less convex and is often muriculate
to punctate or hispidulous. In all species arachnoid
indumentumis presenton the outside of the perianth,
mostly only below the apical part, but in species of
the C. peltata-group,usually also on the apex if the
stigma is peltate (Fig. 6.4). The arachnoidindumen-
tum may occurjust below the apex or extend farther
downward,sometimes even to near the base. In some
species stiff shorthairs may also occur below the ap-
ical partoutside. Arachnoidindumentumis often also
found inside or just below the style channel.In many
species this indumentumappearsto be always pres-
ent, and in others it may be sometimes absent or al-
ways absent (as in several species of the C. peltata-
group). The aperture,which is just large enough to
allow the style (or stigma) through,is slit-shapedor
(almost) circular.The style channel in the thickened
apical partof the perianthvaries in length in relation
to the length of the apical part.The style also varies
in length; long styles are often curved, and they may
bear short hairs. The style is attachedto the ovary
apically or subapically.The stigma varies from pen-
icillate to tongue-shapedto peltate. In species (or in-
dividuals)of the C. peltata-groupwith distinctlypel-
tate stigmas, the arachnoidindumentumextends to
FLORANEOTROPICA
near the apertureof the perianth.In these species (or
individuals)the stigma is alreadyoutside the aperture
long before anthesis, whereas otherwise the stigma
becomes exposed just before anthesis. The presence
of arachnoidindumentumon the apical part to near
the apertureis apparentlycorrelated with a peltate
stigma and a differencein the ontogeny of the flower.
In the C. peltata-groupone can find transitionsfrom
the commonly peltate stigma throughthe subpeltate
to tongue-shapedstigma to the penicillate one, and
these are relatedto extension of arachnoidindumen-
tum from below the apical partof the perianthtoward
the apertureof the perianth(Fig. 6). Tanninis com-
mon in the perianthand is often concentratedin the
upperthickenedpart. However, in some species it is
scantyor in the species of the C.peltata-groupusually
absent.Tanninoften accumulatesin the pericarp,but
it can be totally absent, as in C. longipes.
FRUITS AND SEEDS
The fruitsare achenesenvelopedby somewhaten-
larged, at least in the upperpart, more or less fleshy
perianths,remaininggreenisheven at full maturityof
the infructescence.The fruits are oblongoid, elliptic,
(sub)obovoidor (sub)ovoid. The pericarpis more or
less tuberculatein most species, but in several it is
smooth. The color of the pericarpvaries fromdarkto
pale brown. The testa is very thin. The straightem-
bryo with a ratherlong radicleand equal and flat cot-
yledons is embedded in endosperm. Under certain
conditionsthe viabilityof Cecropiaseeds can be more
than five years. Germinationis triggeredby full sun-
light and changing temperatures (Holthuijzen &
Boerboom, 1982; Vaizquez-Yanes& Smith, 1982).
NATURE AND VARIATION OF
MORPHOLOGICALCHARACTERS
The general picture of morphologicaldifferentia-
tion in the genus is a mixtureof uniformity,as in the
generalhabit and in the pistillate flowers, and of con-
siderable variation, as in leaf charactersand in the
dimensions and numbers of spikes in the inflores-
cence. Partof the variationin leaf characters,such as
the numberand depth of the incisions, the numberof
lateral veins in the midsegment of the lamina, and
their loop-connections before or in the margin, are
clearly relatedto the considerablechange fromjuve-
nile to adult features, often without reaching a state
of stability in features. The variationin the number
of spikes can be related to the basically open inflo-
rescence in which the numberof branchesis not def-
inite and may vary with vigor of growth. Whereas
some of the variationappearsto be relatedto ontog-
eny, variationin otherrespects-such as the length of
ANATOMY 21

Fi(;. 6. Pistillate flowers of Cecropia, different types with regardto stigmas and the presence of arachnoidindu-
mentum.schematic. 1. From C. latiloba. 2. From C. ulei. 3. From C. sciadophY/l1a.4. From C. peltata.

stipules,the composition of the indumentum,and the of parentalcharactershave not been encounteredby

absence,presence,and state of the trichilia-is not or the authors.However, individualsshowing one (or at
is only partlyso. most two) features-e.g., of the indumentum,of a co-
About 50% of the species are morphologically occurring species-are not uncommon (see, e.g.,
uniform;most of them have more or less small ranges comments underCecropiagabrielis and C. metensis).
of distributionandmoreor less narrowecological am- A proof of such results in hybridizationwas found at
plitudes. The more or less conspicuous variation in the campus of the Universidade Federal de Minas
the other species is partly (geographically)clinal; it Gerais (Belo Horizonte, Brazil). A single fruiting
can be partly related to ecological conditions; or it spike from a specimenof C.polystachyain cultivation
cannotbe linked to either distributionor ecology. in Campinas(Sao Paulo, Brazil)was takenby J. Lom-
A remarkable type of variation is the co- bardi(BHCB) and from the seeds severalplants were
occurrenceof color variants in different proportions raisedand four of them were planted.They were adult
and of variantsin the length of spikes (in Cecropia specimens in March2001. Threeof them show all the
obtusifolia). featuresof C. polystachva,but the fourth,a staminate
There are few exceptional characterssuch as the one, almost fully the featuresof the parentalspecies
absenceof a well-developedspathein Cecropiaholo- C. pachystachva.The latterdiffers from other mate-
leuca, trichilia enclosed in "pockets"in C. virgusa, rial of C. pachystachya (indigenous in Campinas)in
connate staminate flowers in C. purpurascens, and the pronouncedlobation of the free part of the seg-
distinctlylargerand pink Mullerianbodies in C. his- ments of the laminaretainedin the adult state. In this
pidissima. There are some charactersshared by few respect, that specimen resembles C. polvstachva.
species which probably indicate taxonomic affinity, Othercharacters,such as numbersor dimensions ex-
such as long irritatinghairs thateasily breakoff in C. ceeding the "normal"rangesof the species, mightalso
garciae, C. hi.spidissima, and C. megastachya, a be linked to hybridization.
group of species showing affinities also in other fea-
tures. However,the presenceof anthersremainingat-
tachedto the filamentsby spiral thickeningsis found ANATOMY
in the clearly relatedspecies C. glaziovii and C. pal- Comparativeanatomicalstudies on the wood and/
mata, but also in C. putumavonis,C. saxatilis, and C. or the leaves, including the genera of the Cecropi-
sciadophvlla, which do not show affinities to each aceae, have been carriedout by Renner( 1907) and by
other or to the formerset of species. Bonsen & ter Welle (I1983).The wood anatomyof the
Hybridizationmay occur ratherfrequently(S. T. neotropical genera was treated by ter Welle et al.
Trimble,pers. comm.). Hybrids with a clear mixture (1992). The anatomyof the leaves of some Cecropia
22
species has been described by Richter (1897). The
idioblasts in the leaves of Cecropiaceae have been
studied by Setoguchi et al. (1993). Members of the
family accumulatesilicon in contrast,butin otherUrt-
icales, abundantcalcium and silicon.
PROTECTION OF FLOWERS
AND INFLORESCENCES
Several featuresof the flowers and inflorescences
probably protect the inflorescences and flowers
againstpenetrationand subsequentdamageby the lar-
vae of insects (cf. Berg, 1990).
Insect larvae are usually present in the spikes of
staminate inflorescences after they have shed the
spathe, but they are absent in the pistillate spikes of
pistillate inflorescences. Andrade (1984a) presented
an account of a species of Ophtalmoborus(Curcu-
lionidae, the larvae of which are found in staminate
inflorescencesof C. pachystachya (= C. lyratiloba).
These larvaedestroy the rachises of the spikes.
Insects (adultsand/orlarvae)can always penetrate
the spikes of staminateinflorescencesthroughthe ap-
erturesof the perianthsand in most species through
the slits among the flowers as well. The slits among
the flowers can be closed off by fusion of the upper
parts of the perianths(in Cecropiapurpurascens)or
by dense, entangledarachnoidindumentumat the lat-
eral surfacesof the perianthsas in some otherspecies
(e.g., C. palmata and C. sciadophylla). A third way
to close slits among the staminateflowers is found in
C. heterochroma,in which the apices of the perianth
interlock.
The perianths of pistillate flowers always bear
long arachnoidindumentumoutside, at least below
the apex. This indumentumis interwovenwith thatof
adjacent flowers and close slits among the flowers.
Penetrationthroughthe apertureof the perianthis pre-
vented by the small diameterof the aperture,just let-
ting throughthe style. Moreover,shortbristles on the
style and the arachnoidindumentumat the inner sur-
face of the perianth,in orjust below the style channel,
will contributeto barringaccess to the interiorof the
flower.In addition,tanninis usually more abundantly
present in the pistillate flowers than in the staminate
ones; it is concentratedin the thickenedapicalpartof
the perianthand in the wall of the ovary and devel-
oping pericarp, and it provides additionalchemical
protectionagainst herbivory(cf. Coley, 1986). These
features probablyexplain the general absence of in-
sect larvaein the spikes of the pistillateinflorescence.
Before anthesis the spikes are not only protected
by the spathe, but initially also in the terminalbud.
FLORA NEOTROPICA
The durationof this protectiondependson the length
of stipules and the length of the inflorescence.In Ce-
cropia hololeuca, which lacks a well-developed
spathe, the protectionof the spikes until anthesis is
carried out only by the stipules. Protectionof inflo-
rescences by the terminalbud is a common featureof
Cecropiaceae.
PHENOLOGY
Most species flower and bear fruit throughoutthe
year,but individualsoften intermittently.In some spe-
cies (e.g., Cecropiaelongata), flowering and fruiting
appearsto be season-dependenteither throughoutor
in partof their ranges of distribution,as is found for
C. obtusa in populations in southeru Para (Brazil),
which bear neitherflowers nor fruitsat the end of the
dry season. According to field notes and data on col-
lection labels, C. palmata and C. sciadophyllaappear
to flower throughoutthe year, but in French Guiana
the fruiting is restrictedto the period October-Janu-
ary (the end of the dry season and the beginning of
the rainy season) (Charles-Dominique,1986). These
data suggest that flowering and fruitingperiods vary
throughouttheir ranges. Milton (1991) reportedsea-
sonal fruiting(December-July)for C. insignis in Pan-
ama. Seasonal floweringis clear in C. latiloba and C.
membranacea,both inhabitantsof seasonally flooded
(riverside) vegetation. These species (at least in the
middle partof the Amazon basin, e.g., nearManaus),
flowerwhen the waterlevel is high and fruitwhen the
water level is going down and river banks become
availablefor seedling establishment.
POLLINATION
Several traits of the staminateflowers and inflo-
rescences described and field observationsindicate
that Cecropia is in general adaptedto wind pollina-
tion, eitherby pendulousspikes which can be moved
by wind to shed the pollen or by the special adaptation
of detachmentof anthersand their secondaryattach-
ment allowing the shedding of pollen by motion of
the anthers.The pollen is dry and easily released by
movement. It is likely that wind pollination is pre-
dominant.Although observationsof pollen transport
by insects are wanting,small beetles and flies use sta-
minate inflorescencesas breedingsites (cf. Andrade,
1984a;Wheeler, 1942) and thus may play a small role
in pollination. Insects (Epitragussp. and Ophtalmo-
borus sp.) eat pollen grainsof C. pachystachya(= C.
lyratiloba) but probably do not pollinate (Andrade,
1984a). In the same Cecropia species pistillate flow-
ers producesmall amountsof nectar(Andrade,1980,
ORGANISMSOTHERTHAN ANTS ASSOCIATEDWITH CECROPIA 23

1984c), probably to catch pollen and not to attract tions by the first authorbut on other evidence, such
insects. as label data and perforatedprostomatain herbarium
material.
2. Species inhabited by Azteca ants, but not
DISPERSAL
throughouttheir geographic ranges; either trichilia
In many species of Cecropia,full-growntrees can (with Mullerianbodies) absentor internodeswith am-
produce millions of seeds (Brokaw, 1998; Martinez- ple pith: Cecropia albicans, C. angustifolia, C. pel-
Ramos & Alvarez-Buylla, 1986). The abundanceand tata, and C. telenitida.
the regularpresenceof fruitspermitsthe genus to play 3. Species inhabitedby other genera of ants; tri-
a role as keystonefood supplyfor frugivorousanimals chilia with Mullerianbodies present,internodeswith
(Vepsalainen, 1999). The diaspores are dispersed scarcepith: always Cecropiahispidissimaand C. me-
mainly by animals:various birds, including insectiv- gastachya, sometimes C. membranacea.
orous ones (Eiseman, 1961; Leck, 1972; Snow & 4. Species never inhabitedby Azteca ants; trichi-
Snow, 1971; Charles-Dominique,1986; Vepsalainen, lia absentor poorly developedand/orinternodesfilled
1999) and toucans(Skutch, 1971);fruitbats (Charles- with ample pith: Cecropiaandina, C. bullata, C. ga-
Dominique, 1986; Fleming, 1988; Fleming & Wil- brielis, C. hololeuca, C. pittieri, C. schreberiana,C.
liams, 1990; Lobovaet al., 2003); monkeys(Eiseman, sciadophylla, C. tacuna, and C. telealba.
1961; Terborgh, 1983; Stevenson et al., 2000); op- The absenceof antsin Cecropiachlorostachyaand
posums (Eiseman, 1961; Medellin, 1994); and fish the presence in C. velutinellais not certain,because
(Gottsberger,1978). They create or continuouslyre- of the few specimens examined.The dense indumen-
plenish the seed banks. tum on the leafy twigs and subpersistentstipules in
Pendulous infructescences with long peduncles these species may prevent or reduce occupation by
and spikes are associated with dispersalby bats, and ants (cf. Davidson et al., 1991).
those with short infructescences with dispersal by In other species, internodesfilled with mucilage
birds. Species occurring along rivers may be dis- or internodeswith thick walls and smooth leafy twigs
persed by water, because parts of matureinfructesc- with a waxy layer (cf. Federle et al., 1997) may pre-
ences drop from the rachis and float for a while, kept vent (or reduce)occupationby ants (or select species)
buoyant by the air in the arachnoid indumentum and attacksby leaf-cutterants (Schimper, 1888). As
among the perianths(see Kubitzki& Ziburski,1994). a waxy surface of the leafy twig and dense villous
indumentumcan be found in the same species, also
SURVEY OF MYRMECOPHYTISM the type of indumentummay deter ants (and other
IN THE GENUS insects).
Specimens of Cecropia species which are nor-
For a thoroughdiscussion of ants and Cecropia,
mally occupied by Azteca ants may experience the
see "Cecropiaand Its Biotic Defenses" by Davidson
absence of these ants or be inhabitedby ants of var-
(this volume, p. 214).
ious other genera (see Wheeler, 1942).
1. Species usually (or often) and throughouttheir
geographicranges inhabitedby Azteca ants; trichilia
with Mullerian bodies present and internodes (usu- ORGANISMS OTHER THAN ANTS
ally) with scarce pith: Cecropia angulata*, C. annu-
ASSOCIATED WITH CECROPIA
lata, C. concolor, C distachya, C. elongata, C. en-
gleriana, C. ficifolia, C. garciae, C. glaziovii, C. Cecropia leaves are favorite food for sloths, ani-
goudotiana, C. granvilleana, C. herthae, C. hetero- mals which can often be observed in Cecropia trees
chroma, C. idroboi*, C. insignis, C. kavanayensis*, (as in C. polystachya, Kemper& Satore, 2000).
C. latiloba, C. litoralis, C. longipes, C. marginalis, An extensive list of animals found on Cecropia
C. maxima,C. membranacea(p.p.;see below), C. me- was presentedby Wheeler(1942). Attentionhas been
C.
tensis, C. montana, C. multisecta, mutisiana, C. drawn to associationof groupsof beetles and species
obtusa, C. obtusifolia, C. pachystachya, C. palmata, of Cecropiaby Andrade(1984a) and Jolivet(1990b).
C. pastasana, C. plicata (often), C. polystachya, C. The latter authoralso drew attentionto the only oc-
purpurascens, C. putumayonis,C. reticulata, C. sa- casional presence of hemiparasiticplants (Lorantha-
rarensis,C. saxatilis*, C. silvae*, C. strigosa, C. sub- ceae) on Cecropia and the possible role of ants to
integra, C. ulei, C. utcubambana,and C. virgusa (44 explain this phenomenon. It is remarkablethat the
species in total). Species with an asterisk (*) were presence of Loranthaceaeis quite rare, but if Loran-
assigned to this category based not on field observa- thaceaeare present,then very locally and abundantly.
24
Hollow internodes often contain scale insects, Coc-
cidae (cf. Wheeler, 1942; Davidson, this volume),
tended by the ants. The coccids are usually attached
to the "ceilings"of the compartmentsof the stem, and
young ones are moved to the next internode,mostly
via the holes made in the nodal septa. The inhabiting
ants obtain sugary exudates from these insects.
ECOLOGY
Cecropia is one of the majorpioneer tree genera
in regions of the Neotropics with more or less wet
lowland and montane forest. There is a high degree
of specializationto certaintypes of habitats,like mon-
tane ratherthan lowland or non-inundatedratherthan
inundatedhabitats.Within the major ecologic cate-
gories species show more definite (fine-tuned)pref-
erences to elevations, duration of inundation, and
otherconditions.Clearecological variationis not un-
common within species.
The ecological aspect in the differentiationand
speciation appearsto be more pronouncedin Cecro-
pia than in the other genera of Cecropiaceae,as well
as in those of Moraceae.
Most of the Cecropia species are lowland species
occurringat elevationsup to 1000(-1300) m. Species
that can be described as submontaneare found at
(700-)1000-1800(-2000) m, and the montane ones
in cloud forest at 1600-2600 m.
The elevational amplitudes are narrow and/or
well-defined for many species (as Cecropiaalbicans
and C. megastachya)or wide and/orwith unclearlim-
its for others (as C. angustifolia). Some predomi-
nantly lowland species, e.g., C. garciae and C. his-
pidissima, extend to (or possibly occur disjunctlyin)
submontanehabitats(see below) and the montaneC.
maximais found in submontanehabitats.
Most of the species are associated with non-
inundated vegetation. Some species, Cecropia lati-
loba and C. membranacea,are clearly elements of
seasonally inundated riverside vegetation (cf. La-
motte, 1990, 1992). These two species of medium-
sized trees occur abundantlyin riversidehabitatswith
periodic inundationof long durationand are distinct
in their requirements:C. membranaceais associated
with nutrient-richandwell-aeratedhabitatsand C. la-
tiloba with poorer and/or less well-aeratedhabitats.
The formercan form extensive stands, the latter oc-
curs often solitary or in small groups. Some other
species may occur in temporaryinundatedvegetation,
as C. metensis in Mauritia-swampsand C. litoralis
(in its Amazonian area of distribution)along black-
water rivers.
A few species are normallyfound on rocks, such
FLORA NEOTROPICA
as Cecropia granvilleana on wet rocky slopes of a
few inselbergs in French Guiana, C. goudotiana on
rocky surfaces in the dry Upper Magdalena valley
(Colombia),and C. saxicola in rocky areasin central
Brazil and Bolivia. Cecropia idroboi is probablyas-
sociated with (sandstone)rocky areasin Caquetaand
Meta (Colombia).
Several species, e.g., Cecropiadistachya, C. her-
thae, C. insignis, C. purpurascens, and C. sciado-
phylla, are species commonly found in (large)treefall
gaps in maturelowland forest (cf. Brokaw, 1986), as
well as in other naturalor man-madeprimaryclear-
ings. These species often become longlived tall trees
(some to 30-40 m) and can form part of the forest
canopy.These species, which can be describedas for-
est (gap) species, usually establishabundantlyin pri-
mary forest clearings (and can form almost pure
stands, cf. Oldeman, 1974: tabs. 68-69), but hardly
succeed in establishingor regeneratingin subsequent
(or secondary) clearing, or after use of the clearing
for agriculture.In contrastto the next category, this
group of species performspoorly, growing slowly, in
greenhouseconditions(at StoreMilde, Norway),pos-
sibly due to absence of mycorrhizalfungi (see Bro-
kaw, 1998: 95).
Others, which can be described as more or less
pronouncedly"weedy,""capoeira,"or "chagra"spe-
cies are Cecropia concolor, C. ficifolia (p.p.), C. Ii-
toralis (p.p.), C. obtusifolia, C. pachystachya,C. pel-
tata, and C. ulei. These are small or medium-sized
and more or less shortlivedtrees, rarelyestablishing
in forest gaps but common and often abundantin sec-
ondarygrowth,abandonedfields, etc.
Many species occur abundantlyin the properhab-
itats throughouttheirrangesof distribution.However,
some species, such as Cecropia annulata, C. putu-
mayonis,and C. utcubambana,are (far)less common
and occur more or less widely dispersedwithin their
ranges of distribution,and, moreover,do not clearly
behave as pioneer trees by forming dense stands in
secondarygrowth.They appearto be associatedwith
the forest habitat,establishingin small gaps or in the
margins of larger treefall gaps and are apparently
shade-tolerant.Cecropiaficifolia behaves similaryin
partof its range, includingthe foothills of the Andes.
The indicateddifferencesin ecological behavioralso
becameevidentin greenhouseexperimentswith some
species of Cecropia,indicatedas "gap"and"pioneer"
species, under different light regimes and nutrient
treatments(Folgarait& Davidson, 1994, 1995).
Thereappearto be two groupsof montane(or sub-
montane) species. One comprises species like Ce-
cropia maxima, C. tacuna, C. telealba, and C. tele-
DISTRIBUTION
nitida, which occur ratherevenly in the forest and do
not evidently show the traitsof pioneer species. They
can become tall trees with broad crowns. The other
group, as representedby C. andina and C. gabrielis,
form usually smaller trees and tend to aggregatein
more or less open sites, such as along streams. In
general,montanespecies are not evidently colonizers
of landslides,which appearsto be in accordancewith
the results of a study by Guariguata(1990) on forest
regenerationon landslidesin PuertoRico.
The categories that can be recognizedwith regard
to ecological behavior can only be circumscribed
roughly, as many species take (or can take) more or
less intermediatepositions. Moreover,they may be-
have differentlyin differentpartsof theirdistribution
ranges, as in the periphery of their ranges. Several
species have (or may have) differentiatedinto eco-
types, ecospecies, or ecological subspecies. This ap-
pears to be clear in some members of the Cecropia
peltata-group (see below). Cecropia litoralis is a
common, "weedy"species of non-inundatedlowlands
of the coastal region of Ecuador and the adjacent
coastal part of Colombia (Narifno).It also occurs in
the Amazonianpartof Ecuadorand adjacentpartsof
Colombiaand Peru.Thereit is apparentlyconfinedto
the bordersof the (more or less) black-waterrivers.
Cecropiaengleriana is a small to medium-sizedtree,
not clearly associated with riverine habitats, in the
southernpartof its range of distribution,Bolivia and
southernPeru.However,in partof the upperAmazon
basin, along the Rio Napo and the upper Amazon
River, this species is found mostly along rivers, in
habitats with short periods of inundation, as a
medium-sizedto quite tall tree.
Differences of abundanceof species in the same
locality and habitatmay be indicative of slight eco-
logical preferences. For example, Cecropia poly-
stachya and C. strigosa co-occur in the lowlands of
eastern Peru, where the former apparentlytolerates
drier climatic conditions than the other.
In the upperAmazonbasin, whereup to 10 species
of Cecropia can be found in the same area, the spe-
cies, or in some cases pairs of species, are ecologi-
cally complementary in occupation of the various
habitats.This is less clear in the Pacific lowlands.
Most of the morphological differentiation is
weakly or not at all relatedto ecological preferences.
A few charactersshow relationsto the ecological cat-
egories as indicatedabove: coriaceous leaves in true
lowland forest (canopy) species and in montanespe-
cies; absence of the full set of myrmecophiloustraits
in montanespecies; dense arachnoidindumentumon
the upper surface of the lamina in montane species;
25
and relatively short and or erect pistillate inflores-
cences in lowland forest (canopy) species and mon-
tane species. But shape of the lamina cannot be as-
sociated with any of the categories. Deeply incised
lamina with numeroussegments as well as shallowly
incised laminaswith few segmentscan be foundboth
in forest species and secondarygrowth species or in
both montaneand lowland species.
Ecological equivalentsof Cecropia in Africa are
the two species of Musanga in the Cecropiaceae(cf.
de Ruiter, 1976; Janzen & McKey, 1977). In Asia,
species of Macaranga Thouars (Euphorbiaceae)are
such equivalents. The genus comprises myrmeco-
phytes with remarkablesimilaritiesto Cecropiain ad-
aptations to and variation in myrmecophytism(see
Fiala & Maschwitz, 1992a, 1992b).
DISTRIBUTION
Forty to fifty percentof the taxa of Cecropiacan
be regardedas montaneor submontaneAndean,with
a distinctconcentrationof species in the northernpart
of the Andes, in Colombia and Ecuador.Takinginto
account the lowland taxa that reach the eastern or
westernfoothills of the Andes, the Andeanregioncan
be regardedas the centerof species richness and also
as the center of speciation. Only approximately25%
of the species occur more or less remotely from the
Andean region. The representationof lowland and
montanespecies in variouspartsof the Neotropicsas
well as the phytogeographicregions for the genus is
presentedin Figure 7.
The size of the ranges of distributionof Cecropia
species varyconsiderably.Some lowlandspecies have
large ranges-e.g., C. membranacea,C. peltata, C.
pachystachya, C. latiloba, and C. sciadophylla-oc-
curringin at least two more or less distinct phytoge-
ographic subdivisions (see below). Cecropia angus-
tifolia is the only montanespecies with a largerange:
from the coastalmountainrangein Venezuelathrough
the Andes to Bolivia, and in CentralAmerica from
Panamato Mexico, at elevations of (500-)800-2500
m. The majorityof the species have medium-sizedto
small rangesand are usuallyconfinedto a single phy-
togeographic subdivision. Some species have very
small ranges, e.g., C. pittieri, an endemic of Cocos
Island, and C. granvilleana,confined to some (two?)
small populationson slopes of granite outcrops (in-
selbergs) in FrenchGuiana.
The distributionof a few species is disjunct:Ce-
cropiamembranacea,occurringin the Amazonbasin,
the lower MagdalenaValley, the Pacific coastal re-
gion; and C. litoralis, common in the coastal region
of Ecuadorbut also occurringalong (black-water)riv-
26
30 ~ ~ ~ ~ ~ ~ ~ ~ ~~-0
2 i0~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
10~~~~~~~~~~~~~~.
10
20- 1;0 260 3~~~7
-0
00 0 02000000000 500... .- '00 0 o/
FIG. 7. Distributionof Cecropia. Dashed lines show the majorboundariesin the distribution.Numbersindicatethe
numberof species per region (boldface for lowland species, italics for montanespecies).
ers in the Amazon basin. In some other cases, more
or less segregateddistributionis relatedto habitatspe-
cialization, such as C. granvilleana, on some insel-
bergs in FrenchGuiana, C. kavanayensison isolated
table mountainsin the Guayana region, C. saxicola
in rocky areas in central Brazil and Bolivia, and C.
latiloba on relatively nutrient-poor and not well-
aeratedperiodically inundatedsoils. In a numberof
montane or submontanespecies, more or less pro-
nounced discontinuitiesin the ranges of distribution
appearto occur in spite of the presence of (presum-
ably) suitablehabitatsin the gaps. For example, in C.
telenitida, not recordedfrom the central part of the
easternslopes of the Andes in Ecuador,C. velutinella,
recordedfrom Pastaza and Morona-Santiagoin Ec-
uador and from San Martfn in Peru, C. montana,
knownfrom Putumayo(Colombia)and adjacentSuc-
umbios (Ecuador) and from Morona-Santiago(Ec-
uador)southwardto Madrede Dios (Peru). An anal-
ysis of the species' distributionthen recognized, and
with the distributiondata then known, was presented
by FrancoRosselli & Berg (1997).
FLORA NEOTROPICA
4-~~~~~~~~~~~~~~~~~~
12 -
s
Some species of Cecropia have been introduced
in neotropical regions outside their naturaldistribu-
tions, e.g., C. peltata in the Bermudasand C. poly-
stachya in Sao Paulo (Brazil), where they are natu-
ralized.
Several species have also been introducedin the
Palaeotropics,e.g., Cecropiapeltata (and possibly C.
pachystachya) in West Africa, Malaysia, and several
Pacific islands, where it is (or they are) are natural-
ized; in Indonesia,Taiwan,and China, where it is (or
they are) probablystill confinedto botanicalgardens.
Cecropia schreberiana was introducedin Mada-
gascar, where it is naturalized,and C. obtusifolia in
Hawai'i, also naturalized.
Cecropiapolystachya is (or was) in cultivationin
botanicalgardensin South Africa and Singapore.
REPRESENTATIONIN THE MAJOR
PHYTOGEOGRAPHIC SUBDIVISIONS
1. West Indies. Two species are found: Cecropia
peltata in Trinidadand in Jamaicaand C. schreberi-
ana in the Lesser and GreaterAntilles, except for Ja-
DISTRIBUTION
maica. The former species has a type which often
lacks trichilia in Jamaica and the widespread type
with trichilia in Trinidad.In C. schreberianatwo in-
fraspecificentities can be distinguished:a subspecies
in the Lesser Antilles and Puerto Rico, and a more
pubescent subspecies in Cuba, Hispaniola, Puerto
Rico, and the Virgin Islands.
2. Central America. Eleven species arerecorded,
of which only Cecropiapittieriis endemic,being con-
finedto Cocos Island.Cecropiaangustifoliais a mon-
tane species with its main areain SouthAmerica.The
other species are lowland elements which also occur
in Pacific Coastalregion of South America. Cecropia
longipes and C. heterochroma have rather small
ranges of distribution,from eastern Panamato adja-
cent Colombia. Cecropia hispidissima, C. garciae,
and C. virgusa range from Ecuadorto eastern Pan-
ama, and C. obtusifoliaand C. insignis from Ecuador
to southernMexico and Honduras,respectively.The
most common species in CentralAmerica is C. pel-
tata, which extends to Jamaica and to the northern
partof South America, far into the Magdalenavalley,
in the Pacific lowlands down to Buenaventura,and
through northernVenezuela to Trinidad, Suriname,
and the northernfringe of the Amazon basin (Brazil).
Cecropiamembranacea,one of the most widespread
species, occurs in the Amazon basin, the Pacific low-
lands, the Cauca valley (up to Valle) and the Lower
Magdalenavalley, and extends just into easternPan-
ama.
3. Pacific Coastal lowlands (Colombia-Ecua-
dor). In addition to the nine species that also occur
in CentralAmerica, this region contains Cecropiali-
toralis, common in Ecuador and extending to the
coastal part of Narifno(Colombia) and, remarkably
enough, also known from the Amazonianregion. Ce-
cropia obtusifolia, already quite variable in Central
America, exhibits confusing variation in Colombia.
Some of the species, such as C. hispidissimaand C.
obtusifolia, extend into submontane conditions in the
westernslopes of the Andes. On the otherhand,some
of the species that can be regardedas (sub)montane,
like C. reticulataand C. angustifolia, extend into the
lowlands, at elevations down to ca. 500 m. The upper
elevational limits of the typically lowland species of
this region vary from 100 to 1200 m, or sometimes
up to 1500 m.
4. Andean region (Venezuela-Bolivia). This re-
gion has the most diverse Cecropiaflora,but it is still
relatively poorly explored, due in part to limited ac-
cess to many areas inhabitedby these species. Many
of these species are known only by a few collections,
and ranges of distributionare often not clear. Several
27
regions, such as in PeruandBolivia, have hardlybeen
explored.
Most species have small to medium-sizedranges
of distribution,mostly confined to either the western
or the eastern slopes of the Andes. Cecropia angus-
tifolia is an exception. In South America it ranges
from the coastal mountainrange of Venezuelato Bo-
livia, occurring in Colombia and Ecuador on both
sides of the Andes, butalso in the Sierrade Macarena,
and in western Ecuadorand in Peru down to eleva-
tions of about 500 m. This species is differentiated
into a numberof regional and ecological forms, dis-
cussed underthe species.
On the western slopes of the Andes, at elevations
of 1700-2400 m, on the slopes at both sides of the
upperdrier partof the Cauca valley the white-leaved
Cecropia telealba is found, and in Ecuadorand ad-
jacent Colombia (Narinio)the white-leaved C. max-
ima. Species at lower elevations on these slopes are
C. bullata and C. gabrielis, both from Pichincha(Ec-
uador)to Antioquia(Colombia); C. megastachya,in
Choc6 and Valle (Colombia);C. plicata, from Antio-
quia to Valle; C. reticulata,from Antioquiato El Oro
(Ecuador);and C. subintegra,from Valle (Colombia)
to Carchi(Ecuador).The totalnumberof montaneand
submontanespecies recordedfor the western slopes
of the Andes is nine.
In the upper Magdalena valley at middle eleva-
tions two species are endemic: at low elevations the
lithophytic Cecropia goudotiana, and at middle ele-
vations C. mutisiana.
In the Planaltoareaof Antioquia,severalmontane
species are represented,the widespreadCecropiaan-
gulata, C. gabrielis and C. plicata, both extending
from the slopes of the Western Cordillera,the en-
demic C. multisecta, and the white-leaved C. teleni-
tida, a species from the Centraland EasternCordil-
leras.
On the eastern slopes of the Andes C. telenitida
extends fromVenezuelato Putumayo(Colombia)and
apparentlydisjunctly from Morona-Santiago(Ecua-
dor) to San Martin(Peru)at elevationsof 1700-2400.
The northernpartof the rangeof C. telenitidalargely
coincide with that of the "palmatisectaform" of C.
angustifolia, and at lower elevations that of C. sar-
arensis (of which the southernlimit is not known). In
southern Colombia (Caqueta) the "palmatisecta
form"of C. angustifoliais replacedby the "hachensis
form,"and two other species reach the northernlimit
of their distribution:the montaneC. andina (extend-
ing to northernPeru, at least to San Martin),and the
submontaneC. pastasana (extendingto centralPeru,
at least to Junin).In the centralpartof Ecuador,two
28
othersubmontanespecies C. utcubambanaand C. ve-
lutinella reach their northernlimits; C. utcubambana
extend to Madre de Dios and C. velutinella to San
Martin,but is not found in southernEcuador.Three
montane species appearto be limited to Peru, C. al-
bicans, rangingfrom San Martinat least to Pasco, C.
chlorostachya,recentlydiscoveredin San Martin,and
C. tacuna, ranging at least from Huainucoto Cuzco.
In Bolivia, the probablyendemic montanespecies C.
elongata is known from Cochabamba,La Paz, and
Santa Cruz. In this area C. strigosa is found at ele-
vations of ca. 900-1800 and is a submontaneelement
ratherthan a lowland one as it is in the northernpart
of its distributionrange.The numberof montaneand
submontanespecies on the eastern slopes of the An-
des is 11 (12 if C. strigosa is included).
5. Amazonian region. In this region 24 species
are found. Some of them are widespreadand extend
to other/adjacentregions: Cecropia distachya, C. la-
tiloba and C. sciadophylla(extendingto [partsof] the
Guianaregion), and C. membranacea(also occurring
west of the Andes). Two species with the main area
of distributionelsewhere occur in the peripheralparts
of the Amazon basin:C. peltata in the northernfringe
and C. pachystachyain the southernfringe. Cecropia
litoralis, a common species of the southernpartof the
Pacific coastal region, is also in the northwesternpart
of the Amazon basin.
The other species are either (largely) confined to
or distinctlyconcentratedin eitherthe upperAmazon
basin or the lower Amazon basin (mostly extending
to the easternpartof the Guianaregion). Elementsof
the upper Amazon basin are Cecropia annulata
(southern), C. engleriana (southernto northern),C.
ficifolia (widespreadand down to the middleAmazon
basin), C. herthae (northern),C. idroboi (northern),
C. marginalis (northern),C. polystachya (southernto
central), C. putumayonis (northern), C. strigosa
(southernto central), and C. utcubambana(northern
to central).The lattertwo species are submontanein
the northernpart of the range. Elements of the lower
Amazon basin are C. concolor (extending to central
Amazonia [Manausregion] and the southwesternpart
of the Amazon basin), C. obtusa (extending to the
Guianas),C. palmata (extendingto the easternpartof
the Guianas, to northeasternBrazil, to centralAma-
zonia [Manausregion], and to Amazonian Bolivia),
Guianas, C. sylvae (with a small distributionrange,
extendingto FrenchGuiana).Cecropiapurpurascens
and C. ulei occupy more or less restrictedareasin the
centralpartof the Amazon basin.
6. Guiana region-Venezuela (Rio Orinoco) to
Brazil (Amapa). Some predominantlyAmazonian
FLORANEOTROPICA
species extend to this region: Cecropiadistachya, C.
latiloba, C. obtusa, C. palmata, C. sciadophylla,and
C. silvae, and, moreover,C. peltata extending from
northernVenezuela.The species confinedto the Gui-
ana region are C. granvilleana(confinedto slopes of
graniteoutcrops,inselbergs,in FrenchGuiana),C. ka-
vanayensis(montanein easternVenezuelaandthe ad-
jacent partsof Guyanaand Brazil [Roraima]),and C.
angulata (also in the easternpartof the region).
7. South America, South of the Amazon Basin.
Five species occurin the region.The most widespread
species is Cecropiapachystachya(easternandcentral
Brazil, northernArgentinaand Paraguay,a member
of the C. peltata-group).Cecropiasaxicola occurs in
centralBrazil andextendsto easternBolivia; it is usu-
ally found in rocky areas. Two species are confined
to more humid areas in eastern Brazil: C. glaziovii
(closely related to C. palmata) and the white-leaved
C. hololeuca (probablyrelatedto an Andeangroupof
montane species). The Amazonian C. palmata ex-
tends south of the Amazon basin to northeasternBra-
zil.
SYSTEMATIC POSITION
Cecropia is one of the six genera which consti-
tuted the tribe Conocephaleae in the classification
presentedby Trecul (1847) and the subfamilyCono-
cephaloideaein the Moraceaein thatpresentedby En-
gler (1889). The other genera included in these taxa
were the Asian genus ConocephalusBlume, the Af-
rican generaMusangawith 2 species andMyrianthus
with 7 species (see de Ruiter,1976), and the neotrop-
ical generaCoussapoawith 48 hemiepiphyticspecies,
Pourouma with 27 species of terrestrialtrees (see
Berg et al., 1990; Berg & Franco Rosselli, 1993,
1996). ChewWee-Lek(1963) includedConocephalus
in PoikilospermumMiquel, a genus until then treated
in the Urticaceae.In his revision of Poikilospermum
20 species were recognized.The studyby ChewWee-
Lek raisedquestionsaboutthe position of the six gen-
era. Chew Wee-Lek suggested transferringthe mi-
crospermousgenera to the Urticaceae and keep the
macrospermousgenera Myranthusand Pouroumain
the Moraceae. Corner(1962) suggested transferring
all six genera to the Urticaceae while Berg (1978a)
placed them in a separatefamily, the Cecropiaceae.
Berg's concept was accepted, e.g., by Cronquist
(1981). Others(e.g., Juddet al., 1989) preferto regard
the group of six genera as a subdivisionof a broadly
construedfamily Urticaceae,also includingthe Mor-
aceae; and Romaniuc-Neto(1999) reducedthe group
again to a subfamily of the Moraceae, but with the
name Cecropioideae.The "technical"characters(to
TAXONOMICDIFFERENTIATION 29

key the three families out) are few and not strong: II. Section Atomentosae. Perianths of the sta-
milky sap (Moraceae), urticaceousstamens (Urtica- minate flowers with short hairs or glabrous.
ceae andMoraceaep.p.), andbasal ovules (Urticaceae Filamentsunequallylong, but in Polystachyae
and Cecropiaceae)versus apical ovules (Moraceae). about equally long.
However, the three families are clearly distinct in D. Centrales. Staminate spikes medium-
overall morphological (and ecological) diversifica- sized, relatively thick. With five species:
tion. The Cecropiaceaealways have adventitiousaer- Cecropia litoralis, three species presently
ial roots, becoming stilt-rootsin most genera or hold in the synonymy of C. peltata, and C. ulei.
fasts for root-climbers(Poikilospermum),or adapta- E. Angulatae. Staminatespikes angular.With
tions to the hemiepiphyticlife form (Coussapoa).The only Cecropiaangulata.
plants contain mucilaginous sap that turns black on F. Elongatae. Staminateand pistillate spikes
exposureto the air.The stipules are fully amplexicaul very long, to 40 cm or more.WithCecropia
(except in some species of Poikilospermum)and con- elongata and C. mexicana (= C. obtusi-
nate, often large and protectingdeveloping inflores- folia).
cences. The plants are always dioecious (unlikeMor- G. Abbreviatae. Spikes usually very short,or
aceae and Urticaceae), and they are microspermous if medium-sized, then "elegant."With 16
or macrospermous(as in Moraceae).Pearlglands are species, including Cecropia angustifolia,
common andthe cystolithsaccumulatesilicon instead C. ficifolia, C. obtusa, C. peltata, and C.
of calcium (Setoguchi et al., 1993). pittieri.
Poikilospermumis morphologicallyand anatomi- H. Polystachyae. Staminate spikes very nu-
cally clearly different from the other five genera merous. Pistillate spikes short and thick.
(Berg, 1989; Renner, 1907). The two macrospermous Stamens short, almost equally long. With
genera, Pouroumaand Myrianthus,constitute a pair four species, including Cecropiamultifiora
of genera with clear morphological and ecological (= C. strigosa).
similarities. The same applies to Cecropia and Mu-
Five species, including Cecropiamontana,could
sanga, but the differencesbetween the inflorescences
not be included in one of these unrankedgroups.
make clear that Musanga cannot be regardedas "a
This classificationwas stronglyinfluencedby the
Cecropia withoutants"(Janzen& McKey, 1977), but
small numberof collections availableat thattime. The
they are probablysister genera. WhereasMusangais
differentiatingcharacterschosen cannotor can hardly
representedby only two species, a lowland and a
be connected to differences in other parts. The pro-
montaneone, speciationbased on bothmorphological
posed subdivisioncannot be used effectively.
and ecological differentiationhas been considerable
The morphological differentiation,as presently
in Cecropia.
known, does not allow the establishmentof a satis-
factory subdivisionof the genus. Therefore,the spe-
cies are presentedin alphabeticalorderin this mon-
TAXONOMIC DIFFERENTIATION
ograph. There are some morphologically isolated
The only attemptto establishan infragenericclas- species, some groupsof two or threespecies showing
sification was made by Snethlage (1923) to accom- clear affinities, some larger groups showing more
modate the 42 species he recognized. It is based on vague or rather clear affinities. The species with a
the differences in indumentumon perianths of sta- more or less isolated position due to the absence of
minateflowers,the length of filaments,andthe length characteristicgeneric featuresareCecropiahololeuca
of spikes of staminateand pistillate inflorescences. and C. sciadophylla. The latteris distinct by the ab-
I. Section Tomentosae. Perianthsof the stami- sence of trichilia,which might not be secondarilylost
nate flowers covered with white felted or as in most other cases and can neither be related to
arachnoidindumentum. occurrence on islands and in montane habitats. Ce-
A. Aequales. Filaments equally long. With cropia hololeuca is distinct primarilyby the absence
Cecropiaconcolor and C. palmata. of trichilia.It also lacks well-developedspathes.The
B. Subequales. Filaments unequally long. absence of well-developed spathes might be second-
With Cecropia glaziovii, C. hololeuca, C. ary. Cecropia hololeuca shows in its habit and its
saxatilis, and C. sciadophylla. white laminas similaritiesto a groupof montaneAn-
C. Arachnoideae. Perianth with long arach- dean species, which may also lack trichilia.The sets
noid indumentum. With only Cecropia of two or three clearly relatedspecies comprise, e.g.,
maxima. C. glaziovii and C. palmata; C. obtusifolia and C.
30
subintegra;C. angustifolia and C. montana; C. lati-
loba and C. utcubambana;C. garciae, C. hispidis-
sima, and C. megastachya;and C. membranacea,C.
polystachya, and C. strigosa. The two largergroups
of species with more or less clear affinitiesare the C.
peltata-groupand the C. telenitida-group.
Cecropiapeltata-group. This group is character-
ized by the presence of peltate stigmas. In contrastto
the more common comose to penicillate stigmas, the
peltate stigma is already outside the aperture of
the perianthat an early stage of developmentof the
flower,and as a consequence,the arachnoidindumen-
tum of the perianthcovers not only the perianthbelow
the apex (as commonly), but also on the apex to near
the aperture.However, this feature is not present in
C. granvilleana,nor is it in all materialrankedunder
C. sararensis and C. schreberiana.In the latter two
species the stigma varies from peltate and subpeltate
to penicillate-comose in combinationwith the pres-
ence of arachnoidindumentumvarying from present
on the apex of the perianthto only below the apex.
In the C. peltata-groupthe trees are usually small to
medium-sized,rarelytall (as sometimes in C. engler-
iana). The stems have prominentscarsof the stipules.
Tanninis absent or scarce in the perianthsof the pis-
tillate flowerandarachnoidindumentumis mostly ab-
sent in the style channel.
The morphologically most distinct taxa of this
groups are Cecropiaengleriana, C. granvilleana,and
C. schreberiana. Cecropia engleriana is distinct by
the large numberof lateral veins in the free part of
the midsegment of the lamina, normally at least 24
pairs,rarelyabout21 pairs,whereastherearebetween
8 and 20, or rarely up to 24 pairs, in all other taxa.
Cecropiagranvilleanais distinctby the small number
of lateralveins in the free partof the midsegmentof
the lamina (9-10 pairs, whereasin the othertaxa nor-
mally more than 10) and the consistent absence of
peltate stigmata.Cecropiaschreberianais distinctin
the remarkableabsence of trichilia (or the occasional
presence of poorly developed ones). Trichilia are
present in all other species of the group. However,
they are absentin partof the collections of C. peltata
made in Jamaica (or also in part of the individuals
introducedin the Old World).In C. schreberiana,the
intemodescontainample brownpith, this being white
and scarce in the other taxa. The indumentumon the
leafy twigs and the base of the petiole can be subvil-
lous. In these threeaberrantfeatures,C. schreberiana
resembles a number of Andean montane species.
Within C. schreberiana two morphological entities
can be distinguished(in the presenttreatmentrecog-
nized as subspecies):subsp.schreberianaglabrousor
FLORA NEOTROPICA
sparsely hairy on the leafy twigs and veins on the
lamina beneathand penicillate-comosestigmata,and
subsp. antillarummore hairy, at least at the venation
of the lamina beneathand transitionsfrom peltate to
penicillatestigmas,therefore,becomingin generalas-
pect quite similarto the neighboringC. peltata. Sim-
ilar variationwith regardto stigma and indumentum
appearsto occurin the submontaneC. sararensis.The
northernglabrousformof C. sararensiscan be readily
distinguishedfrom the otherentities of the C.peltata-
group (C. concolor, C. litoralis, C. metensis, C. pel-
tata, and C. pachystachya), but the southern hairy
form less easily.
The AmazonianCecropiaconcolor can be distin-
guished by the deeply incised lamina, but transitions
to the leaf shape of C. pachystachya occur in the
southernpartof the Amazon basin. Cecropialitoralis
and C. sararensis are distinct from C. metensis, C.
pachystachya, and C. peltata by the relatively long
spikes, in particularin the pistillate inflorescences.
Cecropia metensis is very close to C. pachystachya,
and essentially not distinguishablefrom types of the
latter species with dense arachnoidindumentumon
the petiole and other plant parts. The long whitish
hairsin the trichiliacharacteristicof C. pachystachya
and C. metensis are sometimes found in areas where
the rangesof C.peltata and C. metensismeet (or over-
lap?).
The two taxa with widest distribution,Cecropia
peltata and C. pachystachya,cannotbe distinguished
satisfactorilyfrom each other.Differences in the var-
iation of the length of the stipules, the presence/ab-
sence of lobation of the lamina segments, and the
presence/absenceof long white hairs in the trichilia
can be used to determinethe species morphologically.
Lobation of leaf segments, at least in juvenile
stages, is found in Cecropiapachystachya,C. meten-
sis, and C. concolor.
The taxa of the Cecropiapeltata-groupshow over-
lap in distributiononly in some cases: C. concolor
and C. pachystachyain the southernAmazon basin,
C. concolor and C. engleriana in the southwestern
part of the Amazon basin, and C. engleriana and C.
litoralis in the northwesternpartof the Amazonbasin.
Slight overlapof distributionrangesmay occur for C.
metensisand C.peltata, as well as for C. metensisand
C. sararensis.But in all those cases intermediatesare
absent or rare.
In general, the taxa of the Cecropiapeltata-group
can be regardedas belonging to the "weedy"group
of Cecropia species. Cecropia granvilleana is pro-
nouncedly adaptedto wet granite rock surfaces. Ce-
cropia metensisand C.pachystachyaare largelycom-
VERNACULARNAMES
ponents of savanna,cerrado,and restingavegetation.
Otherspecies show variationwith regardto ecological
requirements.Cecropia engleriana and C. litoralis
are in partsof theirranges of distributionuplandspe-
cies but in otherparts more or less clearly associated
with temporarilyinundatedhabitats.Cecropiapeltata
is found both in regions with wet rain forest and in
dry types of vegetation, as some of the inter-Andean
valleys in Colombia.
The distributionpatternsand ecological differen-
tiation suggest segregationof a very widespreaden-
tity,resultingin variousdegreesof morphologicaldis-
tinctiveness. The taxonomy seems to be expressed
most adequatelyby treatingthe nine taxa as species,
ratherthanto unite some of them fully, reduceothers
to subspecies, and maintainthe rest as species. For
the Cecropia peltata-group a separate key is con-
structedbelow.
Cecropia telenitida-group. It comprises species
of which the leafy twigs and the upper leaf surface
are glabrous (or almost so) and the outer surface of
the stipules and spathesare eitherglabrousor (partly)
villous with long, more or less soft, whitish hairs. In
many representativesof this groupthe upperleaf sur-
face is often or usually covered with more or less
dense arachnoidindumentum.In this groupof species
trichilia can be always absent, occur only in more or
less reducedstates(mostly withoutformingMiullerian
bodies), or occur in states of being absent through
being more or less reduced to occurring in two
patches to well-developed and fused. In some of the
species the differentstates can even be found on the
same tree. Most of these species are not inhabitedby
Azteca ants.The internodesmay containamplebrown
pith in the internodes.The numberof spikes of the
pistillateinflorescencesmay be reducedto one or two.
The group is distinctly concentratedin the northern
Andeanregion and is associatedwith montaneto sub-
montane habitats. It comprises C. albicans, C. bul-
lata, C. gabrielis, C. maxima,C. telealba, and C. te-
lenitida. Cecropia hololeuca (from eastern Brazil),
with its white laminas and pistillate inflorescences
with one or two spikes, could be a member of this
group. Cecropia schreberiana (included in the C.
peltata-group)could be linked to this group,because
of the presenceof amplepith and absenceof trichilia.
The montanespecies C. multisectaand C. plicata, as
well as the lowland species C. insignis, could be
linked to the C. telenitida-group.
A more vaguely definable group, largely associ-
ated with the Guayanaregion and with relativelyfew
and often broad lamina segments as a common fea-
ture, comprises Cecropia angulata, C. ficifolia, C.
31
distachya, C. kavanayensis, C. obtusa, and C. pur-
purascens. It might also include C. saxatilis, which
is, however,clearly distinct in the staminateflowers.
USES
The use of the genus is limitedto largelylocal uses
(cf. Duarte, 1959;Velasquez,1971; Galiano-Sanchez,
1976;Garcia-Barriga,1974). Decoctions of leaves are
made to stimulatethe cardiacsystem, to treatasthma
and pneumonia, to treat diabetes, and as diuretic.
Powder of leaves is used to control Parkinson'sdis-
ease. Extractof roots are used to heal wounds or ec-
zema. Plastersof leaves are madeto reduceswellings.
Ash of leaves (of various Cecropiaspecies, in partic-
ular those C. sciadophylla) is mixed with leaves of
coca (Erythroxylumspp.) to reduceaciditywhen these
leaves are chewed (cf. Plowman, 1981, 1984).
Branchesareused to makepercussioninstruments(in
Nariino,Colombia) or flutes. Cortex fibers are used
for several purposes,such as ropes, bow strings, and
hammocks.Leaves of some species are used as cattle
fodder, to clean utensils, or to polish furniture.The
spikes ("bananitas")are eaten or locally sold on mar-
kets (e.g., in Santa Cruz, Bolivia). Wood is used to
make cheap casketsor matches,or to makerafts.Sev-
eral attemptsto use the wood for the productionof
paper pulp have not been successful, partlybecause
of problemswith the regenerationof the rawmaterial.
Pith of the petioles is used to make ornamentalhead-
gear (in Napo, Ecuador).
VERNACULAR NAMES
Guarumois the commonly used vernacularname
in Spanish-speakingcountries in Central America,
Mexico, the Caribbean,Colombia, and Ecuador.The
variantguarumbo is encounteredin several Central
American countries, yagrumo in Venezuela,and yu-
arumo in Colombia and Ecuador.In the Amazonian
partof Peru,setico (or cetico) is the common vernac-
ular name, but in the Andean partof this countryta-
cuna (or tacona) is in common use. Imbaiiba (also
embalibaor umbauiba)is the common name used in
Brazil, in particularin the Amazonian part of the
country,and ambaiba (or ambaiba) is more common
in the southernpart of the country,as well as in Par-
aguay. Ambaibo is the common vernacularname in
Bolivia. Bois canon is commonly used in the French-
speaking countries of the West Indies and in French
Guiana, whereas trlimpettree is the most common
vernacularname in the English-speakingcountries.
The name snake wood, used in Jamaica,is connected
to the scientific name, which is derived from "Ce-
32 FLORA NEOTROPICA

crops"(or "Kekrops"),a Greekmythologicalcreature, Coilotapalus P. Browne, Civ. Nat. Hist. Jamaica 111.
being half snake and half human.The less commonly 1756, nom. rejic. Type not designated.
used vernacularnames, the common ones with adjec- Ambaiba Barrere,Ess. Hist. Nat. Fr. Equin. 10. 1741;
Adanson, Fam. P1. 2: 377. 1763, as synonym; Kun-
tives, and unusual variants are listed under the spe-
tze, Rev. Gen. 2: 623. 1891.
cies.
Tree,dioecious, terrestrial,usuallywith stilt-roots;
intemodes (mostly) hollow; sap watery,turningblack
CONSERVATION at exposure to the air. Leaves in spirals;lamina sim-
The majority of the species are not endangered. ple, peltate, radiallyveined and incised; petiole usu-
Many of them temporarilyincreasein numberby the ally with one or two patches of dense indumentumof
destructionof forest, but continuousclearing leads to various trichomes (trichilia) at the base; stipules
reductionof numbers,firstof the "forest"species, and large, connate, fully amplexicaul, leaving circular
finally also of the "weedy"species. However, a few scars. Inflorescences axillary, pedunculate digitate
species thatarerestrictedto rockyplaces (as Cecropia clusters of spikes, sometimes 1 spike, (usually) en-
goudotiana), have scatteredoccurrence,and/orpos- closed by a spathe until anthesis. Flowers unisexual,
sess small ranges of distribution(as Cecropiamulti- sessile (or short-pedicellate);tepals 2, rarely3, mostly
secta) can easily become extinct. Second in vulnera- entirely connate, forming a tubularperianth with a
bility are the "small gap" species such as C. narrowaperture;stamens2, rarely3, free; ovaryfree,
putumayonisand C. utcubambana. stigma penicillateto peltate.Fruit a small achene, of-
ten tuberculate,fully enclosed by a fleshy perianth;
seed with endosperm;embryo small, with flat coty-
SYSTEMATICTREATMENT ledons.
Cecropia Loefling, Iter. Hispan. 272. 1758, nom.
cons.; Miquel in Martius, Fl. Bras. 4(1): 139; KEYS TO THE SPECIES
Snethlage, Notizbl. Bot. Gart. Berlin-Dahlem8: A general key to the species would contain too
357. 1923; Cuatrecasas,Revista Acad. Colomb. many weak entries because of dioecy and the varia-
Ci. Exact. 6(22/23): 274. 1945 and 9(36/37): 325. tion of charactersthatcan be used to key out the spe-
1956; Berg, Acta Amazonica 8(2): 149. 1978; cies. Therefore, only keys for countries or larger
Berg & FrancoRosselli in Harling& Andersson, regions have been constructed,as well as a key to the
Fl. Ecuador 48: 3. 1993. Type. C. peltata Lin- species of the C.peltata-group,but nonetheless,weak
naeus. points remainin most of the keys.

Key to the species of CentralAmerica, Mexico, and the West Indies

1. Trichiliaabsent (or poorly developed, in two patches with or withoutMullerianbodies).
2. Intemodes with ample brownpith; Cuba, Hispaniola,PuertoRico, and Lesser Antilles ........... 50. C. schreberiana
2. Internodeswith sparsewhitish pith.
3. Incisions of the lamina down to 2/10-3/10 from the margin;Cocos Island . ........................42. C. pittieri
3. Incisions of the lamina down to 5/10-7/10 from the margin;Jamaica . . 41. C. peltata
1. Trichiliapresent.
4. Trichiliaseparateand enclosed in 2 pockets with slit- shapedaperturesat the base of the petiole;
Panama...................................................................... 61. C. virgusa
4. Trichiliafused, superficial.
5. Laminawithout arachnoidindumentumin the areoles beneath(at most initially with arachnoid
indumentumon the marginand/ormain veins).
6. Lateralveins in the free partof the midsegment(10-)15-25(-50) pairs;Mexico to Panama
................................................................................................................................................37. C. obtusifolia
6. Lateralveins in the free partof the midsegment 10-13 pairs.
7. Laminasegments 6-8; spikes of the staminateinflorescence4-5; spikes of the
pistillate inflorescence(1-)2-4, 3-10 cm long at anthesis, to 15 cm long in fruit;
Panama................................................................ 19. C. heterochroma
7. Laminasegments (7-)8-10; spikes of the staminateinflorescenceca. 30-40; spikes of
the pistillate inflorescence4, 10-16 cm long at anthesis, to 30 cm long in fruit;
Panama................................................................... 31. C. membranacea
5. Laminawith arachnoidindumentumat least in the areoles beneath.
8. Lateralveins in the free partof the midsegment25-50 pairs.
SYSTEMATICTREATMENT 33

9. Leafy twigs and petioles villous to hirsuteor hirtellousto strigose, but without irritating
bristles; Mexico to Panama............................................................... 4. C. angustifolia
9. Leafy twigs and petioles partly setose with irritatinghairs.
10. Trichiliawith bristles among the brownpluricellulartrichomes,the Millerian
bodies pink, ca. 3 mm long; lamina with (9-)10-13(-15) segments, smooth above;
Panama......................................................... 20. C. hispidissima
10. Trichiliawith short brownor long white soft hairs among the brownpluricellular
trichomes,the Mullerianbodies white, ca. 1 mm long; lamina with 14-19 segments,
usually scabridulousabove; Panama.................................................................. 14. C. garciae
8. Lateralveins in the free partof the midsegmentto ca. 25 pairs.
11. Lateralveins in the free partof the midsegment5-8 pairs;spikes of the staminate
inflorescence30-80; peduncle of the pistillate inflorescence30-65 cm long; Panama
............................................................................................................................................27. C. longipes
11. Lateralveins in the free partof the midsegment(8-)10-25 pairs;spikes of the staminate
inflorescenceto ca. 25; peduncle of the pistillate inflorescenceto 15 cm long.
12. Laminausually with 7-10 segments and/orfree partof the midsegmentwith up to 15
pairs of lateralveins.
13. Stipules usually to 10 cm long; Mexico to Panama,Jamaica,and Trinidad..... 41. C. peltata
13. Stipules 10-50 cm long.
14. Stipules 10-15(-25) cm long; spikes of the inflorescencessessile; Panama
.................................................................................................................47. C. reticulata
14. Stipules (15-)20-50 cm long; spikes of the inflorescenceswith stipes;
Hondurasto Panama................................................... 23. C. insignis
12. Laminausually with 10-13 segments and the free partof the midsegmentusually
with 15-25 pairs of lateralveins.
15. Indumentumof leafy twigs and petioles (partly)brownish;lamina (almost)
smooth above; spathesof the inflorescencesdensely hairy outside; Mexico to
Panama......................................................... 4. C. angustifolia
15. Indumentumof leafy twigs and petiole whitish; lamina ? scabrousabove;
spathes of the inflorescences ? sparselyhairy outside; Mexico to Panama
.......................................................................................................................37 C. obtusifolia

Key to the species of Colombia

1. Trichiliaenclosed in 2 pockets with slit-shapedaperturesat the base of the petiole; Pacific region ... 61. C. virgusa
1. Trichiliaexposed (superficial)or absent.
2. Trichiliaabsent.
3. Incisions of the lamina down to the petiole, the segments often petiolulate;Amazonianregion
.51.
.................................................................................................................................................. C. sciadophylla
3. Incisions of the lamina not down to the petiole; Andean region or Pacific region.
4. Indumentumpartlysetose with irritatinghairs;Pacific region ......................................... 20. C. hispidissima
4. Indumentumnot partlysetose, without irritatinghairs;Andean region.
5. Lamina ? scabrousabove; E Cordillera............................................................... 2. C. andina
5. Laminasmooth above; W Cordillera......................... ...................................... 13. C. gabrielis
2. Trichiliapresent.
6. Trichilia separate(in two lateralpatches).
7. Laminawith ? dense arachnoidindumentumabove.
8. Lateralveins in the free partof the midsegment7-16 pairs;arachnoidindumentumin two
distinct layers on the laminabeneath;E and centralCordillera....................................... 57. C. telenitida
8. Lateralveins in the free partof the midsegmentca. 15-25 pairs;arachnoidindumentumnot
in distinct layers on the lamina beneath;W Cordillera...................................................... 56. C. telealba
7. Laminawithout arachnoidindumentumabove.
9. Laminachartaceousto subcoriaceous;spikes of the staminateinflorescenceca. (20-)30-50;
spikes of the pistillate inflorescence4(-7); Amazonianregion...................................... 28. C. marginalis
9. Lamina(sub)coriaceous;spikes of the staminateinflorescence5-9; spikes of the pistillate
inflorescence 1-3(-4); Andean region.
10. Leafy twigs hairy; arachnoidindumentum(almost) absent or in minute patches
surroundedwith thick veinlets on lower surfaceof the lamina .................. ................. 43. C. plicata
34 FLORANEOTROPICA

10. Leafy twigs glabrous;arachnoidindumentumin the areoles or also on veins on the

lower surfaceof the lamina ................................................................... 13. C. gabrielis
6. Trichiliafused.
11. Lamina(usually) with 10-20(-24) segments;lateralveins in the free partof the midsegment
mostly >25 pairs.............................................................. 48. C. sararensis
12. Setose (irritating)hairs presenton young parts;Pacific region.
13. Trichilianot well-defined;Mullerianbodies pink, ca. 3 mm long .......... .......20. C. hispidissima
13. Trichiliawell-defined;Mullerianbodies white, ca. 1 mm long.
14. Lateralveins in the free partof the midsegmentca. 35-50 pairs;lamina usually
scabridulousabove.................................................. 14. C. garciae
14. Lateralveins in the free partof the midsegmentca. 20-30 pairs; lamina smooth
above............................................................ 30. C. megastachya
12. Setose (irritating)hairs absent.
15. Laminasmooth above.
16. Incisions of the lamina down to the petiole; Antioquia, 1300-1600 m ...... 34. C. multisecta
16. Incisions of the lamina not down to the petiole.
17. Laminaoften white above, segments of young leaves patent;spikes of
staminateinflorescence4-6(-8); spikes of pistillate inflorescence(1-)2-3,
the spathe 10-20 cm long ..................... ................................ 29. C. maxima
17. Laminagreen above, segments of young leaves deflexed;spikes of
staminateinflorescence 10-25(-30); spikes of pistillate inflorescence3-8
(-12), the spathe 3-6(-8) cm long...................................................... 4. C. angustifolia
15. Lamina ? scabrousabove.
18. Incisions of the laminadown to the petiole or 1-2 cm from the petiole; stipules
20-30 cm long; Amazonianregion..................................................... 18. C. herthae
18. Incisions of the laminanot down the petiole, or if 1-2 cm from the petiole,
then the stipules <20 cm long.
19. Lateralveins in the free partof the midsegmentusually 30-50 pairs;
spathesand stipules with dense arachnoidindumentumoutside; E
Cordillerato Amazonianregion.................................................... 33. C. montana
19. Lateralveins in the free part usually 10-25 pairs, or if to 35(-43) pairs,
then the spathes and stipules withoutdense arachnoidindumentumoutside.
20. Spathesof the inflorescencessparselyhairy to subglabrousoutside;
Pacific region................................................. 37. C. obtusifolia
20. Spathes ? densely hairy outside; Andean region.
21. Lateralveins in the free partof the midsegment 10-21 pairs;
pistillate inflorescencessoon ? pendulous.
22. Upper marginof the rim of the trichiliumlobate; Upper
Magdalenavalley.................. ........................ 35. C. mutisiana
22. Upperrim of the trichiliumentire;W Cordillera........ 48. C. sararensis
21. Lateralveins in the free partof the midsegmentoften >21;
pistillate inflorescenceserect; widespread.......................... 4. C. angustifolia
1. Laminawith (5-)7-10(-12) segments;lateralveins in the free partof the midsegmentmostly
<25 pairs.
23. Laminawithout arachnoidindumentumin the areoles beneath.
24. Laminacoriaceous, often ? plicate; Andeanregion, 1000-2300 m....................... 43. C. plicata
24. Laminachartaceousto subcoriaceous;plane; Amazonianand/orPacific region.
25. Base of the lamimaentire or shallowly lobed, the upperpart 3-lobed.
26. Spikes of the staminateinflorescence4-5; peduncle of the pistillate
inflorescence25-45 cm long; Amazonianregion ........................... 46. C. putumayonis
26. Spikes of the staminateinflorescence7-12; peduncle of the pistillate
inflorescence3-24 cm long; Pacific region ........................................ 54. C. subintegra
25. Base of the lamina ? distinctlyincised.
27. Lateralveins in the free partof the midsegmentusually 15-25(-40) pairs;
Pacific region.............................................. 37. C. obtusifolia
27. Lateralveins in the free partof the midsegment9-16 pairs.
28. Inflorescences(or at least the spikes) pendulousat anthesis;spikes of
the staminateinflorescenceca. 30-40; spikes of the pistillate
SYSTEMATICTREATMENT 35

inflorescence 10-16 cm long at anthesis (in fruit to 30 cm long);

Amazonianregion and Pacific region, usually riverine......... 31. C. membranacea
28. Inflorescenceserect at anthesis;spikes of the staminateinflorescence
4-25(-30); spikes of the pistillate inflorescence3-10 cm long at
anthesis (in fruit to 15 cm long).
29. Laminawith 6-8 segments, the incisions down to 3/10-5/10
from the margin;Pacific region .................................. 19. C. heterochroma
29. Laminawith (8-)10-14 segments, the incisions down to 7/10-
8/10 from the margin;Andean region ............... ................. 4. C. angustifolia
23. Laminawith arachnoidindumentumin the areoles beneath.
30. Lateralveins of the free partof the midsegmentmarginallyloop-connected.
31. Lateralveins in the free partof the midsegment24-32 pairs;Amazonianregion
.....................................................................................................................II. C. engleriana
31. Lateralveins in the free partof the midsegment8-24 pairs.
32. Lamina(sub)coriaceous,often smooth above.
33. Lateralveins in the free partof the midsegment8-10(-12) pairs;
Amazonianregion............................................. 22. C. idroboi
33. Lateralveins in the free partof the midsegment 12-24 (or more)
pairs.
34. Lateralveins in the free partof the midsegmentusually 12-18
pairs;lamina smooth above; stigma comose; Amazonianregion
.................................................................................................9. C. distachya
34. Lateralveins in the free partof the midsegment> 18 (to 24 or
more) pairs, or if < 18 pairs, then the lamina scabrousabove and
the stigma peltate;not Amazonian.
35. Spathe of the pistillate inflorescence8-15 cm long;
peduncle of the staminateinflorescenceusually >8 cm long
and the spatheca. 10-20 cm long; E Cordillera(E slopes,
300-1400 m)...................................... 48. C. sararensis
35. Spatheof the pistillate inflorescence3-6(-8) cm long;
peduncle of the staminateinflorescenceusually <8 cm long
and the spathe usually 4-10 cm long; Andean region,
widespread,mostly >1500 m ....................... ..............4. C. angustifolia
32. Laminachartaceousto subcoriaceous,mostly ? scabrousabove.
36. Trichiliawith white hairs distinctly longer than the brown
indumentum.
37. Petiole with dense arachnoidindumentum;spikes of the
staminateinflorescence4-20; peduncle of the pistillate
inflorescence3-17 cm long; llanos region............................. 32. C. metensis
37. Petiole without arachnoidindumentum;spikes of the staminate
inflorescenceca. 30-80; peduncle of the pistillate inflorescence
ca. 30-90 cm long; Pacific region.......................................... 27. C. longipes
36. Trichiliawith white hairs about as long as the brown indumentum.
38. Stipules and spathes densely hairy outside; inflorescenceserect;
lateralveins in the free partof the midsegmentoften >20 pairs;
Andes, mostly >1500 m ......................... ................ 4. C. angustifolia
38. Stipules and spathes ? sparselyhairy outside; inflorescences
with at least the spikes pendulousor the pedunclebecoming
deflexed;lateralveins in the free partof the midsegment 12-19
pairs;lowlands.
39. Laminaattachedto the petiole at ca. 1/4 of the total length
from its base; spikes of the staminateinflorescenceusually
ca. 15-20, 14-20 cm long; stigma penicillate;Amazonian
region, inundatedplaces..................................... 25. C. latiloba
39. Laminaattachedto the petiole at clearly >1/4 of the total
length from its base; spikes of the staminateinflorescence
usually ca. 5-15 and 5-12 cm long; stigma peltate;Pacific
region, in non-inundatedand inundatedplaces, and
Amazonianregion, in inundatedplaces........................... 26. C. litoralis
36 FLORA NEOTROPICA

30. Lateralveins of the midsegmentloop-connectedbefore the margin.

40. Trichiliaon a subscrotiform(abaxiallybulging) base of the petiole.
41. Trichilialaterallyextended by lobes; spikes of the staminateinflorescence
4-12, 6-12 cm long; spathe of the pistillate inflorescence5-9 cm long; E
Cordillera.............................................. 47. C. reticulata
41. Trichilialaterallyrounded;spikes of the staminateinflorescence 10-25,
(6-)12-28 cm long; spathe of the pistillate inflorescence 10-17 cm long;
E Cordillera...................................................... 40. C. pastasana
40. Trichilianot on a subscrotiform(abaxiallybulged) base of the petiole.
42. Lamina(sub)coriaceousand usually smooth above.
43. Upper surface of the lamina ? bullate, glabrous;leafy twigs and
petioles villous to subsericeousto subhirsute;W Cordillera............. 6. C. bullata
43. Upper surfaceof the lamina plane or plicate, or if slightly bullateand/
or with villous to subhirsuteindumentumon the leafy twigs and
petioles, then with arachnoidindumentumon the lamina above.
44. Upper surface of the lamina white or whitish with ? dense
(sometimes sparse)arachnoidindumentum.
45. Arachnoidindumentumon the laminabeneathin two
distinctlayers (in the areoles and on the stiff hairs on the
veins); spikes of the pistillate inflorescencesmostly 4-5; E
and centralCordilleras.............. 57. C. telenitida
45. Arachnoidindumentumon the lamina beneathnot in two
distinctlayers;spikes of the pistillateinflorescencemostly 1-
3; W Cordillera.............. 56. C. telealba
44. Upper surfaceof the lamina without (or with very sparse)
arachnoidindumentum.
46. Leafy twigs with ? dense arachnoidindumentum,spikes of
the staminateinflorescence3-9, and those of the pistillate
inflorescence l-3(-4); W Cordilleraand Antioquia.
47. Lamina ? plicate and with 8-10(-I 1) segments;stipes
of the spikes of the staminateinflorescence0.5-2.5 cm
long; spikes of the pistillate inflorescence2-3(-4)
................................................................................... 43. C. plicata
47. Laminaplane and with 7-8 segments;stipes of the
spikes of the staminateinflorescence0.4-0.5 cm long;
spikes of the pistillateinflorescence 1 or 2............ 13. C. gabrielis
46. Leafy twigs without dense arachnoidindumentum,or if
with such indumentum,then spikes of the staminate
inflorescence 10-25(-40) and those of the pistillate
inflorescence3-8(-12).
48. Leafy twigs glabrous(often with a bluish waxy layer)
................................................................................ 13. C gabrielis
48. Leafy twigs hairy.
49. Stipules 22-50 cm long; leafy twigs sparsely
hispidulous;spikes of the pistillateinflorescence8-
18 cm long at anthesis,to 26 cm long in fruit;
Pacific region ........... .................... 23. C. insignis
49. Stipules 6-25 cm long; leafy twigs ? densely
hairy, hispidulousto subvillous;spikes of the
pistillateinflorescence 1-7 cm long at anthesis, to
17 cm long in fruit;Andean region, widespread
. 4. C angustifolia
....................................................................
42. Laminachartaceousto subcoriaceousand usually ? scabrousabove.
50. Lateralveins in the midsegmentmostly 15-35 pairs.
51. Spathesmore or less densely hairy outside; Andean region,
widespread............................. 4. C. angustifolia
51. Spathes sparselyhairy to subglabrousoutside; Pacific region
............................................................................................. 37. C obtusifolia
50. Lateralveins in the midsegment6-15(-17) pairs.
SYSTEMATICTREATMENT 37

52. Upper surfaceof the lamina with ? dense (rarelysparse)

arachnoidindumentum;W and centralCordilleras.............. 57. C. telenitida
52. Upper surfaceof the lamina without(or rarelywith very sparse)
arachnoidindumentum;lowland.
53. Stipules 6-8 cm long.
54. Lateralveins in the free partof the midsegment7-10
pairs; spikes of the staminateinflorescence4; spikes of
the pistillate inflorescenceoften fused in pairs at the
base; lithophytic,Tolima ............................... 16. C. goudotiana
54. Lateralveins in the free partof the midsegment(8-)10-
15(-17) pairs; spikes of the staminateinflorescence10-
60; spikes of the pistillateinflorescencefree at the base;
not lithophytic,widespread....................................... 41. C. peltata
53. Stipules 8-35 cm long.
55. Stipules sparsely hairy to subglabrousoutside,
(usually) green; Amazonianregion, inundatedplaces
........................................................................3 1 .C. membranacea
55. Stipules more or less densely hairy outside, often
reddishor whitish.
56. Stipules 15-25(-35) cm long; Amazonianregion
..........................................................................12 .C. ficifolia
56. Stipules 3-12 cm long; Pacific region, Cauca
valley (to Tolima) and N part of the country.... 41. C. peltata

Key to the species of Venezuela

I. Trichiliaabsent.
2. Laminawith 10-15 segments;lowland, Amazonas, Bolivar,and Delta Amacuro....................... 51. C. sciadophylla
2. Laminawith 7-10 segments;montane,Andes ....................... ........................................ 57. C. telenitida
1. Trichiliapresent.
3. Arachnoidindumentumabsent in the areoles of the lamina beneath.
4. Laminachartaceousto subcoriaceous;stipules and spathesmostly greenish;spikes of staminate
inflorescenceca. 0.2 cm thick; pistillateinflorescencependulousat anthesis;Amazonas, inundated
places....................................................... 31. C. membranacea
4. Lamina(sub)coriaceous;stipules and spathesred; spikes of the staminateinflorescence0.3-0.4(-l)
cm thick;pistillate inflorescenceerect at anthesis (to pendulousin fruit);Amazonasand Bolivar,
non-inundatedplaces............................................................ 9. C. distachya
3. Arachnoidindumentumpresentin the areoles of the laminabeneath.
5. Stipules usually 15-35 cm long; indumentum(of the leafy twig) without uncinatehairs;Amazonas
and Bolfvar,(sub)montane....................................................... 24. C. kavanayensis
5. Stipules usually to 15 cm long; indumentum(of the leafy twigs) with some of the hairs uncinate.
6. Lateralveins of the free partof the midsegmentsubmarginallyloop-connected.
7. Laminawith 6-7(-8) segments, the incisions down to ca. 5/10-9/10 from the margin;free
partof the midsegmentspathulateto obovate and the lateralveins mostly unbranched;
Amazonas.................................................. 12. C.fcifolia
7. Laminawith (7-)8-11 segments, the incisions down to ca. >5/10 from the margin,the free
partof the midsegmentoblong to elliptic to obovate and the lateralveins usually branched.
8. Lamina(sub)coriaceousand mostly smooth above; peduncle and spatheof the
inflorescencesusually >10 cm long; stigma comose and apex of the perianthof the
pistillate flower without arachnoidindumentum;Bolivar and Delta Amacuro,lowland
.........................................................................................................................................3. C. angulata
8. Laminachartaceousto subcoriaceousand ? scabrousabove; peduncle and spatheof
the inflorescencesusually <10 cm long; stigma peltate and apex of the perianthof the
pistillate flower with arachnoidindumentum;widespread,lowland (to montane)........ 41. C. peltata
6. Lateralveins of the free partof the midsegmentmarginallyloop-connected.
9. Lateralveins in the free partof the midsegment9-17 pairs.
10. Petiole with dense arachnoidindumentum;the brown indumentumof the trichilium
intermixedwith distinctly longer white hairs;free partsof the lamina segments often
? lobate; llanos and savannaregions, widespread.............. .................... 32. C. metensis
38 FLORANEOTROPICA

10. Petiole without or with sparse arachnoidindumentum;the brown indumentumof the

trichiliumintermixedwith about equally long (or rarelylonger) white hairs;free parts
of the lamina segment entire;forested regions and riversides.
11. Lamina(sub)coriaceousand usually smooth above; lowland, Amazonas and
Bolivar ............................................................. 9. C. distachya
11. Laminachartaceousto subcoriaceous.
12. Laminaattachedto the petiole at ca. 1/4 of the total length from its base;
stipules and spatheswithout arachnoidindumentumoutside; stigma
penicillate;Amazonas, Apure, Bolivar, and Delta Amacuro,inundatedplaces
......................................................................................................................25 . C. latiloba
12. Laminaattachedto the petiole at clearly >1/4 of the total length from its
base; stipules and spathesoften with (dense) arachnoidindumentumoutside;
stigma peltate;widespread,non-inundatedplaces........................................ 41. C. peltata
9. Lateralveins in the free partof the midsegmentusually 18-32 pairs.
13. Lateralveins in the free partof the midsegment24-32 pairs;Amazonas, lowland
..................................................................................................................................11. C. engleriana
13. Lateralveins in the free partof the midsegment 18-24(-25) pairs;Andes.
14. Spikes of the staminateinflorescenceca. 15-20; peduncle of the pistillate
inflorescence2-5 cm long and the spikes 3.5-7.5 cm long........................... 4. C. angustifolia
14. Spikes of the staminateinflorescence4-8; peduncle of the pistillateinflorescence
8-14 cm long and the spikes 11-33 cm long ..................................................48. C. sararensis

Key to the species of the Guianas

1. Trichiliaabsent;incisions of the lamina down to the petiole and the segments ? clearly petiolulate
.51.
.............................................................................................................................................................. C. sciadophylla
l. Trichiliapresent;incisions of the lamina not down to the petiole and the segments not petiolulate.
2. Laminawith (12-)15-17 segments;lateralveins in the free partof the midsegmentca. 40-45 pairs;
FrenchGuianaand Suriname..................................................................... 52. C. silvae
2. Laminawith 7-11 segments;lateralveins in the free partof the midsegmentca. 10-20 pairs.
3. Trichiliawith whitish hairs 5-9 mm long; FrenchGuiana(inselbergs) ........... ...................... 17. C. granvilleana
3. Trichiliawith whitish hairs much shorter.
4. Lateralveins loop-connectedin the marginof the lamina.
5. Stipules subpersistent;peduncle of the pistillate inflorescence20-40 cm long; staminate
inflorescenceswith 4-6(-1 1) spikes, these 0.8-1.8 cm diam.; FrenchGuianaand E
Suriname............... ............................................... 39. C. palmata
5. Stipules caducous;peduncle of the pistillate inflorescenceto 16 cm long; staminate
inflorescenceusually with > 12 spikes, these 0.2-0.4 cm diam.
6. Lamina(usually) smooth above, (sub)coriaceous;FrenchGuiana.............................. 9. C. distachya
6. Laminascabrousabove, chartaceousto subcoriaceous.
7. Stipules and spathesreddish,without arachnoidindumentumoutside; stigma
penicillate;widespread,in periodicallyinundatedplaces ....................... ................25. C. latiloba
7. Stipules and spathes (usually) greenish or, if reddish,then with + dense arachnoid
indumentumoutside; stigma peltate;Guyanaand Suriname,in non-inundatedplaces
....................................................................................................................................41. C. peltata
4. Lateralveins loop-connected(just) inside the marginof the lamina.
8. Stipules 12-20 cm long; Guyana................................................................... 3. C. angulata
8. Stipules (usually) 3-12 cm long.
9. Upper surfaceof the lamina with ? dense arachnoidindumentum;stigma comose;
widespread.............................................................. 36. C. obtusa
9. Upper surfaceof the lamina without arachnoidindumentum;stigma peltate;Guyanaand
Suriname.............................................................. 41. C. peltata

Key to the species of Ecuador

1. Trichiliaenclosed in 2 pockets with slit-shapedaperturesat the base of the petiole; coastal region ........ 61. C. virgusa
1. Trichiliaexposed (superficial)or absent.
2. Trichiliaabsent.
SYSTEMATICTREATMENT 39

3. Laminausually with 10-15 segments;lateralveins in the free part of the midsegmentca. 25-40
(-45).
4. Leafy twigs hairy;coastal region.................................................................. 20. C. hispidissima
4. Leafy twigs (sub)glabrous;Amazonianregion................................................................. 51. C. sciadophylla
3. Laminausually with 7-9 segments;lateralveins in the free partof the midsegment 10-18 pairs;
montane.
5. Upper surfaceof the lamina smooth, subglabrous;spikes of the staminateinflorescence3-6;
spikes of the pistillateinflorescence 1-2; Andes, W slopes.................................................... 13. C. gabrielis
5. Upper surfaceof the lamina ? scabrous;spikes of the staminateinflorescence5-16; spikes of
the pistillate inflorescence(3-)4-5 spikes; Andes, E slopes.
6. Lamina(sub)coriaceous,the uppersurfaceof the lamina with dense arachnoidindumentum;
main veins sparselyhairy beneath............................................................. 57. C. telenitida
6. Laminachartaceousto subcoriaceous,the uppersurfacewithout arachnoidindumentum;
smaller (and often also larger)veins beneath ? densely hairy ................. ............................ 2. C. andina
2. Trichiliapresent(sometimes poorly developed).
7. Lamina(usually) with 10-20(-24) segments;lateralveins in the free partof the midsegmentmostly
>28 pairs.
8. Setose (irritating)hairs presenton young parts;coastal region.
9. Lamina(usually) ? scabrousabove; trichilia well-defined;Mulllerianbodies white, ca. 1
mm long ............................................................. 14. C. garciae
9. Laminasmooth above; trichilianot well-defined;Mullerianbodies pink, ca. 3 mm long
...................................................................................................................................... .20. C. hispidissima
8. Setose hairs absent.
10. Lateralveins in the free partof the midsegment50-80 pairs;Andes, E slopes.......... 60. C. velutinella
10. Lateralveins in the free partof the midsegment 15-50 pairs.
11. Laminasmooth above.
12. Laminaoften white above, segments of young leaves patent;spikes of staminate
inflorescence4-6(-8); spikes of pistillate inflorescence(1-)2-3, the spathe 10-
20 cm long........................................ 29. C. maxima
12. Laminagreen above, segments of young leaves deflexed; spikes of staminate
inflorescence 10-25(-30); spikes of pistillateinflorescence3-8(-12), the spathe
3-6(-8) cm long .............................................. 4. C. angustifolia
11. Lamina ? scabrousabove.
13. Lateralveins in the free partof the midsegmentusually 30-50 pairs;spathes and
stipules with dense arachnoidindumentumoutside; Andes, E slopes to ca. 1000
m and Amazonianregion ....................................... 33. C. montana
13. Lateralveins in the free partof the midsegmentusually (I0-)15-25 pairs, or if
to 35(-43) pairs, then the spathes and stipules without dense arachnoid
indumentumoutside.
14. Spathesof the inflorescencessparselyhairy to subglabrousoutside; lamina
erect in the bud; coastal region .......... ......................... 37. C. obtusifolia
14. Spathes ? densely hairy outside; lamina often reflexedin the bud; Andean
region.................................... 4. C. angustifolia
7. Laminawith (6-)7-10(-l 1) segments;lateralveins in the free partof the midsegmentmostly <25
pairs.
15. Laminawithout arachnoidindumentumin the areoles beneath.
16. Base of the lamima entire or shallowly lobed, the upperpart3-lobed.
17. Spikes of the staminateinflorescence4-5; peduncle of the pistillateinflorescence25-
45 cm long; Amazonianregion.......................................... 46. C. putumayonis
17. Spikes of the staminateinflorescence7-12; peduncleof the pistillate inflorescence3-
24 cm long; coastal region.................................................. 54. C. subintegra
16. Base of the lamina ? distinctly incised.
18. Lateralveins in the free partof the midsegmentusually 15-25(-40) pairs;coastal
region.............................................. 37. C. obtusifolia
18. Lateralveins in the free partof the midsegment9-16 pairs.
19. Spikes of the staminateinflorescenceca. 30-40; peduncle of the pistillate
inflorescence5-15(-20) cm long; stipules mostly greenish;Amazonianregion
(and coastal region), usually inundatedplaces...................................... 31. C. membranacea
19. Spikes of the staminateinflorescence4-7(-15); peduncle of the pistillate
40 FLORANEOTROPICA

inflorescence 15-50 cm long; stipules mostly reddish;Amazonianregion (to

Andean region), non-inundatedplaces .............................................. 59. C. utcubambana
15. Laminawith arachnoidindumentumin the areoles beneath.
20. Trichiliaseparate.
21. Trichiliaon a subscrotiform(abaxiallybulging) base of the petiole; Andes, E slopes
..............................................................................................................................40. C. pastasana
21. Trichilianot on a subscrotiformbase of the petiole.
22. Leafy twigs glabrousand smooth, mostly with a bluish waxy layer; Andes.
23. Spikes of the staminateinflorescence3-6; spikes of the pistillate
inflorescence 1-2; Andes, W slopes ................................................. 13. C. gabrielis
23. Spikes of the staminateinflorescence(5-)6-16; spikes of the pistillate
inflorescence3-5; Andes, E slopes.
24. Laminachartaceousto subcoriaceous,without arachnoidindumentum
above............................................. 2. C. andina
24. Lamina(sub)coriaceous,with (often dense) arachnoidindumentum
above............................................. 57. C. telenitida
22. Leafy twigs hairy and often ? scabrous,green or reddish-brown;lowlands.
25. Petiole as long as ca. half the diameterof the lamina;spikes of staminate
inflorescence(20-)30-50; stipes of the spikes of the pistillate inflorescence
to 0.5 cm long; Amazonianregion ................................................. 28. C. marginalis
25. Petiole much longer than half the diameterof the lamina;spikes of the
staminateinflorescence5-9; stipes of the spikes of the pistillate
inflorescence0.5-1 cm long; coastal region............................................ 23. C. insignis
20. Trichiliafused.
26. Trichiliaon a subscrotiform(abaxiallybulging) base of the petiole.
27. Trichilialaterallyextendedby lobes; spikes of the staminateinflorescence4-
12, 6-12 cm long; spatheof the pistillate inflorescence5-9 cm long; Andes, W
slopes .................................................... 47. C. reticulata
27. Trichilialaterallyrounded;spikes of the staminateinflorescence 10-25, (6-)12-
28 cm long; spathe of the pistillate inflorescence 10-17 cm long; Andes, E
slopes .................................................... 40. C. pastasana
26. Trichilianot on a subscrotiformbase of the petiole.
28. Lateralveins of the free partof the midsegmentmarginallyloop-connected.
29. Lateralveins in the free partof the midsegment21-32 pairs;Amazonian
region.1................................................. l.C. engleriana
29. Lateralveins in the free partof the midsegment 12-19 pairs.
30. Lamina(sub)coriaceous,usually smooth above; Amazonianregion,
non-inundatedplaces............................................. 9. C. distachya
30. Laminachartaceousto subcoriaceous, ? scabrousabove.
31. Laminaattachedto the petiole at ca. 1/4 of the total length from
its base; spikes of the staminateinflorescenceusually ca. 15-20,
usually 14-20 cm long; stigma penicillate;Amazonianregion,
inundatedplaces .......... ............................... 25. C. latiloba
31. Laminaattachedto the petiole at clearly >1/4 of the total length
from its base; spikes of the staminateinflorescenceusually ca. 5-
15, usually 5-12 cm long; stigma peltate;coastal region, usually
in non-inundatedplaces, and Amazonianregion, in inundated
places .......................................... 26. C. litoralis
28. Lateralveins of the free partof the midsegmentsubmarginallyloop-connected.
32. Lamina ? scabrousabove; Amazonianregion ......................... ..............12. C.ficifolia
32. Laminasmooth above.
33. Laminawith (often dense) arachnoidindumentumabove and leafy
twigs either glabrousor villous; Andes, E slopes .............. ..........57. C. telenitida
33. Laminawithout arachnoidindumentumabove or, if with that
indumentum,then the leafy twigs hispidulous.
34. Incisions of the lamina down to 3/10-6/10 from the margin.
35. Upper surfaceof the lamina usually bullate;spikes of the
staminateinflorescence4-10; spikes of the pistillate
inflorescence 1-2; Andes, W slopes ................................... 6. C. bullata
SYSTEMATICTREATMENT 41

35. Upper surfaceof the lamina smooth; spikes of the staminate

inflorescence20-50; spikes of the pistillate inflorescence4
(-7); Amazonianregion ..................................... 28. C. marginalis
34. Incisions of the lamina (usually) down to 6/10-9/10 from the
margin;upper surface smooth; spikes of the pistillate
inflorescence(usually) 3-8.
36. Spikes of the staminateinflorescence5-9; spikes of the
pistillate inflorescence8-18 cm long; coastal region...... 23. C. insignis
36. Spikes of the staminateinflorescence 10-50; spikes of the
pistillate inflorescence3-8(-12) cm long; Andes and coastal
region..................................... 4. C. angustifolia

Key to the species of Peru

1. Trichiliaabsent.
2. Incisions of the laminadown to the petiole and the segments often petiolulate;Amazonianregion
........................................................................................................................................................
...51. C. sciadophylla
2. Incisions of the laminanot down to the petiole; Andean region.
3. Laminawith 7-10(-1 1) segments.
4. Laminachartaceousto subcoriaceous,without arachnoidindumentumabove ............ ............... 2. C. andina
4. Lamina(sub)coriaceous,with (often dense) arachnoidindumentumabove.......................... 57. C. telenitida
3. Laminawith 10-1 8 segments.
5. Lateralveins in the midsegmentof the lamina 15-35 pairs;uppersurfaceof the lamina ?
scabrous;Andes............................................................... 55. C. tacuna
5. Lateralveins in the midsegmentof the lamina 10-16 pairs;the uppersurfaceof the lamina
smooth; Andes.............................................................. 1. C. albicans
1. Trichiliapresent.
6. Trichiliaseparate(in 2 lateralpatches).
7. Trichiliaon a subscrotiform(abaxiallybulging) base of the petiole; Andes.............................. 40. C. pastasana
7. Trichilianot on a subscrotiformbase of the petiole.
8. Lamina(sub)coriaceous,with (often dense) arachnoidindumentum(and thus usually white)
above; Andes ............................................................... 57. C. telenitida
8. Laminachartaceousto subcoriaceous,without arachnoidindumentumabove.
9. Leafy twigs (sub)glabrous,smooth, with a bluish waxy layer; Andes .............. .................... 2. C. andina
9. Leafy twigs hairy.
10. Petiole as long as ca. half the diameterof the lamina;terminalbuds usually curved, not
inflated;Amazonianregion ........................................................ 28. C. marginalis
10. Petiole much longer than half the diameterof the lamina;terminalbuds straight, ?
inflated;Amazonianregion to Andes ......................... ............................... 53. C. strigosa
6. Trichiliafused.
11. Laminawith 10-20(-24) segments.
12. Terminalbuds + inflated;lateralveins in the free partof the midsegment8-17 pairs.
13. Laminacoriaceous, with ? dense arachnoidindumentumand smooth above; Andes
(2200-2500 m) ............................................................................ 1. C. albicans
13. Laminachartaceousto subcoriaceousand without arachnoidindumentumand scabrous
or smooth above; Amazonianregion to Andes (to 1900 m) 53. C. strigosa
12. Terminalbuds not inflated;lateralveins in the free partof the midsegment 15-80 pairs.
14. Lateralveins in the free partof the midsegment50-80 pairs;Andes ........... ........... 60. C. velutinella
14. Lateralveins in the free partof the midsegment 14-50 pairs.
15. Incisions of the lamina down to the petiole or at most 2 cm from the petiole;
Amazonianregion ............................................................. 18. C. herthae
15. Incision of the lamina down to >2 cm from the petiole.
16. Lateralveins in the free partof the midsegmentusually 30-50 pairs;leafy
twig whitish, green, or brownish;Amazonianregion.................................. 33. C. montana
16. Lateralveins in the free partof the midsegmentca. 15-25 pairs, or if to 35
(-43) pairs, then the leafy twig usually turningblackish.
17. Upper segments of the lamina lobate; lamina erect in the bud; peduncle
of the inflorescencesusually >8 cm long; Amazonianregion (to Andes,
to 1350 m)44.................................................. C. polystachya
42 FLORANEOTROPICA

17. Upper segments of the lamina entire;lamina often reflexedin the bud;
peduncle of the inflorescencesusually to 8 cm long; Andes (mostly 1200-
2400 m, sometimes down to 400 m) ....................... ......................... 4. C. angustifolia
11. Laminawith 7-10(-1 1) segments.
18. Laminawithout arachnoidindumentumin the areoles beneath.
19. Base of the lamima entire or shallowly lobed, the upperpart3-lobed; Amazonianregion
.................................................................................................................................46. C. putumayonis
19. Base of the lamina ? distinctlyincised.
20. Spikes of the staminateinflorescenceca. 4-7(-15); peduncle of the pistillate
inflorescence 15-50 cm long; stipules mostly reddish;Amazonianregion, non-
inundatedplaces........................................................ 59. C. utcubambana
20. Spikes of the staminateinflorescenceca. 25-50; peduncle of the pistillate
inflorescenceca. 5-15(-20) cm long; stipules mostly greenish.
21. Terminalbuds ? inflated;stipes of spikes of the staminateinflorescence(0.5-)
1-2.5 cm long; spikes of the pistillate inflorescence3-7(-9); Amazonian
region to Andes (to 1900 m), non-inundatedplaces .................................... 53. C. strigosa
21. Terminalbuds not inflated;stipes of spikes of the staminateinflorescence0.5-
1 cm long; spikes of the pistillate inflorescence(2-)4; Amazonianregion,
usually inundatedplaces................................................................ 31. C. membranacea
18. Laminawith arachnoidindumentumin the areoles beneath.
22. Lateralveins in the free partof the midsegmentmarginallyloop-connected.
23. Lateralveins in the free partof the midsegment24-32 pairs.
24. Inflorescencespendulous,the peduncle usually 10-19 cm long; Amazonian
region ................................................. IIC. engleriana
24. Inflorescenceserect, the peduncle usually 1-8 cm long; Andes (mostly 1200-
2200 m)........................................................ 4. C. angustifolia
23. Lateralveins in the free partof the midsegment 12-24 pairs.
25. Incisions of the lamina down to 1.5 cm from the petiole.
26. Inflorescencespendulous,the peduncles usually 10-22 cm long;
Amazonianregion .............................................. 8. C concolor
26. Inflorescenceserect, the peduncles usually 1-8 cm long; Andes to
Amazonianregion ([400-]1200-2400 m) . ...........................4. C. angustifolia
25. Incisions of the lamina down to at most 4 cm from the petiole.
27. Lamina(sub)coriaceous,usually smooth above.
28. Free partsof the uppersegments of the laminaobovate to
subobovate;Amazonianregion ........................................... 9. C. distachya
28. Free partsof the uppersegments of the lamina oblanceolate;Andes
(mostly 1200-2400 m) .................................................... 4. C. angustifolia
27. Laminachartaceousto subcoriaceous,mostly ? scabrousabove.
29. Spatheswith dense arachnoidindumentum;peduncles usually 10-20
cm long, and lateralveins in the free partof the midsegment21-32
pairs............................................ 11. C. engleriana
29. Spathes without or with sparse arachnoidindumentum;peduncles to
10 cm long, and/orlateralveins in the free part of the midsegment
to 20 pairs.
30. Laminaattachedto the petiole at ca. 1/4 of the total length
from its base; spikes of the staminateinflorescenceusually ca.
15-20, usually ca. 14-20 cm long; stigma penicillate;
Amazonianregion, inundatedplaces..................................... 25. C. latiloba
30. Laminaattachedto the petiole at clearly > 1/4 of the total
length from its base; spikes of the staminateinflorescence
usually ca. 5-15 and/orusually 5-12 cm long; stigma peltate
(lowland) or comose (montane).
31. Stipules and spatheschartaceousto membranaceouswhen
dry, sparsely hairy; stigma peltate;Amazonianregion,
inundatedplaces ........... .......................... 26. C. litoralis
31. Stipules and spathes subcoriaceouswhen dry, densely
hairy; stigma comose; Andes (mostly 1200-2400 m)
. 4 . C angustifolia
..................................................................................
SYSTEMATICTREATMENT 43

22. Lateralveins in the free partof the midsegmentsubmarginallyloop-connected.

32. Trichiliaon subscrotiform(abaxiallybulging) base of the petiole.
33. Laminawith ? dense arachnoidindumentumabove; spikes of the staminate
inflorescence 10-25; Andes (900-2300 m) ............................................... 40. C. pastasana
33. Laminawithout arachnoidindumentumabove; spikes of the staminate
inflorescenceca. 25-100; Amazonianregion and Andes (to 1900 m) ........ 53. C. strigosa
32. Trichilianot on a subscrotiformbase of the petiole.
34. Leafy twigs (sub)glabrous,smooth, bluish by a waxy layer; Andes ............... 2. C. andina
34. Leafy twigs ? hairy.
35. Upper lamina segments lobate; Amazonianregion and Andes (to 1350 m)
....................................................................................... 44. C. polystachya
35. Upper lamina segment entire.
36. Lamina(sub)coriaceous.
37. Stipules subpersistent;Mulllerianbodies pinkish when fresh;
spikes of the staminateinflorescence7-8; spikes of the
pistillate inflorescence4, often curved;Andes .............. 7. C. chlorostachya
37. Stipules caducous, or if subpersistent,then the Mullerianbodies
white, the spikes of the staminateinflorescence(usually) 10-
25(-40), and the spikes of the pistillate inflorescenceoften >4,
straight,and often thickenedtowardthe base.
38. Lateralveins in the free partof the midsegmentusually 12-
18 pairs; spikes of the pistillate inflorescence2-4, usually
8-15 cm long at anthesis;Amazonianregion............... 9. C. distachya
38. Lateralveins in the free partof the midsegmentusually 15-
35 pairs; spikes of the pistillate inflorescenceoften >4, 1-
7 cm long at anthesis;Andes .................................... 4. C. angustifolia
36. Laminachartaceousto subcoriaceous.
39. Petiole as long as ca. half the diameterof the lamina;stipules
ca. 15-45 cm long, glabrousinside; terminalbuds usually
curved, glabrousinside; Amazonianregion...................... 28. C. marginalis
39. Petiole much longer than half the diameterof the lamina;
stipules often < 15 cm long, or if longer,then often hairy inside;
terminalbuds straight.
40. Lateralveins in the free partof the midsegment
unbranched;incisions down to ca. 5/10, or if more deeply,
then the midsegmentbroadlyspathulate;spikes of the
staminateinflorescence8-13 and the peduncle of the
staminateinflorescenceusually 10-20 cm long;
Amazonianregion ........... ........................ 12. C. ficifolia
40. Lateralveins in the free partof the midsegmentmostly
branched;incisions down to 5/10-8/10, the midsegment
obovate, elliptic, subobovate,oblong, or oblanceolate;
spikes of the staminateinflorescence4-7 or ca. 25-100.
41. Lateralveins in the free partof the midsegment
usually ca. 20-35 pairs, the midsegmentusually
oblanceolate;lamina often reflexedin the bud; Andes
(mostly 1200-2400 m) ................................ 4. C. angustifolia
41. Lateralveins in the free partof the midsegment
usually 8-17 pairs, the midsegmentusually obovate
to elliptic or subobovate;lamina erect in the bud;
lowland to submontane.
42. Terminalbuds + inflated;lamina often ?
bullate above; indumentumof the leafy twig
puberulousto hirtellousor to subhirsute;with
straighthairs;Amazonianregion to Andes (to
1900 m)........................... 53. C. strigosa
42. Terminalbuds not inflated;laminanot bullate
above; indumentumof the leafy twig partly
44 FLORA NEOTROPICA

(sub)setuloseor puberulousto hispidulouswith

uncinatehairs;Amazonianregion.
43. Spikes of the staminateinflorescenceca. 30-
40; peduncleof the pistillateinflorescence5-
15(-20) cm long; leafy twig partly
(sub)setulose;Amazonianregion, usually
inundatedplaces .... ..... 31. C. membranacea
43. Spikes of the staminateinflorescence4-7
(-15); peduncle of the pistillate
inflorescence 15-50 cm long; leafy twig not
(sub)setulose;Amazonianregion, non-
inundatedplaces .... ..... 59. C. utcubambana

Key to the species of AmazonianBrazil

1. Trichiliaabsent;incisions of the lamina down to the petiole and the segments ? clearly petiolulate
.51.
.............................................................................................................................................................. C. sciadophylla
1. Trichiliapresent;incisions of the lamina not down to the petiole and the segments not petiolulate.
2. Laminawith 12-15 segments.
3. Lateralveins in the free partof the midsegmentca. 40-45 pairs;NE Amazon basin .......... ............ 52. C. silvae
3. Lateralveins in the free partof the midsegmentca. 20-25 pairs;NE to centralAmazon basin......... 58. C. ulei
2. Laminawith up to 10(-12) segments.
4. Laminaincised down to 1.5(-2.5) cm from the petiole; lower, centraland SW Amazon basin.... 8. C. concolor
4. Laminaincised down to at most 4 cm from the petiole.
5. Lateralveins of the free partof the midsegmentmarginallyloop-connected.
6. Lateralveins in the free part of the midsegmentof the lamina (21-)24-32 pairs;western
Amazon basin.1................................................................ L C. engleriana
6. Lateralveins in the free partof the midsegment 10-17 pairs.
7. Laminaattachedto the petiole at ca. 1/4 of the total length from its base, chartaceousto
subcoriaceous,scabrousabove; stipules caducous;widespread,inundatedplaces ...... 25. C. latiloba
7. Laminaattachedto the petiole clearly > 1/4 of its total length from its base,
(sub)coriaceous,smooth above, or if + scabrous,then the stipules often subpersistent;
widespread,non-inundatedplaces.
8. Inflorescencespendulousat anthesis;spathes white due to dense arachnoid
indumentumoutside; stipules often subpersistent.................................................. 39. C. palmata
8. Inflorescenceserect (at least at anthesis);spathesnot white, without dense arachnoid
indumentumoutside; stipules caducous.................................................................. 9. C. distachya
5. Lateralveins of the free partof the midsegmentsubmarginallyloop-connected.
9. Indumentum(of the leafy twig) without uncinatehairs;Roraima,submontane....... 24. C. kavanayensis
9. Indumentum(of the leafy twig) with some of the hairs uncinate.
10. Lower surface of the lamina without arachnoidindumentumin the areoles.
11. Lamina(sub)coriaceous;stipules red; widespread,non-inundatedplaces......... 9. C. distachya
11. Laminachartaceous(to subcoriaceous);stipules mostly green; widespread,
inundatedplaces .............. ............................................ 31. C. membranacea
10. Lower surfaceof the laminawith arachnoidindumentumin the areoles.
12. Lamina(sub)coriaceous;petiole and usually also the uppersurfaceof the lamina
with dense arachnoidindumentum;lower to centralAmazon basin ........... ......... 36. C. obtusa
12. Laminachartaceousto subcoriaceous;petiole and usually also the uppersurface
of the lamina without or with sparse arachnoidindumentum.
13. Midsegmentof the lamina oblong to elliptic; lateralveins in the midsegment
usually branched;spathes with ? dense arachnoidindumentumoutside;N
fringe of the Amazon basin................................................ 41. C. peltata
13. Midsegmentof the lamina broadlyspathulateto obovate or ovate; lateral
veins of the midsegmentusually unbranched;spatheswithoutor with sparse
arachnoidindumentumoutside.
14. Midsegmentof the lamina ovate; stipules 8-16 cm long; central
Amazon basin............................................ 45. C. purpurascens
14. Midsegmentof the laminabroadlyspathulateto obovate;stipules
usually ca. 15-25 cm long; W (to central)Amazon basin .................. 12. C. ficifolia
SYSTEMATICTREATMENT 45

Key to the species of Bolivia

1. Trichiliaabsent;incisions of the lamina down to the petiole and the segments ? clearly petiolulate
.51.
.............................................................................................................................................................. C. sciadophylla
1. Trichiliapresent;incisions of the lamina not down to the petiole and the segments not petiolulate.
2. Trichiliaseparate(2 lateralpatches); submontane........................................................................ 53. C. strigosa
2. Trichiliafused.
3. Laminawith 10-16 segments;montane.
4. Spikes of the inflorescences 18-40 cm long at anthesis ..............................................0...........I. C. elongata
4. Spikes of the inflorescences 1.5-13 cm at anthesis,in fruit to 17 cm long......................... 4. C. angustifolia
3. Laminawith (5-)7-10(-12) segments;lowland (or montane).
5. Lateralveins of the free partof the midsegmentmarginallyloop-connected.
6. Lateralveins in the free partof the midsegmentusually 24-32 pairs;lowland, widespread
..........................................................................................................................................11. C. engleriana
6. Lateralveins in the free partof the midsegment8-24 pairs.
7. Incisions of the lamina down to 1.5(-2.5) cm from the petiole; lowland (widespread)
.........................................................................................................................................8. C. concolor
7. Incisions of the lamina down at most to ca. 4 cm from the petiole.
8. Lateralveins in the free partof the midsegment8-10 pairs;lowland (SantaCruz)
.................................................................................................................................49. C saxatilis
8. Lateralveins in the free partof the midsegment 10-20 pairs.
9. Laminaattachedto the petiole at ca. 1/4 of the total length from its base,
chartaceous;lowland, inundatedplaces (Beni and Pando)................................ 25. C. latiloba
9. Laminaattachedto the petiole at clearly > 1/4 of its total length from its base,
subcoriaceousto coriaceous;lowland, non-inundatedplaces.
10. Stipules with dense arachnoidindumentumoutside; trunkwith prominent
stipularscars;lowland (La Paz) ................................. ................... 5. C. annulata
10. Stipules without or with sparse arachnoidindumentumoutside; trunk
withoutprominentstipularscars.
11. Inflorescencespendulous;spatheswith dense arachnoidindumentum
outside; stipules often subpersistent;lowland (SantaCruz)............. 39. C. palmata
11. Inflorescencesinitially erect, pendulousin fruit;spatheswithout or
with sparse arachnoidindumentumoutside; stipules always caducous;
lowland (La Paz and SantaCruz) ................................................ 9. C. distachya
5. Lateralveins in the free partof the midsegmentsubmarginallyloop-connected.
12. Laminawithout arachnoidindumentumin the areoles beneath.
13. Spikes of the staminateinflorescenceca. 4-7(-15); peduncle of the pistillate
inflorescence 15-50 cm long; stipules mostly reddish;lowland (Pando)........ 59. C. utcubambana
13. Spikes of the staminateinflorescenceca. 25-50; peduncle of the pistillate
inflorescenceca. 5-15(-20) cm long; stipules mostly greenish.
14. Terminalbuds ? inflated;stipes of spikes of the staminateinflorescence(0.5-)1-
2.5 cm long; spikes of the pistillate inflorescence3-9; submontane............... 53. C. strigosa
14. Terminalbuds not inflated;stipes of spikes of the staminateinflorescence0.5-1
cm long; spikes of the pistillate inflorescence(2-)4; lowland, usually inundated
places ................................................... 31. C. membranacea
12. Laminawith arachnoidindumentumin the areoles beneath;non-inundatedplaces.
15. Upper segments of the lamina lobate; lowland (widespread)............. ............... 44. C. polystachya
15. Upper segments of the lamina entire.
16. Lamina(sub)coriaceous;spikes of the staminateinflorescence4-8, the peduncle
3-9 cm long; spathe of the pistillate inflorescence6-8 cm long; lowland (Santa
Cruz) .................................................... 49. C. saxatilis
16. Laminachartaceousto subcoriaceous;spikes of the staminateinflorescence8-
100, or if <8, then the peduncle usually 10-21 cm long; spatheof the pistillate
inflorescence8-18(-23) cm long.
17. Terminalbuds ? inflated;lateralveins of the midsegmentmostly branched;
spikes of the staminateinflorescence(15-)25-100; submontane............ 53. C. strigosa
17. Terminalbuds not inflated;lateralveins of the midsegmentoften
unbranched;spikes of the staminateinflorescence8-15; lowland (Beni and/
or Pando).
46 FLORANEOTROPICA

18. Stipules (12-)18-25(-35) cm long; midsegmentof the lamina usually

broadlyspathulate......... ................................... 12. C.ficifolia
18. Stipules 4-12 cm long; midsegmentof the lamina oblong to obovate
...................................................................................................59. C. utcubambana

Key to the species of Extra-AmazonianBrazil, Paraguay,and Argentina

1. Trichiliaabsent;uppersurfaceof the lamina white due to dense arachnoidindumentum;E Brazil...... 21. C. hololeuca
1. Trichiliapresent;uppersurface of the lamina not white, without dense arachnoidindumentum.
2. Lateralveins in the free partof the midsegment8-10 pairs;centralBrazil............... ......................... 49. C. saxatilis
2. Lateralveins in the free partof the midsegment 10-20(-23) pairs.
3. Stipules white to greenish, with + dense arachnoidindumentumoutside; stigma peltate, and
arachnoidindumentumcovering the apex of the perianthof the pistillate flower;anthers0.5-0.7
mm long; spikes of the staminateinflorescenceoften >10 and often <1 cm diam.; Brazil
(widespread),Argentina,and Paraguay............................. .................................... 38. C. pachystachya
3. Stipules usually red(dish),withoutor with sparse arachnoidindumentumoutside; stigma comose to
penicillate, and arachnoidindumentumnot covering the apex of the perianthof the pistillateflower;
anthers0.8-2 mm long; spikes of the staminateinflorescenceoften <10, often >1 cm diam.
4. Spathes with dense arachnoidindumentumoutside (white); NE Brazil (Paraiba,Pernambuco,
and Sergipe)............................................................... 39. C. palmata
4. Spathes without or with sparse arachnoidindumentumoutside (reddish);Bahia to Rio Grande
do Sul .............................................................. 15. C. glaziovii

Key to the species of the Cecropiapeltata-group

1. Trichiliaabsent or poorly developed (in 2 patches and/orwithout Mullerianbodies).
2. Internodeswith ample brownpith; Cuba, Hispaniola,PuertoRico, Lesser Antilles ......... .........50. C. schreberiana
2. Internodeswith scarce whitish pith;Jamaica.......................................................................... 41. C. peltata
1. Trichiliapresentand well-developed (fused).
3. Incisions of the lamina (at least some) down to near (at most 1.5 cm or rarely2.5 cm from) the petiole;
lower to SW Amazon basin ........................................................................ 8. C concolor
3. Incisions down to 9/10, at least 2 cm from the petiole.
4. Lateralveins in the free partof the midsegment(21-)24-32 pairs;upperAmazon basin, from
Bolivia to Colombia .................................................................... 11. C. engleriana
4. Lateralveins in the free partof the midsegment9-20(-24) pairs.
5. Trichiliawith white (or brownish)unicellularhairs ? dense and distinctly longer than the
brown indumentum.
6. Stipules and mostly also the petiole with dense arachnoidindumentum.
7. Stipules to 13 cm long; lamina segments sometimes lobate; savannasfrom Colombiato
Guyana............................................................... 32. C. metensis
7. Stipules to 22 cm long; lamina segments often lobate;extra-AmazonianBrazil,
Paraguay,Argentina,Paraguay,and S fringe of the Amazon basin..................... 38. C. pachystachya
6. Stipules and petiole withoutor with sparse arachnoidindumentum.
8. Stipules 3-4 cm long; lateralveins in the free partof the midsegment9-10 pairs;
French Guiana,inselbergs......................................................... 17. C. granvilleana
8. Stipules 3-10(-12) cm long; lateralveins in the free partof the midsegment(8-)10-15
(-17) pairs;Jamaica,CentralAmerica, N South America, eastwardto Suriname,
Trinidad,Tobago, N fringe of AmazonianBrazil (Para,Roraima)............. .................. 41. C. peltata
5. Trichiliawith (inconspicuous)short white hairs.
9. Stipules usually to 10 cm long and spathesusually to 7 cm long; segments of the lamina
entire;Jamaica,CentralAmerica, N South America, eastwardto Suriname,Trinidad,
Tobago, N fringe of AmazonianBrazil (Para,Roraima)..................................................... 41. C. peltata
9. Stipules usually 10-20 cm long and spathesmostly 8-19 cm long, if shorter,then the
segments of the lamina often lobate to sinuate.
10. Spathes often ,0 cm long; segments of the lamina often lobate to sinuate;centralto E
Brazil to Argentina,Paraguay,and S fringe of the Amazon basin .......... .......... 38. C. pachystachya
10. Spathes usually 10-19 cm long; segments of the laminaentire.
11. Spathes (usually) densely hairy inside; and/orleafy twigs sparselyhispidulousto
SYSTEMATICTREATMENT
glabrous;E Cordillera,Colombia to Venezuela,mostly submontaneto montane
..........................................................................................................................
11. Spathes sparselyhairy inside; leafy twigs densely hispidulous;Pacific Coastal
lowlands of Ecuadorand Colombia and NW Amazon basin (Colombia,Ecuador,
and Peru).......................................................
1. Cecropia albicans Trecul, Ann. Sci. Nat. Bot.,
Ser. 3, 8: 82. 1847 (Aug). Type. Peru. Without
locality, without date (Y), Ruiz & Pavo'ns.n. (ho-
lotype: FI-W, photograph seen; isotypes: F, G,
LE).
Tree, to 20 m tall; branches(very) short and de-
partingat angles of ca. 900 at the apex of the tree or
longer and then departingin acute angles at the lower
part of the tree. Leafy twigs 3-10 cm thick, blackish
or reddishto green and then with a bluish waxy layer,
densely (sub)hirsuteor (sub)glabrous.Laminacoria-
ceous, ca. 25 X 25 cm to 95 x 95 cm; the segments
11-13, the upper ones in deeply incised laminas ob-
long (to subovate), the incisions down to 4/10 or to
8/10; apices acute to subacuminate;upper surface
smooth, subglabrousor with ? dense arachnoidin-
dumentum;lower surface rathersparsely to densely
tomentoseon the veins, arachnoidindumentumin the
areoles and on the smaller veins or also on the main
veins; lateralveins in the free partof the midsegment
10-16 pairs, submarginallyloop-connected, in the
lower partoften not distinctly so, often ? branched;
petiole ca. 30-75 cm long, puberulousandwith sparse
to rather dense arachnoid indumentum or subgla-
brous;trichiliaabsentor presentand fused, the brown
indumentum intermixed with short to rather long
white to pale brown (unicellular)hairs; stipules ca.
(lO-)25-45 cm long, red to pinkish, sometimes with
a bluish waxy layer, with only moniliform brown
pluricellularhairs outside, glabrous inside; terminal
buds inflated.Staminateinflorescencesin pairs,erect,
usually subtendedby pinkish or greenish, caducous
bractsto 4 cm long, attachedat or ca. 1 cm above the
base of the peduncle; peduncle 3-7 cm long,
(sub)glabrous;spathe ca. 4-10 cm long, yellowish to
pinkish, with only moniliform brown pluricellular
hairs outside, glabrousinside; spikes (7-)15-24, 3.5-
9 x 0.3-0.7 cm, sessile or with stipes 0.2-0.3 cm long
and sparsely hairy; rachis hairy. Staminateflowers:
perianthtubular, 1-1.5 mm long, sparsely and min-
utely puberulouson the lower part of the apex, the
apex plane, ? clearly 2-lobed; filamentsflat;anthers
ca. 0.8 mm long, appendiculate,detachedat anthesis,
remainingattachedto the filamentby 2 filiformcon-
nectionsbetween the connectiveandthe uppermargin
of the filament.Pistillate inflorescencesin pairs,erect,
usually subtended by pinkish, caducous bracts to 4
47
48. C sararensis
26. C. litoralis
cm long, attachedat or ca. 1 cm above the base of the
peduncle;peduncle 1.5-2.5 (or to 14) cm long, sub-
glabrous; spathe 5-11 cm long, the color and indu-
mentum as in the staminateinflorescence;spikes 4-
7(-9), 4-10 X 1-1.2 cm, to 15 X 2 cm in fruit,sessile;
rachis glabrous.Pistillateflowers: perianthca. 2 mm
long, with arachnoidindumentumbelow the apex out-
side, also in the lower partof the style channelinside,
the apex ? convex, punctateto submuriculate;style
rather long, straight, unilaterallyhairy; stigma co-
mose. Fruit oblongoid, ca. 2.5 mm long, ? tubercu-
late.
Distribution (Fig. 8.2). Peru (Pasco to San Mar-
tin), in cloud forest, at 2200-2500 m.
Specimens examined. PERU. HuANUCO:Rd. Huan-
uco-Tingo Maria,Carpish,ca. 2500 m, 23 Nov 1997 (6),
Berg et al. 1730 (BG, COL, MOL); rd. Hudnuco-Tingo
Maria,below Carpish,ca. 2400 m, 23 Nov 1997 (Y fl-fr),
Berg et al. 1732 (BG, COL, MOL). PASCO:Prov. Oxa-
pampa,rd. Oxapampa-Lluapi,km 3-4, ca. 2250 m, 29 Nov
1997 (Y fl-fr),Berg et al. 1757 (BG, COL,MOL),(6), Berg
et al. 1758 (BG, COL, MOL); Prov. Oxapampa,rd. Oxa-
pampa-Cerrode Pasco, 20-30 km W of Oxapampa,2000-
2500 m, 3 Feb 1983 (Y fr), Gentry et al. 39956 (BG, MO,
MOL,UMS). SAN MARTiN: Prov.Rioja,rd.Rioja-Poma-
cocha, ca. km 379, ca. 2200 m, 8 Dec 1997 (d), Berg et al.
1800 (BG, COL,MOL);Prov.MariscalCaceres,Abiseo Na-
tional Park,Las Palmas, E of Gran Pajatenruins, 2250 m,
17 Aug 1986 (? fl), Young4071 (F, MOL). WITHOUT LO-
CALITY: (6), Ruiz& Pav6n s.n. (BM,K).
The materialcollected in Pasco, matchingthe type
material,has been made from trees with the typical
Cecropia tree shape, although the primarybranches
are short and do not branch,the stems are broadened
towardthe apex, the leafy twigs aredensely hairy(and
blackish), the lamina of adult trees is incised <5/10,
and trichilia are present. Berg et al. 1800 from San
Martin largely matches that from Pasco, but Young
4071 from San Martin matches the Huanuco material.
The materialcollected in Huanucotaken from trees
more or less fastigiatein shape, with lateralbranches
departing(at one to threelevels) in acute angles from
the lower partof the stem, the stem is not broadened
towardthe apex, the leafy twigs are glabrouswith a
bluish waxy layer, the lamina is incised >5/10, and
trichiliaare absent.In otherfeaturesthe two types are
largely similar, occur at similar elevations, and co-
occur with C. tacuna both in Pasco and Huanuco.
48 FLORA NEOTROPICA

tAX~~~~~~~~

FIG. 8. Distribution maps. 1. Cecropia latiloba, circles; C. tacuna, hexagons; C. virgusa, triangles. 2. C. albicans,
hexagons; C. heterochroma, triangles; C marginalis, circles. 3. C. litoralis, circles; C. longipes, hexagons; C. mutisiana,
triangles. 4. C. obtusa. 5. C. chlorostachya, hexagon; C. obtusifolia, circles; C. pittieri, triangle. 6. C. granvilleana, open
circles; C. hololeuca, hexagons; C membranacea, circles; C. saxatilis, triangles. 7. C. garciae, triangles (question mark for
uncertain locality); C. montana, circles.
SYSTEMATICTREATMENT
Young4071 also has glabrous leafy twigs and also
lacks trichilia. Juvenile specimens of the Pasco type
has laminaswith incisions to 9/10, the free partof the
midsegmenthas ca. 17-20 pairs of lateral veins, the
dense subhirsuteindumentumfoundin the leafy twigs
occurs also on the petioles and on the main veins of
the lamina beneath,and the stipules are pinkish. The
peduncleof the pistillateinflorescenceis usually very
short (to 2.5 cm), but in Young4071 the peduncles
are much longer (to 9 cm). The trees in Pasco are
usually inhabitedby ants, those in Huainuconot.
2. Cecropia andina Cuatrecasas,RevistaAcad. Col-
omb. Ci. Exact. 6(22/23): 286. 1945. Type. Co-
lombia. Putumayo:Nr. source of Rio Putumayo,
SibundoyValley,2200 m, 3 Jan 1941 (Y) Cuatre-
casas 11662 (holotype:COL; isotype: F). Fig. 9
Tree,to 20 m tall. Leafy twigs 2-4 cm thick,bluish
due to a waxy layer, glabrous;pith brown and copi-
ous. Lamina subcoriaceousto chartaceous,ca. 40 X
40 cm to 80 X 80 cm (to 130 X 130 cm), the segments
7-9, the free parts of upper segments elliptic to su-
bobovate, the incisions down to 4/10-7/10 or in the
lower part of the lamina <4/10; apices acuminateto
obtuse;uppersurface ? scabrous,sometimes ? bul-
late, hirtellous to puberulousto hispidulous subhir-
sute; lower surfacehirtellousto subtomentoseon the
veins or the primaryveins subglabrous,the arachnoid
indumentum in the areoles, often very short and
sparseor only nearthe margin,sometimesabsent;lat-
eral veins ca. 10-15 pairs, submarginally loop-
connected, most of them branched;petiole 30-85 cm
long, (sub)glabrous,often ? bluish due to a waxy
layer;trichiliaabsent,or if present,then separateand
usually poorly developed, the brownindumentumin-
termixedwith dense shortandratherlong white hairs;
stipules ca. 15-30 cm long, red to greenish,glabrous
outside, hairy inside. Staminate inflorescences soli-
tary or in pairs, the peduncle erect, the spikes pen-
dulous; peduncle 6-10 cm long, glabrous or with
sparse long white hairs, often bluish due to a waxy
layer; spathe 18-30 cm long, green to yellowish or to
pink, almost glabrousoutside, densely (sub)sericeous
inside; spikes 6-16, 9-27 X 0.3-0.6 cm, the stipes
0.6-1 cm long, glabrous;rachis with shorthairs.Sta-
minateflowers sessile or short-pedicellate;perianth
tubular,1.2-1.6 mm long, glabrous,the apex slightly
convex to almost plane, slightly 2-lobed, smooth;fil-
aments flat; anthers0.6-0.8 mm long, appendiculate,
detachedat anthesis,reattachedto the marginsof the
apertureby the appendages.Pistillate inforescences
solitaryor in pairs,pendulousor the peduncleinitially
49
erect; peduncle 6-14 cm long, glabrous,often bluish
due to a waxy layer; spathe ca. 15-22 cm long, the
color and indumentumas in the staminateinflores-
cences or glabrous inside; spikes (3 or) 4, 12-20 X
0.8-1 cm, to 35 X 1.2 cm in fruit, subsessile or with
stipes to 1.5 cm long and glabrous;rachis hairy or
glabrous.Pistillate flowers: perianthca. 2 mm long,
with arachnoidindumentumbelow the apex outside,
also in the style channel inside, the apex convex to
dome-shaped,smooth; style short, hairy. Fruit ellip-
soid to ovoid, ca. 1.2-1.5 mm long, smooth.
Distribution (see Fig. 10.3). FromColombia(Ca-
queta and Putumayo) to Peru (Amazonas and San
Martin),easternslopes of the Andes, in (cloud)forest,
at 1600-2300 m.
Representative specimens examined. COLOMBIA.
CAQUETA: Mun. Florencia, below Gabinete,km 47, 2000
m, 16 Oct 1993 (9 fl-fr), Franco et al. 4503 (BG). HUILA:
30 km NW of Palermo,ca. 2300 m, 10 Oct 1944 (9 fl-fr),
Little8788 (COL,F, U). PUTUMAYO:Rd. Mocoa-Sibundoy,
El Mirador,2000 m, 5 May 1994 (9 fl-fr), Franco et al.
5529 (BG).
ECUADOR. LOJA: Rd. Loja-Zamora, QuebradaNa-
vidad,30 Jan 1996 ( 9 fl), Merinoet al. 4779 (LOJA).NAPO:
Bermejo, valley of Rio Cosanga, 1800 m, 27 Dec 1939 (9
fl-fr), Asplund 10150 (S); Cosanga, ca. 20 km S of Baeza,
ca. 1850 m, 20 Sep 1977 (d), Berg et al. 439 (BG, COL,
QCA, U); rd. Baeza-Cotundo, 1990 m, 12 Aug 1975 (st),
Little et al. 150 (Q, QAME, QCNE); Cant6n Quijos, be-
tween Baeza & Cuyuja,2400 m, 13 Jan 1992 (9 fl), Palacios
et al. 9627 (BG, QCNE);nr.Baeza, ca. 2000 m, 23-24 May
1992 (9 fl), Valenciaet al. 2306 (BG, QCA). TUNGURA-
HUA: Rio Pastaza,Hda. Rio VerdeGrande,ca. 1500 m, 26
Jul 1939 (9 fl-fr), Asplund 7849 (F, S), 4 Feb 1956, (6),
Asplund 19205 (S); Hda. Michai, ca. 1500 m, 4 Feb 1956
(6), Asplund 19228 (S). ZAMORA-CHINCHIPE:Rd.Loja-
Zamora,nr. Sabanilla,ca. 1700 m, 3 Jan 1991 (d), Berg et
al. 1652 (BG, LOJA, QCA), (9 fl), Berg et al. 1653 (BG,
LOJA,QCA); 18 km E of Loja, 1800 m, 3 Sep 1975 (9 fl-
fr), Little et al. 213 (COL, LOJA, Q, QAME, QCNE); rd.
Loja-Zamora,ca. km 13 E of the pass, before jct. with old
rd., ca. 2030 m, 8 Mar 1989 (st), 0llgaard et al. 90836
(AAU, LOJA,QCA).
PERU. AMAZONAS: Prov. Bongara, Dtto. Sipabamba,
Shilla, 1900 m, 5 May 1981 (9 fr), Younget al. 357 (BG,
MO, NY. TEX). SAN MARTiN: Prov. Rioja, rd. Rioja-Po-
macocha, nr. km 392, 1600-1700 m, 9 Dec 1997 (9 fl-fr),
Berg et al. 1796 (BG, COL, MOL), (d), Berg et al. 1797
(BG, COL, MOL); rd. Pedro Ruiz-Moyobamba, km 390,
Venceremos,1770-2150 m, 5-7 Aug 1983 (st), D. N. Smith
et al. 4657 (MO).
This species shows close affinities to Cecropia
pastasana, as discussed under the latter. Cecropia an-
dina can be distinguished from C. pastasana by the
absence of trichilia (or by very poorly developed
50 FLORA NEOTROPICA

f~~~~~~~~~~~~~~~~~~~~~~~

4 ~ ~~ ~ ~ ~\~ ~ ~~~-

FIG. 9. Cecropia andina. 1. Lamina, reduced. 2. Apex of lamina and venation (Berg et al. 1652). 3. Stipules and
young leaf (Berg et al. 1653). 4. Lower surfaceof lamina.5. Staminateinflorescencewith spathe.6. Staminateinflorescence
at anthesis. 7. Staminateflower and stamen (Berg et al. 1652). 8. Filament after detachementof anther(Berg et al. 1797).
9. Pistillate inflorescence with spathe. 10. Pistillate inflorescence at anthesis (Berg et al. 1653). 11. Pistillate flower and
pistil. 12. Fruit(Franco et al. 4503).
SYSTEMATICTREATMENT 51

X~~~~~~~~~~~~~~

4.0-~~~~~~~~~~~~~~~~~~~~

FI(G.10. Distributionmaps. 1. Cecropiaangustifolia,circles (open circles for materialof not entirelycertainidentity);

C. kavanxensis,triangles;C. purpurascens,hexagons. 2. C. annulata, open circle; C.ficifolia, circles; C. telealba, triangles.
3. C. ancdina,triangles;C. distachsva,circles. 4. C. engleriana, circles; C. maxima,triangles.5. C. hispidissima.6. C. herthae,
triangles;C. idroboi, hexagons; C. insignis, circles.
52 FLORANEOTROPICA

ones) and usually also by the smooth leafy twigs with long, glabrous; rachis glabrous. Staminateflowers:
a bluish waxy layer.In the southernpartof the species perianthtubular,1-2 mm long, sparselyminutelypu-
range (Zamora-Chinchipeand Peru), in adult speci- berulous below the apex, the apex almost plane; an-
mens the incisions of the laminarun down to at most thers0.6-0.8 mm long, appendiculate,detachedat an-
5/10 of the distance between margin and umbilicus thesis, reattachedto the marginsof the apertureby the
(or less in the lower partof the lamina)and the lower appendages (?). Pistillate inflorescences in pairs or
surface of the lamina is conspicuously hairy. In the solitary, initially erect, in fruit pendulous;peduncle
(possibly disjunct)northernpartof the species range, 3-14(-18) cm long, puberulousto hispidulous;spathe
the lamina is more deeply incised and less conspicu- 6-13 cm long, the color unknown,puberulousto hir-
ously hairy beneath. In the Sibundoy valley (Putu- tellous, or also with ? dense arachnoidindumentum
mayo, Colombia)and nearLoja (Ecuador),C. andina outside, glabrousinside; spikes (2-)4(-6), (3-)7-9 X
extends somewhat onto the western side of the wa- 0.4-0.6 cm, to 16 X 1.2 cm in fruit,subsessile or with
tershed. stipes to 0.6 cm long and subglabrous;rachis hairy.
Vernacular name. Peru:chanchan(Amazonas). Pistillate flowers: perianth ca. 1.5 mm long, with
arachnoidindumentumbelow the apex, also below
the style channel inside, the apex convex, glabrous
and smoothor muriculate;style short;stigmacomose.
3. Cecropia angulata I. W. Bailey, Bot. Gaz. (Craw- Fruit oblongoid, ca. 2.5 mm long, tuberculate.
fordsville) 74: 378. t. 1-7. 1922; Veldsquez,Acta
Bot. Venez. 6: 39, t. 4. 1971. Type. Guyana.Kar- Distribution (see Fig. 11.2). Guyana and north-
tabo region, Barakara,Jul-Aug 1920 (d), Bailey eastern Venezuela,in forest and in secondarygrowth,
9 (lectotype, here designated:GH). at low elevations.

Tree,to 20 m tall. Leafy twigs 3-6 cm thick,green, Representativespecimensexamined.VENEZUELA.

BOLivAR: Dtto. Sifontes, Concesi6n MineraCristina Cua-
puberulousto hispidulous,some of the hairsuncinate.
tro, 11 Aug 1985 (Y fl-fr), Aymardet al. 4146 (MO, US);
Lamina (sub)coriaceous,ca. 55 X 55 cm to 100 X Dtto. Sifontes, 3-12 km SE of CampamentoCVG Las Flo-
100 cm (to 120 X 120 cm), the segments (8-)9-1 1, res, 16-17 Sep 1989 (Y fl-fr), Colella et al. 1444 (BG,
the free partsof uppersegmentsobovate,the incisions VEN); Dtto. Roscio, San Martinde Turumban, Aug 1979
down to ca. 7/10-9/10; apices obtuse to rounded;up- (Y fl-fr), Delascio et al. 8010 (VEN); Mun. Sucre, 2 km S
per surface usually smooth, sometimes scabridulous, of SantaMaria de Erebato,Jan 1989 (Y fl-fr), Fernandezet
usually subglabrousor sometimes scabrous,initially al. 5007 (BG, MO); Mun. Cedenio, Serraniade Maigualida,
often also with sparse arachnoidindumentum;lower 20 km E of San Jose de Kayama, Apr 1989 (Y fl), Ferndndez
surface puberulous on the veins, and with sparse 5505 (MO,NY);rd.El Dorado-Santa Elenade Uairen,be-
longer uncinatehairs, with arachnoidindumentumin tweenkm 88 & PiedraLa Virgen,30 Jan 1981 (Y fl-fr),
the areoles and on the smaller veins, or sometimes Marcano Berti et al. 144-981 (U, VEN); Rio Caura,Cafno
La Ceiba (Cafio Cumaca),9-26 May 1988 (d), Stergios et
extending to the main veins; lateralveins in the free
al. 12888 (BG, NY, VEN); Reserva Forestal del Imataca,
part of the midsegment 15-20 pairs, submarginally
Sep 1967 (? fl-fr), Velasquez252 (US, VEN), (Y fl), Velds-
loop-connected, unbranched or the lower ones quez253 (VEN). DELTAAMACURO: Depto. Antonio Diaz,
branched;petiole 50-110 cm long, sparsely puberu- nr. Curiapo,Feb 1987 (Y fl-fr), Ferndndez3964 (BG, MO,
lous to hirtellous or toward the apex often densely NY, VEN); Cafno Araguao, Aug 1954 (Y fr + d), Gines
hirtellous,or also with ? dense ( ? persisting)arach- 5156 (US); Rio Winikina, 16 Jan 1983 (st), Wilbert68325
noid indumentum;trichilia fused, the brown indu- (MYF).
mentum intermixed with short white hairs; stipules GUYANA. KartaboRegion, (partly in) Barakara,Jul-
12-20 cm long, color unknown, outside ? densely Aug 1920 (st orjuv), Bailey 3 & 17 (GH); NorthwestDistr.,
puberulousto hirtellousor sometimesalso with arach- Port Kaituma,Aug 1967 (st), Davis 239 & 240 (K, NY);
noid indumentum,inside (rather)sparselyhairy.Sta- Northwest Distr., Waini R., 3-18 Apr 1923,De La Cruz
minate inflorescences in pairs, the peduncle curved 3752 (F, GH, MO, NY, US); MazaruniStation, 19 Jun 1942
(9 fl-fr), Fanshawe 732 (FD 3468) (K, NY); EssequiboR.,
downwardand the spikes pendulous at anthesis;pe-
Bartica, 12 Aug 1950 (d), Fanshawe 2990 (FD 6320) (K,
duncle (6-)10-15 cm long, puberulousto hispidulous
NY, S, U); Demerara-Mahaica Region, 1-2 km S of Loo Cr.,
or also with arachnoidindumentum;spathe 12-16 cm on Linden Hwy., 2 Mar 1989 (9 fl-fr), Gillespie et al. 803
long, color unknown, outside sparsely to rather (BG); Kangaruma,23-27 Jun 1921 (9 fl-fr), Gleason 198
densely puberulousto hirtellous or also with arach- (GH, NY, US); Kamarang,21 Aug 1977 (d), Maas et al.
noid indumentum,inside glabrous;spikes ca. 8-20, 2589 (NY, U); EssequiboR., Kurupukari,3 Oct 1937 (9 fl),
4-14 x 0.3-0.6 cm long, ? angular;stipes 0.8-2 cm A. C. Smith2164 (A, G, K, MO, NY, P,S, U, US); nr.Mabura
SYSTEMATICTREATMENT 53

4 ..~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~.....

A~~~~~~~~~

0~~~
C. 0tuabn,crls
hexagons;~'
FI.11 piat,tiages* cidpyla crls.2
itrbtinmps .Cerpi lzivi,hxgos

anulta ranle;C le, icls.3.Cbulaa ice utsca

exgos;Csiva, 0rage;Cguoin,oe

heagn; ucbabaa crce.0

Hill sawmill, 31 Oct 1982 ( fl-fr), Stofferset al. 159 (MO, C. angulata C. obtusa
U); Upper Mazarunibasin, Kukui R., Suruagupuh,12 Sep Stipules 12-20cm 7-12(-15) cm
1960 (9 fl-fr), Tillettet al. 45398 (BG, K, NY); Kamarang Segments (8-)9-11 6-8(-9)
R., above Utschi mouth, 25 Oct 1960 (9 fl-fr), Tillettet al. Incisions 7/10-9/10 5/10-7/10
45804 (BG, NY).
(-8/10)
This species appearsto be closely related to Ce- Pairs lat. veins 15-19 9-15
cropia obtusa. It matches C. obtusa in many features, Lamina (Sub)coriaceous Chartaceousto
but can be told apartby the differencespresentedbe- subcoriaceous
low: d peduncle (6-)10-15 cm 4-8.5 cm
54
Withthe listed differencesthe two taxa can be dis-
tinguished,althougha few specimens (such as Maas
et al. 7501, includedin C. obtusa) tend to be more or
less intermediate.The flowers of C. angulata resem-
ble those of both C. obtusa and C. ficifolia. In the
shape, the texture, and the smooth upper surface of
the lamina, C. angulata also resembles C. distachya,
from which it can be usuallydistinguishedby the sub-
marginallyloop-connectedlateralveins and the more
or less dense arachnoidindumentumon the petiole,
featuresonly occasionally found in C. distachya. Al-
though C. angulata shows strongmorphologicalsim-
ilarities to both C. distachya and C. obtusa, it should
be maintainedas a distincttaxon on the species level,
at least for the time being, awaitingcomparativefield
studies.
Vernacular names. Venezuela:kama'-in-yek(Bo-
livar), waru (WinikinaWarao,Delta Amacuro).Guy-
ana: congo pump, congopom.
4. Cecropia angustifolia Trecul,Ann. Sci. Nat. Bot.,
ser. 3, 8: 83. 1847 (Aug). Type. Peru.Withoutlo-
cality, (Y), Ruiz & Pav6n s.n. (holotype: FI-W,
photographex FI-W in G; isotypes: B, G, OXF).
Figs. 12, 13, 14
CecropiaacutifoliaTrecul,Ann.Sci. Nat.Bot., ser.3,
8: 81. 1847(Aug).Type.Peru.Withoutlocality,(d
+ Y),Ruiz& Pav6ns.n. (FI-W,photographs seen,
mixedcollectionconsistingof leavesof C. angusti-
folia andinflorescencesof C. strigosa,the leaf on
the sheetwiththe staminateinflorescence heredes-
ignatedas thelectotype).
Cecropia digitata Klotzsch, Linnaea 20: 534. 1847
(Oct).Type.Peru.Withoutlocality,(6), Ruiz& Pa-
vons.n. (holotype:B, destroyed,
photograph ex B in
F, G, MO;isotypes:B, G, OXF).
CecropiatubulosaKlotzsch,Linnaea20: 534. 1847
(Oct).Type.Peru.Huanuco:Chinchoaor Cuchero,
Ruiz& Pavons.n. (holotype:B-n.v.,destroyed; is-
otypes:probablytheholotypeandisotypesof C.an-
gustifolia).
Cecropiapolyphlebia Donnell Smith, Bot. Gaz. (Craw-
Bot.40:
fordsville)27:442. 1899;Burger,Fieldiana,
127, t. 23. 1977.Type.CostaRica. San Jose:La
Palma,1460m, Sep 1898(9), Tonduz12642 = in
herb.Donnell-Smith 7411 (holotype:US; isotypes:
F, G, GH,K).
CecropiasylvicolaStandley& Steyermark, Publ.Field
Mus.Nat.Hist.,Bot.Ser.23: 153.1944.Type.Gua-
temala.AltaVerapaz: Rd.Tacticto thedivideon rd.
90762(holotype:
to Tamahu, Apr1941(d), Standley
F).
CecropiacoriaceaCuatrecasas,RevistaAcad.Colomb.
Ci. Exact.6(21): 66. 1944;Not. Fl. Colomb.VI.
FLORA NEOTROPICA
(Trab.Com. Bot. etc. Colomb.) 41. 1944. Type. Co-
lombia. Huila: Between Gabinete & Andalucia,
2200-2300 m, 25 Mar 1940 (Y), Cuatrecasas8697
(holotype:COL; isotype: F).
Cecropiastrigilosa Cuatrecasas,RevistaAcad. Colomb.
Ci. Exact. 6(22/23): 282. 1945. Type. Colombia.
Valle:Rio Calima,El Cairo,between Darien & Me-
diacanoa, 1700 m, 6-7 Jan 1943 (5), Cuatrecasas
13872 (COL), (9) Cuatrecasas 13939 (not traced),
Cuatrecasas 13872 here designated as lectotype
(COL).
Cecropia caucana Cuatrecasas,Revista Acad. Colomb.
Ci. Exact. 6(22/23): 289. 1945. Type. Colombia.
Cauca:Timbio,Perez Arbeldez& Cuatrecasas6117
(holotype:COL; isotype: US).
Cecropia danielis Cuatrecasas,Revista Acad. Colomb.
Ci. Exact. 6(22/23): 290. 1945. Type.Colombia.An-
tioquia:San Antonio del Prado, 1870 m, 7 Aug 1943
(5), Hno. Daniel 2843 (holotype:COL;isotypes: F,
GH, MEDEL).
Cecropiapalmatisecta Cuatrecasas,Revista Acad. Col-
omb. Ci. Exact.6(22/23): 297. 1945;Velasquez,Acta
Bot. Venez. 6: 49, t. 10. 1971. Syntypes. Colombia.
Caqueta:Sucre, La Portada, 1200-1350 m, 5 Apr
1940 (5), Cuatrecasas 9157 (COL), (9), Cuatre-
casas 9158 (not traced),Cuatrecasas9157, here des-
ignated as lectotype (COL).
CecropiahachensisCuatrecasas,RevistaAcad. Colomb.
Ci. Excact. 6(22/23): 298. 1945; Berg & FrancoRos-
selli, Fl. Ecuador48: 23. 1993. Type. Colombia.Ca-
quetd:Rio Hacha, Caj6n de Pulido, 1700-1750 m,
26 Mar 1940 (9), Cuatrecasas 8782 (holotype:
COL; isotype: F).
CecropiamoniquiranaCuatrecasas,Revista Acad. Col-
omb. Ci. Exact. 6(22/23): 298. 1945. Type. Colom-
bia. Boyaca: Nr. Moniquira, 1580 m, 25 Feb 1940
(9), Perez Arbelaez & Cuatrecasas8135 (holotype:
COL; isotypes: F, US).
Cecropia philipsonii Cuatrecasas,Revista Acad. Col-
omb. Ci. Exact. 9(36/37): 382. 1956. Type. Colom-
bia. Meta: Sierra de la Macarena,North Ridge, 29
Dec 1949 (5), Philipson & Idrobo 1983 (holotype:
US; isotypes: BM, COL).
Cecropiavillosa C. C. Berg & P. Franco,Fl. Ecuador48:
53. 1993. Type. Ecuador. Zamora-Chinchipe:Rd.
Loja-Zamora, nr. Sabanilla, 1600-1700 m, 3 Jan
1991 (9 fl-fr), Berg et al. 1651 (holotype: QCA;
isotypes: BG, GB, LOJA).
Tree, to 20(-25) m tall. Leafy twigs (1-)2-5 cm
thick, green to brownish to often (turning) blackish,
hispidulous to hirtellous to strigose to subvelutinous
or to villous, usually also with dense brown to black-
ish pluricellular hairs, sometimes also with arachnoid
indumentum; internodes with sparse (whitish) pith or
sometimes partly to entirely filled with brown pith.
Lamina chartaceous to coriaceous, ca. (20 X 20 cm
to) 45 X 45 cm to 75 X 75 cm (to 90 X 90 cm), the
SYSTEMATICTREATMENT 55

FIG. 12. Cecropia angustifolia. 1. Lamina, reduced (Berg 1279). 2. Lamina (Berg 1248). 3. Apex of lamina and
venation. 4. Stipules and base of petiole with trichilium (Berg et al. 44J). 5. Stipules and young leaf (Berg 1281). 6. Pair
of staminateinflorescenceswith spathesand base of petiole with trichilum(Hurtadoet al. 2581). 7. Staminateinflorescence
at anthesis and base of petiole with trichilium (Gentry et al. 63245). 8. Staminateflower and stamen (Berg et al. 1633). 9.
Pistillate inflorescence with spathe (Dodson et a!. 10541). 10. Pistillate inflorescence at anthesis and base of petiole with
trichilium(Berg et a!. 1204). 11. Pistillate flower and pistil (Berg et a!. 1632). 12. Fruit(Franco et a!. 4475).
56 FLORA NEOTROPICA

/4~~~~~~~~~~~~~~~~~~~~~~~~~~~4

I /~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~

FIG. 13. Cecropiaangustifolia. 1. Lamina,reduced.2. Apex of laminaand venation.3. Stipules,young leaf, staminate
inflorescences with spathes, and base of petiole with trichilium. 4. Pair of staminate inflorescences with spathes and just at
anthesis. 5. Staminate flower. 6. Stamen and filament (Berg et al. 1753). 7. Pair of pistillate inflorescences with spathes and
base of petiole with trichilum. 8. Pairs of pistillate inflorescences at anthesis and base of petiole with trichilium. 9. Pistillate
flower and style. 10. Fruit (Berg et al. 1750). From Caldasia 24: 232. 2002.
SYSTEMATICTREATMENT 57

FIG. 14. Cecropia angustifolia. 1. Lamina, reduced. 2. Apex of lamina and venation. 3. Stipules. 4. Staminate
inflorescencewith spatheand young leaf. 5. Staminateflowerand stamen(Berg et al. 1650). 6. Pairof pistillateinflorescences
at anthesis. 7. Pistillate flower. 8. Fruit(Berg et al. 1651). (From Fl. Ecuador48: 54. 1993, modified.)
58
segments (8-)10-14, in young leaves often reflexed,
the free partsof uppersegments (ob)lanceolateto ob-
long to subobovate, the incisions down to (5/10-)
6/10-9/10 (or to near the petiole); apices obtuse to
rounded or subacute to short-acuminate;upper sur-
face scabrous or smooth, densely to sparsely hispi-
dulous or puberulous or partly to hirtellous or to
(sub)strigillose, also with ? dense brown pluricel-
lularhairs and sometimes sparsearachnoidindumen-
tum, the "umbilicus"sometimes villous; lower sur-
face sparsely to densely (minutely)puberulous(with
straightto uncinatehairs) to hirtellousto subtomen-
tose or to subhirsuteon the (main) veins, with arach-
noid indumentumconfined to the areoles, extending
to the smaller veins, or (almost) absent;lateralveins
in the free part of the midsegment (12-)15-35(-43)
pairs, submarginallyor marginally loop-connected,
branched or unbranched;petiole 20-70 cm long,
sparsely to densely puberulousto hirtellous to sub-
hirsuteor to villous, or sometimes also with sparseto
dense arachnoid indumentum; trichilia fused, the
brown indumentumintermixedwith short (or some-
times rather long) white to brownish (unicellular)
hairs; stipules 6-25(-30) cm long, caducous or sub-
persistent, green to yellowish, reddish to purplish,
brownish or black, sparsely to densely white to
brownishpuberulousto hirtellousor partlyto largely
subhirsuteto sericeous to subvelutinousor to villous,
also with ? dense brown pluricellularhairs outside,
sparselyto densely white- to pale brown-sericeous(or
glabrous) inside. Staminate inflorescences in pairs,
erect, often subtendedby subpersistentbracts,to 5 cm
long; peduncle 1-8(-12) cm long, puberulousto sub-
hispid to hirtellous to subhirsuteor to subvelutinous
and often (initially) with dense brown pluricellular
hairs and often also sparse arachnoidindumentum,
sometimes subglabrous;spathe4-14 cm long, green-
ish to yellowish to brownish or reddish to purplish,
densely white- to brownish-puberulousto -hirtellous
or partly to -subhirsuteto -strigose or to villous and
with dense brown pluricellular hairs or also with
sparseto ratherdense arachnoidindumentumoutside,
sparselyto densely (sub)sericeousor glabrousinside;
spikes 10-25(-50), (l.5-)3-11(-13) X 0.3-0.8 cm,
sessile or with stipes to 0.7(-1.2) cm long,
(sub)glabrousto puberulousto hirtellous;rachishairy
(or glabrous).Staminateflowers:perianthtubular,1-
2 mm long, with short and stiff hairs on the ribs,
arachnoidindumentumbelow the apex, or glabrous,
the apex plane to convex, smooth or muriculate,the
aperturecircular or slit-shaped; filaments flat or ?
swollen; anthers0.5-0.9 mm long, appendiculateor
not, detached and reattachedto the margins of the
FLORANEOTROPICA
apertureby the appendagesor remainingattachedto
the filament at anthesis. Pistillate inflorescences in
pairs, erect, often subtendedby subpersistentbracts,
to 3 cm long; peduncle 1-8(-12) cm long, the indu-
mentum as in the staminateinflorescence;spathe 3-
6(-8) cm long, color and indumentumas in the sta-
minateinflorescence;spikes 3-8(-12), 1-7 X 0.4-0.8
cm, to 17 X 0.8-2 cm in fruit,often broadenedtoward
the base, sessile; rachis hairy (or glabrous).Pistillate
flowers: perianth 1.5-2.5 mm long, with arachnoid
indumentumbelow the apex outside, also in the lower
partof and/orbelow the style channelinside, the apex
convex to almost plane, muriculateto minutely his-
pidulous or glabrous, the aperturecircular or slit-
shaped;style rathershort;stigmacomose. Fruitellip-
soid to oblongoid to ovoid 1.5-2.5 mm long,
tuberculateor smooth.
Distribution (see Fig. 10.1). From Guatemala(or
probably Mexico: Puebla) to the coastal mountain
range of Venezuelaand throughthe Andes (and some
adjacent low mountain ranges) to Bolivia, in wet
montaneor wet to relatively dry submontaneforest,
at ca. (400-)800-2400 m.
Representative specimens examined. MEXICO.
PUEBLA: Nr. Cuerrero, 15 mi E of Tezuitlan,16 Jul 1947
(6 + 9 fl-fr), Converses.n. (US); rd. Tlapacoyan(Veracruz)
to SanJoseAcateno,1000m, 22 Oct 1968(6), Pennington
et al. 9273 (A, K, NY).
GUATEMALA.ALTA VERAPAZ:E of Tactic,rd.to Ta-
mahu, 1500-1550 m, 9 Apr 1939 (st), Standley 71248 (F);
rd. betweenTactic& the divideon rd. to Tamahu,1500-
1600m, 1-7 Apr1941(6), Standley90576 (F).HUEHUE-
TENAGO: Nr. Maxbal,17 mi N of Barillas,Sierrade los
Cuchumatanes, 1500m, 15-16 Jul 1942(juv),Steyermark
48888(US).
HONDURAS. CORTES:ParqueNacionalCusuco,nr.
Centrode Visitantes,18 kmW of SanPedroSula,1620m,
21 Mar 1993 (6), Dario 355 (BG, EAP).OLANCHO: La
Muralla,ca. 10kmN of LaUni6n,ca. 1350m, 26 Oct1996
(6), Berg 1806 (EAP).
NICARAGUA. JINOTEGA:Macizosde PeniasBlancas,
nr.FincaEl Cielo,1200-1400m, 13-18 Jan1979(3 fl-fr),
Stevens11688(BG,MO);rd.Matagalpa-Jinotega,ca.4 km
NW of Aranjuez,ca. 1400m, 8 Dec 1983(9 fl), Stevens
22520 (MO,NY,TEX).MATAGALPA: BetweenDisparate
de Potter& Arajuez,1300m, 12 Jan1963(d), L. 0. Wil-
liams et al. 23663 (F, G).
COSTA RICA. ALAJUELA: Rio PetiasBlancas,1540-
1575 m, 18 Jun 1985 (9 fr), Haber et al. 1658 (MO); 2 km
N of La Balsa de San Ram6n, 800 m, 13 Feb 1977 (d + 9
fl-fr), Lent 4101 (AAU,MO,NY). CARTAGO: 10 km S of
Tapantf,aboveRio Grandede Orosi,1600m, 1 Sep 1971
(9 fl-fr), Burgeret al. 8428 (BM, EAP, G, MO, NY, PMA);
CerroDean,3 kmE of Cachi,1400m, 23 Apr1969(9 fl),
Lent1614(S); 0.5 km S of PetiasBlancasde Cachi,1400
SYSTEMATICTREATMENT
m, 5 Sep 1971 (Y fl-fr), Lent 2120 (BM, EAP, G); S limit
of Monumento Nacional Guayabo, sector Las Ventanas,
1100 m, 7 Jul 1992 (5), Rivera 1934 (K); 15 km NW of
Turrialba,ca. 1000 m, 13 Aug 1967 (Y fr), Taylor4393
(NY). HEREDIA: 3 km W of VaraBlanca, 1750 m, 8 Aug
1971 (? fl-fr), Lent 2052 (GH, U). PUNTARENAS: Cerro
Pando, above Rio Cot6n & Rio Negro, 1000-1800 m, 19-
21 Feb 1982 (st), Barringer et al. 1607 (LL); Monteverde,
1500 m, 15 Apr 1981 (5), Haber 455 (MO). SAN JOSE:
Cord. de Talamanca,between Eusa Mato & El Empalme,
2200-2400 m, 3 Mar 1966 (9 fl), Blum et al. 2225 (MO);
Cerro El Espino (Alto Mata de Cania-Azahar),Patarra,
1600-1800 m, 13 Nov 1983 (5), Chacon et al. 1563 (BG,
MO); Las Tablas,Rfo Cotoncito, 10 Dec 1983 (5), Chacon
et al. 1816 (BG, COL, MO, NY); Rfo ParaBlanco, 1500 m,
4 Feb 1973 (5), Lent 3154 (COL, U).
PANAMA. CHIRIQUI: Cerro Horqueta,ca. 2300 m, 24
Jul 1966 (2 fl-fr), Blum et al. 2600 (MO); CerroColorado,
above 1000 m, 9 Aug 1984 ( 9 fl-fr), Schmalzelet al. 2026
(BG).
COLOMBIA. ANTIOQUIA: Mun. La Ceja, 3 km N of
La Ceja, 2400 m, 21 Jan 1994 (5), Callejas et al. 11111
(BG, COL, HUA); Mun. Medellin, SantaElena, 1700 m, 24
Mar 1994 (5), Franco et al. 4577 (BG, HUA), (9 fl-fr),
Franco et al. 4578 (BG, HUA), (5), Franco et al. 4580 (BG,
HUA), (5), Franco et al. 4582 (BG, HUA); Mun. Cocormn,
Vrda.Viao, 1950 m, 26 Mar 1994 (5), Franco et al. 4591
(BG, HUA); Mun. Frontino,Cgto. Nutibara,rd. Nutibara-
La Blanquita, 1700 m, 4 Mar 1995 (9 fl-fr), Franco et al.
5547 (BG, COL, HUA, LP, MO, US); Mun. Frontino,rd.
Nutibara-Frontino,km 14, 1750 m, 5 Mar 1995 (d), Franco
et al. 5564 (COL, HUA, LP, MO, US); Mun. Anorf, Vrda.
El Roble, 1400 m, 8 Mar 1995 (5), Franco et al. 5568 (BG,
COL, HUA, LP,MO, US); Mun. Anorf,rd. Anorf-El Roble,
1600 m, 8 Mar 1995 (9 fl-fr),Francoet al. 5570 (BG, COL,
HA, LP, MO, NY, QCA). BOYACA: Mun. Pajarito,below
Corinto, 1200 m, 29 Jan 1995 (9 fl-fr), Franco et al. 4641
(BG, COL, HUA, MO, US); between Moniquira& Gach-
antivd, 1200-1600 m, 14 May 1996 (st), Rangel et al. 13234
(COL). CALDAS: Manizales, 2153 m, Jul 1938 (5), Duque-
Jaramillo4048 (COL, NY). CAQUETA:Mun. Florencia,be-
low Gabinete,km 17, 2200 m, 15 Oct 1993 (5), Franco et
al. 4493 (BG), km 45, 2150 m, 15 Oct 1993 (9 fl-fr),Franco
et al. 4495 (BG, COL), 15 Oct 1993 (5), Franco et al. 4496
(BG); Mun. Florencia,rd. Florencia-Gabinete,km 49, 1900
m, 16 Oct 1993 (5), Franco et al. 4502 (COL), ca. 1050 m,
18 Oct 1993 (9 fl-fr), Franco et al. 4513 (BG, COL), ca.
1750 m, 18 Oct 1993 (9 fl-fr), Franco et al. 4515 (BG,
COL);trailLas Guacamayas-Ramos,km 59, ca. 1600 m, 29
Apr 1944 (9 fl-fr),Little 7763 (COL, US). CASANARE:Nr.
Sacama, 1100 m, 27 Jan 1995 (5), Franco et al. 4637
(COL); Sacama, Vrda. Guavarin,1750 m, 27 Jan 1995 (9
fl-fr), Franco et al. 4638 (BG, COL, HUA, MO, US).
CAUCA: Between Popayan & Piendam6, 1800 m, 31 Dec
1942 (9 fl), Cuatrecasas13855 (F, US, VALLE);valley of
Rio Palac6, rd. to Totor6, Quebradade Palac6, 1820-1960
m, 15 Oct 1961 (5), Cuatrecasaset al. 26487 (COL,P,US),
(9 fl-fr), Cuatrecasas et al. 26491 (COL, P, US); rd. La
Plata-Purace, 12 km from Santa Cecilia, Reserva Meren-
59
berg, 15 Jul 1977 (6), Idrobo8753 (COL);Rio San Joaquin,
1400-1500 m, 29-30 Jun 1922 (6), Killip 7874 (GH, K,
NY, US); Popayan,21 Jul 1960 (9 fl), Schultes22508 (COL,
GH); El Tambo, 1700 m, 8 Oct 1939 (9 fl), Sneidern2218
(A, LL, MICH, NY, S, TEX, US). CESAR: Sierrade Perijd,
E of Manaure,El Podrido, 1550-1600 m, 16 Nov 1959 (9
fl-fr), Cuatrecasas et al. 25393 (COL, US), 17 Nov 1959
(6), Cuatrecasaset al. 25420 (COL, US); Mun. Manaure,
rd. Manaure-ElCinco, 1850 m, 10 Nov 1993 (d), Franco
et al. 4522 (BG), (9 fl), Franco et al. 4523 (BG). CHOC6:
Mun. San Jose del Palmar,Vrda. El Sinai, 1650 m, 19 Mar
1994 (9 fl-fr), Franco et al. 4562 (BG, HUA); EmisoraLa
Sirena, 3 km W of La Mansa, 2300-2400 m, 16 Jan 1979
(9), Gentry et al. 24195 (BG, COL, MO). CUNDINA-
MARCA:Mun. Tena,Vrda.El Rosario, rd. to Lagunade Pe-
dro Palo, 2100 m, 7 Aug 1984 (9 fl), Barrera 228 (COL);
Granada,Subia, 2225 m, 12 Mar 1993 (6), Franco et al.
4343 (BG, COL); Guayabetal,rd. Guayabetal-Villavicen-
cio, 1030-1150 m, 29 May 1993 (9 fl-fr),Francoetal. 4456
(BG) & 4457 (BG); rd. Caqueza-PuenteQuetame, 13 Mar
1985 (9 fl-fr), Galeano et al. 642 & 645 (COL);rd. Bogota-
Lagunade Pedro Palo, ca. 2100 m, 18 Mar 1985 (9 fl-fr),
Galeano et al. 675 (COL). HUILA: Finca Merenberg,E of
Volcan Purac6,nr. Cauca border,2290 m, 3 Apr 1986 (st),
Gentry et al. 53945 (BG). MAGDALENA:SierraNevada de
Santa Marta,above Minca, 1680-1690 m, 16 Jan 1989 (9
fl), Gentryet al. 64746 (MO);SierraNevadade SantaMarta,
Finca Los Arroyitos, 1600-1700 m, 4 Oct 1972 (d), Kirk-
bride 2381 (BG, COL, NY, US); Mun. Santa Marta, San
Lorenzo, 2240 m, 14-15 Sep 1969 (st), Lozano C. 1078
(COL). META: Mun. Villavicencio,rd. to Guayabetal,900-
1000 m, 13 Feb 1995 (9 fl-fr), Franco et al. 4677 (BG,
COL, HUA, NY, US); Sierrade La Macarena,Macizo Ren-
jifo, 1300-1900 m, 6-20 Feb 1951 (9 fr), Idroboet al. 1092
(COL, LE, U, US), (6), Idroboet al. 1093 (COL, GH, US);
Sierrade La Macarena,CentralMtns., NorthRidge, 29 Dec
1949 (9 fl-fr), Philipson et al. 1982 (BM, COL, US). NA-
RINO: Mun. Ricaurte,ReservaNaturalLa Planada,1800 m,
May 1989 (9 fr), Beltran 13 (MO, PSO); Mun. Ricaurte,
nr. Ricaurte, 1400 m, 8 Oct 1993 (6), Franco et al. 4465
(BG); Mun. Ricaurte,Reserva La Planada, 1800 m, 8 Oct
1993 (st), Franco et al. 4468 (BG); Mun. Ricaurte,nr. Ri-
caurte, 1400 m, 8 Oct 1993 (9 fr), Franco et al. 4475 (BG);
Samaniego, 1535 m, Aug 1964 (6), Mora 0. 3185 (PSO);
Mun. San Francisco,rd. Pasto-Mocoa, between El Mirador
& San Francisco, 1500-2000 m, 27 Nov 1967 (9 fl-fr),
Mora 0. 4474 (COL);Mun. Leiva, nr.Leiva, 1500 m, 1 Sep
1991 (6), RamfrezP 4167 (COL).NORTE DE SANTANDER:
Ocafnaregion, nr. Apasica, Quebradadel Guarumal,1900-
2000 m, 26 Sep 1969 (6), Cuatrecasaset al. 27950 (COL,
NY, P, US); Ocafiaregion, Cercanfasde Apasica, La Playa,
1700 m, 26 Sep 1969 (9 fl-fr), Cuatrecasas et al. 27953
(COL, NY, P, US); Mun. Toledo,Vrda.El Encanto, 1800 m,
25 Jan 1995 (6), Franco et al. 4625 (BG, COL, HUA, US).
PUTUMAYO: Mocoa, Vrda. La Campucana,Finca La Mar-
iposa, 1350 m, 20 Apr 1994 (d), Franco et al. 5186 (BG,
COL), (9 fl-fr),Franco et al. 5206 (BG, COL),28 Apr 1994
(st), Franco et al. 5458 (BG), (6), Franco et al. 5497 (BG),
(9 fl-fr), Franco et al. 5498 (BG); Mun. Mocoa, between
60
La Campucana& San Antonio, 1000 m, 2 May 1994 (5),
Franco et al. 5515 (BG); Mun. Mocoa, Cgto. El Pepino, rd.
El Pepino-Sibundoy,Vrda. Las Mesas, 1250 m, 3-4 May
1994 (5), Franco et al. 5528 (COL); rd. Mocoa-Pitalito,
Sajonia, 1600 m, 5 May 1994 (9 fl-fr), Franco et al. 5536
(BG); rd. San Juan-Pitalito,between Suacira& Santo Do-
mingo, 2000 m, 5 May 1994 (9 fl-fr), Franco et al. 5541
(BG). QUINDIO: Mun. Montenegro, Vrda. Morelia Baja,
1240 m, 21 Apr 1991 (9 fl), Agudelo et al. 1429 (COL);
Mun. Pijao, rd. Pijao-Chili, 2800 m, 13 May 1990 (5), W-
lez et al. 1693 (COL); Mun. Pijad, Vrda. Carniceres,1840
m, 24 Aug 1993 (9 fr), Wlez et al. 3370 (COL). RISAR-
ALDA: Mun. SantaRosa, Vrda.El Crucero,Hda. La Colina,
1725 m, 24 Jul 1980 (5), Idrobo et al. 9975 (COL, U).
SANTANDER: Mun. Oiba, Vrda. El Guayabito,ca. 1400 m,
22 Jan 1995 (9 fl-fr), Franco et al. 4605 (BG, COL, HUA,
US); nr. Aratoca,ca. 1800 m, 22 Jan 1995 (9 fl-fr), Franco
et al. 4606 (BG, COL, HUA, US). TOLIMA: Mun. Santa
Isabel, Finca La Cima, 1860 m, 6 Aug 1980 (9 fl), Idrobo
et al. 10680 (COL). VALLE: Rio Sanquinini,La Laguna,
1250-1400 m, 10-20 Dec 1943 (5), Cuatrecasas15540 (F,
VALLE),(9 fr), Cuatrecasas15550 (COL,F, US); Rio Cali,
confluence of Rio Pichindecito& Rio Pichinde, 1580-1650
m, 7 Nov 1944 (9 fl-fr), Cuatrecasas 18785 (COL, F,
VALLE), (9), Cuatrecasas 18786 (F, US, VALLE),(9 fl-
fr), Cuatrecasas18787 (F, US, VALLE);Rio Pichinde,Que-
brada de Juntas, 2100 m, 23 Sep 1946 (st), Cuatrecasas
22015 (MO, VALLE);Monte La Guarida, above La Car-
bonera,between Las Brisas & Alban, 1950-2000 m, 16-24
Oct 1946 (5), Cuatrecasas 22511 (F, US); Quebradadel
Tigre, between Las Brisas & La Marina, 1670-1730 m, 28
Oct 1946 (9 fr), Cuatrecasas22682 (F, US, VALLE);Alto
del Dinde, between Cartago & Alcala, 1200-1260 m, 16
Nov 1946 (5), Cuatrecasas22891 (F, US, VALLE),(9 fr),
Cuatrecasas22902 (F, US); Mun. Argelia, rd. El Cedro-La
Argelia-La Aurora, 1900 m, 11 Oct 1983 (5), Devia 313
(COL, MO, TULV); Mun. Calima, Vrda. Campo Alegre,
Finca Los Bambues, 900 m, 5 Oct 1983 (9 fl), Devia 406
(BG, MO, TULV);Reserva Forestalde Yotoco, 1200-1900
m, Jul 1993 (5), Escobar M. s.n. (VALLE);Mun. Trujillo,
rd. Trujillo-Rio Frio, km 1, 1260 m, 11 Sep 1993 (9 fl-fr),
Franco et a!. 4177 (BG), (5), Franco et al. 44811 (BG);
Mun. Trujillo,rd. Trujillo,Naranjal,km 20, Rfo Sanquinini,
1I Sep 1993 (d), Franco et al. 4488 (BG); Mun. Yotoco,
rd. Media Canoa-Dari6n,El Caney, 1500 m, 15 Mar 1994
(5), Franco et al. 4525 (BG), (9 fl-fr), Franco et al. 4526
(BG), (d), Franco et al. 4527 (BG); 1 km E of Bitaco, ca.
1500 m, 16 Nov 1963 (9 fr), Hutchinsonet al. 3009 (F, NY,
US), (d), Hutchinsonet al. 3028 (COL, K, MO, NY, P, US),
(d), Hutchinsonet al. 3035 (COL, MICH,NY, US); Monte
Frio, Yanaconas, 1700-1850 m, 27 Feb-i Mar 1939 (5),
Killip et al. 33712 (US); Mun.Darien,Vrda.El Vergel,1550,
20 Jun 1970 (9 fl-fr),Mahecha 281 (UDBC); Mun. Darien,
Alto Calima, 1970 (9 fl-fr), Mahecha s.n. (UDBC 5780);
rd. Cali-Pichinde, 1700 m, 15 Nov-6 Dec 1979 (d), Rooden
et al. 251 (BG, MO, NY, U).
VENEZUELA. ARAGUA: ParqueNacional Henri Pit-
tier, Rancho Grande, 1400 m, 10 Aug 1953 (9 fr), Little
15437 (NY, VEN); rd. El Pauji-El Soccoro, S of El Consejo,
FLORA NEOTROPICA
1350-1400 m, 16 Jul 1979 (9 fl-fr), Steyermarket al.
118157 (U, VEN); nr. Estaci6n Rancho Grande,800-1300
m, Aug 1966 (6), Veldsquez92 (US, VEN, (9 fl-fr), Velds-
quez 93 (US, VEN), 26 Dec 1975 (6), Wood429 (VEN).
BARINAS:Mun. Pedreza, Alto del Aguada, 1200-1300 m,
18 Feb 1955 (9 fl), Bernardi1966 (NY); Altamira-Calderas,
850-1600 m, 1 Jun 1957 (9 fr), Bernardi6878 (NY). DIs-
TRITO FEDERAL: Nr. Tovar, 1860 (6), Fendler s.n. (G);
ParqueNacional El Avila, QuebradaQuintero,Altamira,6
Sep 1975 (9 fl-fr), Manara s.n. (VEN); Cerro Naiguata,
QuebradaBesenilla, 900-1000 m, 3 Nov 1963 (6), Stey-
ermark91899 (U); between rd. Tovar-Junquito& Hda. El
Lim6n, 1300-1500 m, 12 Feb 1966 (9 fl), Steyermark
94778 (NY, US, VEN); ParqueNacional Avila, nr.Estaci6n
Los Venados, 900-1000 m, Mar 1967 (d), Veldsquez244
(US, VEN). FALCON:Dtto. Bolivar, Sierra de San Luis,
CerroGalicia, 1500 m, 29 Mar 1984 (9 fr), Plowmanet al.
13448 (BG, K, F, MO, NY), (6), Plowmanet al. 13449 (BG,
F, K, MO, NY). MERIDA: Mun. El Carrizal, El Rinc6n,
1400-1600 m, 23 Jun 1954 (9 fl-fr),Bernardi1295 (F, NY);
Mesa Bolivar, 1450 m, I Nov 1959 (9 fl-fr),Bernardi7629
(VEN); Mun. Zerpa, Dtto. Andres Bello, La Azulita, 1500
m, 31 Oct 1979 (9 fl-fr),MarcanoBerti et al. 280-979 (U);
above La Azulita, 1375-1920 m, 25 Apr 1944 (st), Steyer-
mark56101 (VEN); nr. Merida,Feb 1970 (9 fr), Veldsquez
2093 (US). MIRANDA: Los Guayabitos, SE of Baruta, 11
Oct 1969 (9 fl-fr), Steyermark91644 (NY, US); Los Guay-
abitos, 1200-1400 m, Nov 1967 (6), Veldsquez249 (VEN),
(9 fl-fr), Veldsquez250 (VEN). SucRE: Peninsulade Paria,
NW of Irapa,1060-1270 m, 3 Mar 1966 (9 fl), Steyermark
94971 (F, US); Peninsulade Paria, Dtto. Cagigal, between
El Paujil& El Brasil, 850-890 m, 21 Feb 1980 (9 fr), Stey-
ermarket al. 121434 (MO, NY, U, VEN). YARACUY:Sierra
Aroa, Cerro Tigre, 10 km E of Aroa, 1000 m, 3 Apr 1980
(9 fl-fr), Liesner et al. 9987 (MO, U, VEN). ZULIA: Dtto.
Mara,nr.Puesto El Bosque, 1450-1600 m, 10-15 Dec 1982
(9 fl-fr), Buntinget al. 12087 (MO, U).
ECUADOR. CARCHI: Nr. Guadalupe,RR Ibarra-San
Lorenzo, 900 m, 8 Feb 1981 (9 fl-fr), Berg 1248 (AAU,
BG, COL, MO, QCA, U); ca. 6 km above Maldonado,2275
m, 23 May 1993 (st), Boyle et al. 1916 (BG). COTOPAXI:
Rd. Quevedo-Latacunga,nr. Esperanza(El Tigre), 1500 m,
16 Feb 1981 (6), Berg 1263 (BG, COL, NY, QCA, U); rd.
Quevedo-Latacunga,nr. El Palmar,ca. 800 m, 17 Feb 1981
(9 fl-fr), Berg 1279 (AAU, BG, COL, MO, QCA, U), (6),
Berg 1281 (AAU, BG, COL, GB, MO, NY, QCA, U). EL
ORO: Rd. Pifias-SantaRosa, km 12.5, ca. 600 m, 7 Oct 1979
(st), Dodson et al. 8941 (MO), 7 Oct 1979 (6), Dodson et
al. 8953 (MO, QCNE);Viscaya, nr. Zaruma,1250-1500 m,
24 Sep 1980 (9 fl-fr), Dodson et al. 10541 (MO, QCNE).
IMBABURA: Between Parambas& Collapi, 1000 m, 4 Jul
1949 (6), Acosta S. 12777 (F). LOJA: Nr. Alamor,ca. 1250
m, 29 Dec 1997 (9 fl-fr), Lozano et al. 901 (BG, LOJA).
Los Rios/PIcHINCHA:El Centinela, at crest of Montafias
de Ila, rd. PatriciaPilar-24 de Mayo, km 12, 600 m, 27 Nov
1978 (st), Dodson et al. 7305 (BG, QCNE), (6), Dodson et
al. 7335 (BG, QCNE). MANABi: MachalillaNationalPark,
San Sebastian,550 m, 21 Jan 1991 (6), Gentryet al. 72522
(MO, QCNE). MORONA-SANTIAGO:Rd. Lim6n-Quenca,
SYSTEMATICTREATMENT
ca. 15 km, Plan del Milagro, 1700 m, 30 Jan 1981 (9 fl-fr),
Berg 1233 (BG, COL, QCA, U); rd. Gualaquiza-SanJuan
Bosco, ca. km 13, ca, 1300 m, 16 Feb 1994 (9 fl-fr), Berg
et al. 1692 (BG, QCA). NAPO: Cant6n Baeza, Bermeja,
2200-2650 m, 14 Dec 1993 (9 fl), Alvarez 881 (QCA,
QCNE);Cant6nArchidonard. Hollin-Loreto, km 32, Chal-
lua Yacu, 1200 m, 14-16 Sep 1989 (9 fl-fr), Hurtadoet al.
2457 (MO, QCNE), 14-16 Sep 1989 (d), Hurtado et al.
2581 (BG, MO, QCNE); El Codo Alto, 1300, 12 Sep 1990
(i), Jaramillo et al. 12580 (QCA); Cant6n El Chaco, Rio
Quijos, Finca El Ave Brava, 1800-1900 m, 7-10 Sep 1990
(i), Palacios 5363 (BG, QCNE, MO). PASTAZA:Nr. Mera,
Rio Mangayuca,ca. 1100 m, 9 Dec 1955 (9 fl-fr),Asplund
18745 (S); Upper Rio Pastaza, San Francisco,ca. 1100 m,
4 Feb 1956 (d), Asplund 19215 (S); Tnte. Hugo Ortiz, 20
km NNE of Puyo, 950 m, 30 Jan 1981 (9 fl), Berg et al.
1117 (AAU, BG, COL, GB, MO, QCA, U); 2 km NE of
Mera, 1100 m, 27 Feb-19 Mar 1985 (9 fl-fr), Neill et al.
5994 (BG, MO, QAME, QCNE). PICHINCHA:Nr. Tandapi,
1350-1800 m, 11 Sep 1977 (6), Berg et al. 417 (BG, COL,
MO, NY, QCA, U), (9 fl-fr), Berg et al. 418 (AAU, BG,
COL, MO, QCA, U); old rd. Quito-Santo Domingo de los
Colorados, km 59, nr. Palmeras,ca. 1800 m, 14 Dec 1990
(9 fl-fr), Berg et al. 1646 (BG, QCA); old rd. Quito-Santo
Domingo de los Colorados, km 59, Reserva Floristica-
Ecol6gica Rio Guajalito,1800-2000 m, 24 Aug 1985 (9 fl-
fr), Jaramillo et al. 8145 (BG, G, GB, GH, K, MO, NY,
QCA, US); 5-6 km above Nanegalito, 1700 m, 21 Jul 1980
(9 fl-fr), Holm-Nielsen et al. 24409 (AAU). SUCUMBiOS:
Rd. Baeza-Lago Agrio, nr. Santa Rosa, 1500-1600 m, 21
Sep 1977 (d), Berg et al. 441 (BG, COL, QCA); Cant6n
Gonzalo Pizarro, Campo Bermejo 6 Norte, 30 km NW of
Lago Agrio, 1050 m, 31 Mar 1990 (st), Cerc'net al. 9418
(BG); 3.5 km N of Borja, 1850 m, 20 Sep 1980 (d), Holm-
Nielsen et al. 26350 (AAU). TUNGURAHUA:Rd. Bafios-
Puyo, between Rio Verde& Rio Negro, ca. 1400 m, 10 Jan
1981 (9 fl-fr), Berg 1204 (BG, COL, QCA); Banios,Los
Llanganates,18 km from Topo, 1800-2000, 9 Mar 1995 (9
fl-fr), H. Vargas et al. 355 (BG, QCNE). ZAMORA-
CHINCHIPE: Rd. Loja-Zamora, nr. San Ram6n (hydro-
electric station), ca. 1900 m, 3 Jan 1991 (d), Berg et al.
1650 (BG, LOJA,QCA); rd. Loja-Zamora,km 28, 1900 m,
15 Feb 1994 (9 fl), Berg et al. 1688 (BG); rd. Loja-Zamora,
nr. Sabanilla, 1600-1700 m, 4 Sep 1975 (9 fl-fr), Little et
al. 227 (COL, LOJA,Q).
PERU. CAJAMARCA:La Palma, 10 km NW of Chirinos,
1780 m, 5 Feb 1988 (st), Gentry et al. 61217 (BG, MO);
Jaena,E.P.S.El Champe, 1500-1800 m, 3 May 1980 (9 fr),
Rios T 100 (MOL);Prov.Cutervo,Quebradade SantaRosa,
W of rd. San Andres-Santo Tomas, 2400 m, 4 Nov 1991
(J), Sanchez V et al. 6060 (F). Cuzco: Prov.Paucartambo,
rd. Pilcopata-Paucartambo,km 17, 900 m, 2 Aug 1988 (9
fl-fr), Berg et al. 1630 (BG, COL, Centrode Medicina An-
dina, Cuzco); Prov. Paucartambo,rd. Pilcopata-Paucar-
tambo, ca. 1600 m, 2 Aug 1988 (9 fl-fr), Berg et al. 1632
(BG, COL, Centrode MedicinaAndina,Cuzco); Prov.Pau-
cartambo,rd. Pilcopata-Paucartambo,ca. 1600 m, 2 Aug
1988 (d), Berg et al. 1633 (BG, COL, Centrode Medicina
Andina,Cuzco); Prov.Quispicanchis,Camanti,Mariri,720
61
m, 9 Sep 1990 (Y fl-fr), Timand913 (BG, MO), (d), Timand
918 (MO); Prov. Quispicanchis, between Inambari &
Quince Mil, 500-600 m, s.d. (Y fl-fr), Vargas C. 16517
(US). HUANUCO: Rd. Huanuco-TingoMaria, below Chin-
chao, ca. 1800 m, 24 Nov 1997 (6), Berg et al. 1737 (BG,
COL, MOL);nr.Tingo Maria,9 Aug 1940 (Y fl-fr),Asplund
12928 (S). JUNiN: Above San Ram6n, 1400-1450 m, 8-12
Jun 1929 (6), Killip et al. 24737 (F, NY, US); La Merced,
1350 m, 27 Aug-I Sep 1923 (9), Macbride5741 (F). MA-
DREDE Dios: Prov. Manu, nr. Shintuya,ca. 500 m, 30 Jul
1988 (juv), Berg et al. 1610A (BG). PASCO:Rd. Oxapampa-
Llaupi-Cerrode Pasco, ca. 13 km, ca. 2100 m, 29 Nov 1997
(6), Berg et al. 1759 (BG, COL, MOL); rd. Oxapampa-
Llaupi-Cerrode Pasco, ca. 13 km, ca. 1950 m, 29 Nov 1997
(9 fl-fr), Berg et al. 1762 (BG, COL, MOL); 20 km NE of
Villa Rica, rd. to Cacazu, 1600 m, 4 Jul 1988 (6), Gentry
et al. 63245 (BG, MO, MOL); Prov. Oxapampa,Gran Pa-
jonal, W of Chequitavo,1200 m, 10 Apr 1984 (9 fl-fr),D. N.
Smith6905 (BG, MO, MOL). PUNO:Prov. Sandiabetween
Quinsacu& Munu,5 Aug 1965 (6), VargasC. 16381 (US).
SAN MARTiN:Prov. Rioja, rd. Rioja-Pomacocha, nr. km
391-392, ca. 1750 m, 8 Dec 1997 (d), Berg et al. 1798
(BG, COL, MOL); Prov. Rioja, rd. Rioja-Pomacocha, nr.
km 378, ca. 2200 m, 8 Dec 1997 (6), Berg et al. 1801 (BG,
COL, MOL), km 390-394, 1910-2040 m, s.d. (9 fl-fr),
D. N. Smith4525 (BG, MO). UCAYALI: Rd. Pucallpa-Tingo
Maria,ca. km 145, near Huipoca, ca. 400 m, 26 Nov 1997
(9 fl-fr), Berg & Franco 1750 (BG, COL, MOL); rd. Pu-
callpa-Tingo Maria,nr. Boquer6n,ca. 400 m, 26 Nov 1997
(d), Berg & Franco 1753 (BG, COL, MOL). WITHOUT
LOCALITY: 1784 (st), Ruiz & Pavon s.n. (G).
BOLIVIA. BENI:Prov. Ballivian, rd. Yucumo-Caran-
avi, ca. km 13, ca. 900 m, 4 Mar 1994 (st), Berg 1713 (BG,
COL, LPB); Prov. Ballivian, SerraniaPil6n Lajas, 24 km
SW of Yucumo,rd.Yucumo-Quiquibey,km 24, 850-900 m,
3-7 May 1991 (st), Killeen et al. 3072 (BOLV,USZ); rd.
Yucumo-Rurrenabaque,km 28, Rio Hondo, 12 Mar 1989
(9 fl-fr), D. N. Smith et al. 12856 (LPB). COCHABAMBA:
Rd. Cochabamba-VillaTunari,km 96, 1800 m, 6, Dec 1995
(6), Berg et al. 1724 (BG, BOLV,COL, LPB, USZ); Prov.
Chapare,Locotal, 1600 m, 5 Feb 1929 (9 fl-fr), Steinbach
9030 (BM, E, F, G, GH, K, MO, NY, S, U), 20 Feb 1929
(st), Steinbachs.n. (US). LA PAZ:Prov. Sud Yungas,Chu-
lumani,rd. to Ocobaya, 1800 m, 28 Jun 1987 (9 fl-fr),Beck
12082 (BG, LPB); rd. Yucumo-Caranavi,nr. La Cascada,
ca. 5 km SSW of Quiquibey,ca. 1100 m, 4 Mar 1994 (9 fl-
fr), Berg 1715 (BG, COL, LPB); San Carlos, Rio Mapiri,
750 m, 7 Sep 1907 (st), Buchtien2118 (NY, US); Prov.Sud
Yungas,Rio Boopi, San Bartolom6,1-22 Jul 1939 (6), Kru-
koff 10304 (A, F, G, K, MICH, MO, NY, S, U, US); Prov.
Larejaca,Tuiri, 12-30 Sep 1939 (6), Krukoff10811 (A, G,
K, MICH, MO, NY, S, U, US).
This species is highly variable not only morpho-
logically but also ecologically, considering the wide
elevational range inhabited and the occurrence in rel-
atively dry habitats in some regions and in very wet
habitats in others. Attempts to distinguish (at least at
the subspecific level) formal entities related to distri-
62
bution and ecology have failed, partlybecause of the
ratherconfusing patternsin the morphologicaldiffer-
entiationand partlybecause of lack of sufficientcol-
lections from many parts of its range of distribution.
But it is possible to recognize some more or less dis-
tinct informal entities, representingextremes of the
variation.
In material from Venezuela and the eastern Cor-
dillera in Colombia, southwardto Caqueta,and from
the Sierrade la Macarena,the indumentumis (rather)
sparse and the upper surfaceof the lamina is usually
smooth. The lamina is usually coriaceous to subcor-
iaceous, and the reticulum veinlets are often thick,
and, therefore,the areoles small beneath.This mate-
rial includes the types of Cecropiacoriacea, C. mon-
iquirana, C. palmatisecta, and C. philipsonii.
Another type is found in the western part of Co-
lombia from Antioquia to Narinloand adjacentEc-
uador(Carchi).It is largely associatedwith relatively
dry areasin the westernCordillera,the CaucaValley,
andpartsof the centralCordillera.It is distinctlyhairy
(hirtellousto subtomentose)on the veins of the lam-
ina beneath,scabrousabove, and at least in the south-
ern and northernpartof its rangeof distributionoften
without arachnoidindumentumin the areoles of the
laminabeneath.This type is most clearly represented
by the material described as Cecropia caucana and
C. danielis, but it also includes the type material of
C. strigilosa. In the Cauca Valley the stipules and
spathes are usually reddish to purplish, and the pis-
tillate inflorescencescan be ? pendulousat fruit,due
to relatively long peduncles. South of the Cauca
Valley (Narinloand adjacentEcuador)the stipulesand
spathesare greenishto yellowish, the peduncleof the
pistillate inflorescencesis short, the arachnoidindu-
mentumin the areoles is often lacking, the internodes
can be completely filled by brownpith, and the num-
ber of lamina segments is relativelysmall, mostly 8-
9, as in the type of C. caucana. In (northern)Antio-
quia (as in the type of C. danielis), the smaller veins
of the lamina are often prominentbeneath, whereas
normally (almost) plane, the arachnoidindumentum
is often lacking in the areoles of the lamina beneath,
but in contrastto the materialoccurringsouth of the
Cauca Valley, the numberof lamina segments is 9-
11. In adjacentwetterpartsof ColombiaandEcuador,
the lamina has usually 11-13 segments and the in-
dumentumon the veins beneath is short and incon-
spicuous.
On the easternslopes of the Andes, from Caqueta'
to Morona-Santiago,three morphologicaltypes can
be distinguished.One of them resemblingthe type of
the wetter partsin western Colombia and Ecuador,a
FLORA NEOTROPICA
second one with relatively few (ca. 9) lamina seg-
ments and densely hairy, sericeous to subhirsuteon
the stipules and often also on the spathes,and a third
one (only found in Caqueta')with ratherdense arach-
noid indumentumon the leafy twigs and the petioles.
The second type has been described as Cecropia
hachensis and occurs in two color morphs,at least in
parts of its range of distribution:a green morph and
a reddishto purplishone.
Material in the coastal region of Ecuador,in El
Oro and Manabf,at 550-1500 m, shows similarities
to Cecropia montana in the whitish stipules and
spathes and in the presence of ratherlong white uni-
cellularhairsin the trichilia,but the numberof lateral
veins in the free part of the midsegmentis up to ca.
20 pairs, thus distinctly less than the minimumnum-
ber in C. montana.
In the southernpart of the species range at least
three types can be recognized. The most prominent
one is found in Bolivia and adjacent Peru. It has
blackish and subpersistent stipules and incisions
down to ca. 6/10-7/10 or incisions down to near the
petiole. More northwardin Peru, the stipules vary in
color from brownto greenishand are eithercaducous
or subpersistent.The lattertype comprisesthe type of
the species. In some specimensfrom Bolivia andPeru
(as in the type of the species) the lamina has narrow
segments, to 4 cm broad in medium-sizedleaves. In
Bolivia (Cochabamba)a purplish morph occurs be-
sides the predominantgreen morph. In the majority
of the materialfrom Peru,the numberof lateralveins
in the free partof the midsegmentvaries from ca. 23
to 35 pairs, but in the third type the number varies
between 15 and 20 pairs.The thirdtype is represented
by Berg et al. 1632, 1750, and 1753, the latter two
found at ca. 400 m.
A remarkabletype is representedby materialfrom
southern Ecuador and northen Peru on the eastern
slopes of the Andes and from Costa Rica and western
Panama. Material from Ecuador was recently de-
scribed as Cecropiavillosa (Berg & FrancoRosselli,
1993). Although in general features quite similar to
C. angustifolia, it is distinct in villous indumentum
and internodesentirelyfilled with brownpith.Further
studies on the genus revealed that this material did
not differ clearly from collections from Costa Rica
which were named C. polyphlebia and were made
from trees not inhabited by ants (Longino, 1989a);
the Ecuadoriantrees are also not occupied by ants.
Ants are absent in spite of the common presence of
trichilia, producing abundantMiillerian bodies (cf.
Janzen,1973: 17, fig. 1). The absenceof ants is likely
caused by the ample pith in the internodes.In C. an-
SYSTEMATICTREATMENT 63

gustifolia the pith in the intemodes is usually very stipules 12-20(-28) cm long, orange-redto pinkish
sparse,rarelyample (as in Berg et al. 1759). The un- or partly whitish, caducous, (appressed-)puberulous
usual disjunctionbetween two sets of specimens and to hirtellous, or on the ribs to subhirsute,also with
the fact that they fill gaps in the distributionof C. dense arachnoidindumentumand ratherdense brown
angustifolia led finally to the decision to include the
pluricellularhairs outside, ? densely hairy inside.
materialinitially recognizedas C. polyphlebiaand C. Staminateinflorescencessolitary or in pairs, the pe-
villosa in C. angustifolia. This villous type tends toduncle erect, the spikes ? spreadingto pendulous;
have more lateralveins [ca. 25-35(-40) pairs] in the peduncle 9-13 cm long, with sparse arachnoidindu-
free partof the midsegmentof the laminathanis usual mentumand brown pluricellularhairs; spathe 12-15
in C. angustifolia. It is remarkablenot only that thiscm long, white, with dense arachnoidindumentum
villous type with ample pith arose disjunctly,but alsoand sparsebrownpluricellularhairsoutside, glabrous
that both in Ecuadorand CentralAmerica it is mor- or sparselyhairy inside; spikes ca. 10-25, 6-17 X ca.
phologically linked to a hairy type without ample 0.3 cm (yellow to pale orange), with stipes 0.8-1.3
pith. In Ecuador, this type includes the type of C. cm long and sparsely puberulousin the upper part;
hachensis, and in CentralAmerica it is represented rachis hairy. Staminateflowers: perianthtubular,1-
by collections with mostly brownish indumentum 1.5 mm long, glabrous, the apex slightly convex,
from Nicaraguaand Honduras. smooth; filaments flat; anthersca. 0.8 mm long, ap-
This species is closely related to Cecropia mon- pendiculate,detachedat anthesis (?). Pistillate info-
tana. The differencesbetween the two species aredis- rescences solitary or in pairs, pendulous; peduncle
cussed underthe latter. 9.5-19 cm long, sparselypuberulousto hirtellous(to
subhispid);spathe not seen; spikes 4-5(-6), 9-17 X
Vernacular names. Colombia: guarumomoreno
(0.5-)0.7-0.8 cm, sessile or with stipes to 0.5 cm long
(Caldas); llarumo blanco, llarumo negro (Cauca);
and minutelypuberulous;rachishairy.Pistillate;flow-
guarumonegro (Norte de Santander);patade gallina,
ers: perianthca. 1.5-2 mm long, with arachnoidin-
orumo (Santander).Venezuela:jagrumo(Falc6n,Me-
dumentumbelow the apex or also on the marginof
rida). Peru: yungapara (Cajamarca); guarumbo,
the apex outside, also below the style channelinside,
purma(San Martin).
the apex ? convex, punctate to muriculate; style
rathershort;stigmatongue-shapedto subpeltate.Fruit
ellipsoid, ca. 1.8 mm long, smooth.
5. Cecropia annulata C. C. Berg & P.Franco,Novon
Distribution (see Fig. 10.2). Bolivia (La Paz and
6: 245. 1996. Type. Bolivia. La Paz: Nr. Sapecho,
Beni), in non-inundatedforest and secondarygrowth,
1 Mar 1994 (Y fl-fr), Berg 1704 (holotype:LPB;
at low elevations.
isotypes: BG, COL, MO, NY).
Specimensexamined.BOLIVIA.BENI:Prov.Balli-
Tree,to 25 m tall; trunkwith prominent(annular) vidn,Serranfadel Pil6n,8-10 km from,Yucumo,19 May
scars of the stipules. Leafy twigs 2-4.5 cm thick, 1989(9 fl-fr),D. N. Smithet al. 13264(BG,BOLV,LPB,
(dark) green, hispidulous with curved to uncinate USZ).LA PAZ: Prov.SudYungas,nr.Sapecho,26 Feb-i
hairs. Lamina subcoriaceousto coriaceous, ca. 30 X Mar 1994 (st), Berg 1698A(BG),(d), Berg 1699 (AAU,
30 cm to 75 X 75 cm, the segments 8-10, the free BG,COL,LPB,MO),(6) Berg1701(BG,COL,LPB);nr.
parts of the upper segments obovate to elliptic, the Tucupi(= Tullupi),ca. 30 kmSE of PalosBlancos,nr.Rio
upper ones sometimes slightly lobate, the incisions Beni,5 Mar1994(9), Berg1717(AAU,BG, COL,LPB,
down to 6/10-8/10(-9/10); apices obtuse; uppersur- MO,NY).
face smooth to scabridulous, sparsely to rather This species is probablyclosely relatedto Cecro-
densely minutely puberulous to strigillose on the pia polystachya. It differs from the latterin the mar-
(main) veins, initially also with sparse arachnoidin- ginally loop-connectedlateralveins in the free partof
dumentum;lower surfacesparselypuberulousto stri- the midsegment, the (very) sparse arachnoidindu-
gillose with straight,curved, or uncinatehairs on the mentumon the petiole, the (rather)short white (uni-
(main) veins, with arachnoidindumentum(almost) cellular)hairsin the trichilia,and the entire(or rarely
confined to the areoles or almost absent;lateralveins slightly) lobate midsegment. However, C. annulata
11-16(-20) pairs, marginally loop-connected, the also resembles C. concolor subsp. engleriana, from
lower ones branched; petiole ca. 25-70 cm long which it differs in the less deeply incised lamina,the
sparsely (minutely) puberulousand also with sparse smallernumberof lateralveins in the free partof the
arachnoidindumentum;trichiliafused, the brownin- midsegment, and the non-peltatestigmas. The char-
dumentumintermixedwith (rather)shortwhite hairs; acters of this species look like a mixtureof those of
64 FLORA NEOTROPICA

the two related species named above and might in- stipes 0.3-1.5 cm long and subsericeous or with
dicate an origin by hybridization.The species is rel- arachnoidindumentum;rachis hairy.Staminateflow-
atively rare in secondary growth. In the field, it caners: perianthtubular,ca. 2 mm long, glabrousor pu-
be easily recognized by the orange to pinkish young berulous below the apex, the apex plane to slightly
leaves. The filamentsof the stamenscan be distinctly convex, hispidulousto muriculate;filamentsswollen;
shorterthan the perianthor they are longer, with the anthers 0.5-0.7 mm long, appendiculate,remaining
apices beyond the aperture;this might imply that the attachedto the filamentat anthesis(?) Pistillate inflo-
anthersare not detachedbefore anthesis. rescences (usually) solitary, erect at anthesis to pen-
dulous in fruit;peduncleca. 5-12 cm long, at least in
Vernacular name. Bolivia: ambaibonegro(Beni).
the upperpartor the lower parthirsuteto subvillous,
also with short hairs and/or sparse arachnoidindu-
mentum;spathe 10-14 cm long, the color and indu-
6. Cecropia bullata C. C. Berg & P. Franco,Fl. Ec- mentumas in the staminateinflorescence;spikes 2 or
uador48: 14, t. 3. 1993. Type.Ecuador.Pichincha: 1, 8-12 X ca. 1 cm, to 21 X ca. 2-3.5 cm in fruit,
Rio Guajalito,ca. 2 km N of Palmeras,rd. Quito- subsessile or with stipes ca. 0.5 cm long andhirtellous
San Juan-Chiriboga-Enpalme,km 59, ca. 1850 m, to hirsute;rachishairy.Pistillateflowers: perianthca.
15 Dec 1990 (Y), Berg et al. 1648 (holotype: 4-5 mm long, with arachnoidindumentumbelow the
QCA; isotype: BG). Fig. 15 apex, sometimes also sparsely in the style channel
inside, sometimes also short stiff hairsjust below the
Tree, to 20 m tall. Leafy twigs 2-4 cm thick,
apex outside, the apex slightly convex, muriculateto
densely white to brownishvillous to subsericeousor
smooth or hispidulous around the aperture; style
to subhirsute;internodesentirely or partlyfilled with
ratherlong, straight,hairy; stigma penicillate. Fruit
brown pith. Lamina subcoriaceous (when dry often
oblongoid, ca. 3 mm long, smooth.
brittle),ca. 30 X 30 cm to 70 X 70 cm, the segments
7-9, the free parts of the upper segments elliptic to Distribution (see Fig. 11.3). On the western
obovate, the incisions in the upperpartof the lamina slopes of the western Cordillerain Colombia, from
down to 5/10-6/10, the lowerpartof the laminalobed; Antioquiato Narinlo,and in northwesternEcuador,in
apices (of the uppersegments) acuminate;uppersur- (sub)montane(cloud) forest, at ca. 1700-2000 m.
face smooth, bullate, (sub)glabrous; lower surface
Specimens examined. COLOMBIA.ANTIOQUIA:
with concave areoles, ? densely (sub)hirsuteto hir- Mun.Frontino,Cgto.Nutibara, rd.Nutibara-La Blanquita,
tellous to subtomentoseor sparsely hirtellous to pu- Murriregion,1600m, 4 Mar1955(9 fl-fr), Franco et al.
berulous (to subglabrous)on the veins, on the main 5550 (BG, COL,HUA),1650m, (5), Franco et al. 5553
veins often also with elongate brown pluricellular (COL,HUA).CHOCO: Mun.SanJosedel Palmar, Vrda.El
hairs, with (sparse) arachnoidindumentumin the ar- Sinai, ca. 1800 m, 19 Mar 1994 (d), Franco et al. 4559
eoles, mostly extendingto the reticulumor also dense (BG,HUA),(9 fl-fr),Francoet al. 4561 (BG,HUA).NA-
arachnoidindumentumlargely covering the area be- RINO: Mun.Ricaurte, ReservaNaturalLaPlanada, ca. 1700
tween the secondary veins, often extending to the m, 19 Jun 1990 (5), Benavides 11138 (PSO); Mun. Ri-
main veins; lateralveins in the free part of the mid- caurte,rd. to ReservaNaturalLa Planada,1800m, 8 Oct
1993(9 fr), Franco et al. 4467 (BG),22 Feb 1995(9 fl-
segment 10-20 pairs, submarginallyloop-connected,
fr), Franco et al. 4698 (COL); La PlanadaReserva, 7 km
unbranched(or the lower ones branched);petiole ca. from Chucunes, 1800 m, 7 Jan 1988 (9 fl-fr), Gentryet al.
40-55 cm long, puberulousto hirtellousor also whit- 60519 (BG, MO, PSO); Mun. Ricaurte,ReservaNaturalLa
ish to brownishhirsuteto (sub)villous,often also with Planada,20 Jul 1987 (9 fl-fr), Restrepoet al. 378 (MO), 14
ratherdense arachnoidindumentum(or subglabrous); Jan 1989 (5), Restrepo441 (MO). VALLE: Mun. El Silen-
trichilia fused, the brown indumentum intermixed cio, Hda. Himalaya,W of Yumbo, 1840 m, 4 Feb 1989 (5),
with sparse short whitish to brownish (unicellular) Gentry et al. 65484 (BG, MO); Mun. Cali, rd. Cali-Bue-
hairs; stipules (8-)20-36 cm long, pink to dull red, naventurakm 18, rd. to Dapa, km 4, Finca Zingara,1900 m,
densely whitish to brownish villous to subhirsuteto 10 Jul 1944 (9 fl-fr), Giraldo-Gensini et al. 411 (NY,
TULV).
subsericeous or also with short arachnoidindumen-
ECUADOR.CARCHI: Cant6n Mira, N of El Carmen,
tum outside, sparsely hairy to glabrous inside. Sta-
rd. to Chical, 2000-2200 m, 10 Feb 1992 (9 fl-fr), Gudinio
minateinflorescencesin pairsor solitary,the peduncle et al. 9720 (QCNE);Cant6nMira,rd. to Chical,N of El
erect, the spikes erect to ? spreading;peduncle ca. Carmen,2000-2200 m, 10 Feb 1992 (9 fl-fr), Palacios et
6.5 cm long, white-hirsute;spathepink to dull red, ? al. 9720 (BG, QCNE). PICHINCHA:Nr. Saloya, 1800 m, 9
densely villous to subhirsuteto hirtellousoutside,gla- Sep 1943 (st), Acosta S. 5848 (F); rd. Atenas-Tandapi,1700
brous inside; spikes 4-10, 6-15 X 0.7-0.8 cm, with m, 3 Nov 1991 (d), Jaramilloet al. 14521 (NY, QCA); rd.
SYSTEMATICTREATMENT 65

)~~~~~~~~~~~~~~~~~~~~~~~~~~~r

I~~~~~~~~~~

(I

FIG. 15. Cecropia bullata. 1. Lamina, reduced. 2. Apex of lamina and venation (Berg et al. 1648). 3. Stipules. 4.
Fruit~~~~~~~~~~~~~~~
(Fac ta.4599(rmF.Ecao 8 5 19,mdfe.
Staminateinflorescence with spathe. 5. Staminateinflorescenceat anthesis. 6. Staminateflower and stamen (Franco et al.
4559). 7. Pistillate inflorescence with spathe (Franco et al. 4461). 8. Pistillate inflorescenceat anthesis and base of petiole
with trichilium(Zak 1312). 9. Pistillate inflorescence in fruit and trichilium (Franco et al. 4467). 10. Pistillate flower. 11.
Fruit(Franco et al. 4561). (From Fl. Ecuador48: 15. 1993, modified.)
66
Quito-San Juan-Chiriboga-Empalme,Rio Guajalito,ca. 2
km N of Palmeras,km 59, 1800-1900 m, 29 Feb 1992 (d),
Jaramillo et al. 14651 (QCA); km 59, 15 km NE of rd., ca.
1700 m, 23 Sep 1986 (Y fl-fr), Zak 1312 (BG, MO, NY,
QCA).
This species is closely related to both Cecropia
telealba and C. gabrielis, from which it can be easily
distinguishedby the surfacesof the lamina:distinctly
bullateabove andwith concave areolesbeneath.From
C. telealba it can also be distinguishedby the absence
of dense arachnoidindumentumon the uppersurface
of the laminaand from C. gabrielis by the hairy (vil-
lous to subhirsute)leafy twigs. The leaves are more
distinctly bullate and with less dense arachnoidin-
dumentumon the laminabeneathin the southernpart
of the rangeof distribution(EcuadorandNarifio)than
in the northernpartof the range (Valle to Antioquia).
Since the villous indumentumis sometimes less pro-
nouncedlypresentin the northernpartandthe stipules
and spathes of C. gabrielis can be (partly) sparsely
(sub)villous in Antioquia, C. bullata can (only) be
distinguishedby the presenceof villous indumentum
on the leafy tiwgs (and mostly also on the petioles).
The trees are not inhabitedby ants.
7. Cecropia chlorostachya C. C. Berg & P. Franco,
Caldasia 24: 233. 2002. Type. Peru. San Martin:
Prov. Rioja, rd. Rioja-Pomacocha, nr. km 391-
392, ca. 2000 m, 8 Dec 1997 (Y fl-fr), Berg &
Franco 1802 (holotype: MOL; isotypes: BG,
COL). Fig. 16
Tree, to 22 m tall. Leafy twigs 2-3.5 cm thick,
whitish (due to the indumentum),green, white- to
brownish-subvillous to -subhirsute to -hirtellous,
sometimes only near the scars of the stipules, also
with dense arachnoidindumentumand brown pluri-
cellular hairs. Lamina (sub)coriaceous,ca. 50 X 50
cm to 60 X 60 cm, the segments 7-10, the free parts
of upper segments elliptic to oblong, the incisions in
the upperpartof the lamina down to ca. 6/10, apices
acuminate; upper surface (almost) smooth, rather
sparsely puberulousto hirtellous, also sparse arach-
noid indumentum;lower surfacerathersparselysub-
villous to subhirsuteto hirtellous on the veins, with
arachnoidindumentumin the areoles, extending to
the main veins, on the main veins also brown pluri-
cellular hairs, most densely near the petiole; lateral
veins in the free part of the midsegment ca. 15-20
pairs, submarginallyloop-connected,mostly unbran-
ched; petiole ca. 25-55 cm long, white subvillous to
subhirsuteandwith dense arachnoidindumentum;tri-
chilia fused, the brown indumentumintermixedwith
FLORANEOTROPICA
shortand (partlywith) long white hairs,the Miillerian
bodies pinkish;stipules 15-32 cm long, subpersistent,
pinkish, densely (sub)villous, also with dense arach-
noid indumentumand brown pluricellularhairs out-
side, sparselyto (partly)densely hairy inside. Stami-
nate inflorescencesin pairs, erect, or the spikes ?
spreading;peduncle ca. 3-6 cm, white, subhirsuteto
hirtellous and with dense arachnoid indumentum;
spathe ca. 12-15 cm long, white, (sub)villous, with
dense arachnoidindumentumand with brown pluri-
cellularhairs outside, inside hairy;spikes 7-8, 15-17
X 0.6-0.8 cm, before anthesis yellow, at anthesis
green, with stipes 0.5-1 cm long andhirtellous;rachis
hairy with stiff and crinkled (arachnoid)hairs. Sta-
minateflowers:perianthtubular,1.8-2 mm long, with
arachnoid indumentum below the apex, the apex
plane, sparsely muriculate;anthersca. 0.8 mm long,
with short appendages,detachedat anthesis,remain-
ing attachedto the filamentby 2 filiformconnections
between the connective and the upper marginof the
filament.Pistillate inflorescencesin pairs or solitary,
erect to ? spreadingwith the spikes straightto curved
upward,subtendedby whitish, caducous bracts,to 3
cm long; peduncleca. 7-10 cm long, the indumentum
as in the staminateinflorescence; spathe 10-12 cm
long, pinkish, the indumentumas in the staminatein-
florescence;spikes 4, 4-12 X ca. 0.-0.5, to 15 X 1
cm in fruit, subsessile or with stipes to 0.4 cm long
with dense arachnoidindumentum;rachis hairy with
stiff and crinkledhairs.Pistillateflowers: flowersof-
ten basally connate; perianthca. 2.5 mm long, with
arachnoidhairsbelow the apex outside, andbelow the
style channelinside, the apex plane to slightly convex
and muriculate;style short;stigma comose. Fruit el-
lipsoid to oblongoid, ca. 1.5-2 cm long, (almost)
smooth.
Distribution (see Fig. 8.5). NorthernAndeanPeru
(San Martin and Amazonas), in cloud forest, at ca.
1750-2000 m.
Specimens examined. PERU. AMAZONAS: Prov.
Chachapoyas, RodriguesMendoza,Cochamal, Montania de
Yanamonte, 4 Jul 1991(? fl-fr),C. Diazet al. 4567 (MO).
SAN MARTiN: Prov.Rioja,rd.Rioja-Pomacocha, ca. 1750
m, 8 Dec 1997(d), Berget al. 1799(BG);Prov.Rioja,rd.
PedroRuiz-Moyobamba, km390,Venceremos, 2100m, 7-
9 Aug 1983 (i), D. N. Smithet al. 4686 (BG).
This species is distinguished by the dense and
white indumentumon variousparts,the subpersistent
stipules, and the green spikes of the staminateinflo-
rescence. Green spikes sometimes occur in Cecropia
distachya, but only before anthesis (as in Berg et al.
1597). The species is probablynot myrmecophytic.
SYSTEMATICTREATMENT 67

FIG. 16. Cecropia chlorostachya. 1. Lamina, reduced (D. N. Smith et al. 4686). 2. Apex of lamina and venation. 3.

Stipuesadbaeofetioewihtrihilim(CDia et al. 4567).4.Stminatinflrescecew

ils c a sa Smith et al. 4686).6.Staminateflower(lngi

stn7Smafanarecnone(gt.J . .Pt i
a
'I" ? \
'0

FIG. 16. Cecropia chlorostachya.1. Lamina,reduced (D. N. Smithet al. 4686). 2. Apex of lamina and venation. 3.
Stipules and base of petiole with trichilium(C. Diaz et al. 4567). 4. Staminateinflorescencewith spathe.5. Pairof staminate
inflorescences at anthesis and base of petiole with trichilium (D. N. Smith et al. 4686). 6. Staminateflower (longitudinal
section). 7. Stamen and filament after detachmentof anther(Berg et al. 1799). 8. Pistillate inflorescence with spathe (C.
Diaz et al4567). 9. Pair of pistillate inflorescences at anthesis and base of petiole with trichilium. 10. Pistillate flower and
style. 11. Fruit(Berg et al. 1802). (From Caldasia24: 234. 2002.)
68 FLORA NEOTROPICA

8. Cecropia concolor Willdenow, Sp. P1. 4(2): 652. ? densely sericeous inside. Staminateinflorescences
1806; Berg, Acta Amazonica8(2): 164. 1978.Am- in pairs or solitary,pendulous;peduncle (3.5-)10-19
baiba concolor (Willdenow) Kuntze, Revis Gen. cm long, sparsely puberulousto hirtellous and with
P1. 2: 624. 1891. Type. Brazil. Para:Without lo- sparseto ratherdense arachnoidindumentum;spathe
cality, (Y), Sieber in herb.Hoffinannseggs.n. (ho- 10-20 cm long, white to yellowish or pinkish,puber-
lotype: B in herb. Willdenow, destroyed, photo- ulous to hirtellousand often with dense arachnoidin-
graphs in F, G, MO, US), herewith replaced by: dumentum outside, glabrous or (densely) sericeous
Brazil. Para:Mun. Belem, Mocambo, 8 Oct 1995 inside; spikes (8-)15-20, (1-)6-15 X 0.2-0.4 cm,
(6), Berg 1721 (neotype: MG; isoneotypes: B, with stipes 0.2-2 cm long and hairy or subglabrous;
BG, COL, INPA, NY, RB, U, UB). Fig. 17 rachishairy.Staminateflowers:perianthtubular,0.8-
CecropialeucocomaMiquel,in Martius,Fl. Bras.4(1): 1.5 mm long, glabrous, the apex almost plane; fila-
142. 1853. Ambaiba leucocoma (Miquel) Kuntze, ments slightly swollen; anthers0.5-0.6 mm long, ap-
Revis. Gen. P1.2: 624. 1891.Type.Brazil.Ama- pendiculate, detached at anthesis, reattachedat the
zonas:Nr.Manaus(= Barra),Dec 1850-Mar1851 marginsof the apertureby the appendages(?). Pistil-
( i ), Spruce1322ors.n.(holotype: M;isotypes:BM, late inflorescencesin pairs or solitary,pendulous;pe-
E, F, G, GH,K, LE,NY,P). duncle (3.5-)10-22 cm long, puberulousto hirtellous
CecropiaobovataRusby,Bull.NewYorkBot.Gard.6: and with sparse to ratherdense arachnoidindumen-
498. 1910.Type.Bolivia.LaPaz:SanBuenaventura, tum; spathe 10-17 cm long, the color and indumen-
15 Nov 1901(Y), R.S. Williams 645 (holotype:NY;
tum as in the staminateinflorescence;spikes 4, (2-)
isotypes:BM,K).
Cecropia maranhensis Snethlage, Notizbl. Bot. Gart. 5-18 X 0.4-1 cm, to 29 X 1-1.5 cm in fruit, sessile
Berlin-Dahlem 9: 172. 1924.Syntypes.Brazil.Mar- or with stipes to 1(-1.5) cm long and hairy: rachis
anhao:Turiaqu,2-3 Jan 1924 (d) (Y), Snethlage hairy. Pistillate flowers: perianth 1.5-2.5 mm long,
384 (B, destroyed,photographs ex B in F,G,MICH, with arachnoidindumentumon and below the apex,
MO),(cd),Snethlage 388 (B,destroyed, photographs absent inside, the apex plane; style short; stigma
ex B in F, G, GH,MICH,MO;isotype:F). (sub)peltate. Fruit ellipsoid to ovoid to oblongoid,
Tree, to 15 m tall; stem with prominentstipular 1.5-2.5 mm long, ? tuberculate.
scars. Leafy twigs 1.5-4 cm thick, green with yellow Distribution (Fig. 18.1). Lower and centralAm-
spots to purplish-brown,hispidulous,with curved or azon basin, extending to Bolivia and southernAma-
uncinate hairs, or also with sparse to rather dense zonian Peru (Cuzco and Madrede Dios), common in
arachnoid indumentum. Lamina subcoriaceous to secondarygrowth,in non-inundatedplaces or (in the
chartaceous,ca. 30 X 30 cm to 65 X 65 cm, the southwesternpart of its range) sometimes in tempo-
segments (7-)8-10(-11), the free parts of the upper rary inundatedplaces, at low elevations.
segmentsoblong to subobovateto oblanceolate,entire
or ? sinuate (or if juvenile, then to lobate), the in- specimensexamined.PERU.Cuzco:
Representative
cisions down to 1.5(-2.5) cm from the petiole; apices Prov.Paucartambo,betweenPilcopata& Atalaya,29 Jul
1988 (d), Berg et al. 1605 (BG, COL, Centrode Medicina
short-acuminateto (sub)acuteto obtuse;uppersurface
Andina,Cuzco);Prov.Paucartambo, rd.Pilcopata-Paucar-
smooth to scabridulous,sparsely puberulousto sub- tambo,km 12,2 Aug 1988(9 fl-fr),Berget al. 1628(BG,
hispidulous, also with sparse to ratherdense arach- COL,Centrode MedicinaAndina,Cuzco);Prov.Paucar-
noid indumentum,along the main veins often with tambo,rd.Pilcopata-Atalaya, halfway,24 Oct 1984(9 fr),
brown pluricellular hairs; lower surface minutely Maaset al. 6160(BG,U);Prov.Paucartambo, betweenTono
puberulouson the main veins, with arachnoidindu- & Pifiipifii(= Kosnipata),23 Nov 1965 (6), VargasC.
mentum in the areoles, on the smaller veins, or, at 16900 (US). MADRE DEDIos: Prov.Manu,nr.Shintuya, 30
least initially, also on the main veins; lateralveins in Jul 1988(9 fl-fr),Berget al. 1609 (BG,COL,Centrode
the free partof the midsegment 16-24 (or if juvenile MedicinaAndina,Cuzco);ParqueNacionalManu,Rfo
to 30) pairs, marginally loop-connected, mostly Manu,Cocha CashuStation, 19 Sep 1986 (6), Foster11435
branched;petiole 25-50 cm long, sparselypuberulous (BG, MOL,US), (9 fl), Foster11444 (BG);ParqueNa-
cional Manu,Rio Manu,Tayakome,29 Sep 1986 (9), Foster
and/or with sparse to (rather)dense arachnoidindu-
et al. 11528 BG, US).
mentum; trichilia fused, the brown indumentumin- BRAZIL.AMAPA: Estradado Marucca, 6 Nov 1979
termixed with (sparse) short white hairs; stipules 7- (6), Rabelo 62 (MG). AMAZONAS: Manaus,INPA, 29 Aug
15 cm long, white to pale yellow or pinkish,caducous 1973 (d), Berg 245 (U), (9 fl-fr), Berg 246 (A, MO, U,
or subpersistent,puberulousor sparselysericeous,of- km65, 21 Sep 1973(st),Berg
US);rd.Manaus-Caracarai,
ten with ? dense arachnoidindumentum,sometimes 279 (U), (9 fl-fr), Berg 280 (K,U); Rio Solimoes,Espfrito
(also) with ? dense brownpluricellularhairsoutside, Santo, between Boca de Januaca& Boca de Manaquiri,3
SYSTEMATICTREATMENT 69

I9,'

XIE ~~ ~

6 H '10 8 9
FIG. 17. Cecropia concolor. 1. Lamina, reduced. 2. Apex of lamina and venation (Berg et al. 1628). Cecropia
engleriana. 3. Stipules and bases of petioles with trichilia (Berg et al. 1629). 4. Staminateinflorescence with spathe and
bases of petioles with trichilia (Alarcon 74). 5. Staminate inflorescence at anthesis (Berg 1605) 6. Staminateflower and
stamen (Franco et al. 4688). 7. Pistillate inflorescenceafter anthesis and base of petiole with trichilium(Gentry 12442). 8.
Pistillate flower (Berg et al. 1629). 9. Pistil (Berg et al. 1104). 10. Fruit(Berg 1700).
70 FLORA NEOTROPICA

v~~~~~~~~~

1~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~

FIG. 18. Distributionmaps. 1. Cecropia concolor, triangles; C. elongata, hexagons; C. pachystachya,circles; both
C. concolor and C. pachystachya, crosses. 2. C. megastachya, hexagons; C. metensis, triangles; C. palmata, circles; C.
strigosa, open circles. 3. C. peltata, circles; C. schreberianasubsp. schreberiana,triangles; C. schreberianasubsp. antil-
larum,hexagons;both subsp.schreberianaand subsp.antillarum,crosses. 4. C.polystachya,circles; C. velutinella,triangles.
5. C. putumayonis,circles; C. reticulata, triangles. 6. C. gabrielis, triangles; C. sararensis, circles. 7. C. telenitida. 8. C.
pastasana, circles; C. subintegra,triangles.
SYSTEMATICTREATMENT
Oct 1973 (Y fr), Berg et al. P17590 (C, COL, F, INPA, K,
MG, MO, NY, P, S, U, US); Manaus, Reserva Ducke, 28
Sep 1962 (i), Duarte 7190 (INPA, K, RB); Manaus, Feb
1904 (cd + Y fl-fr),Huber(MG) 4175 (G, MG);nr.Manaus,
rd. Taruma-PontaNegra, km 13, 6 Sep 1973 (Y fl), Lleras
et al. P17553 (C, F, INPA, K, MG, MO, NY, S, U); Mun.
Manaus,ca. 90 km N of Manaus,Distrito Agropecuariada
SUFRAMA, 16 Jul 1992 (Y fl), Nee 43005 (INPA,K, NY);
nr. Manaus, Ponta Negra, 16 Mar 1967 (3), Prance et al.
4665 (F, INPA, NY, S, U); rd. Manaus-Itacoatiara,km 26,
Reserva FlorestalDucke, 26 Apr 1994 (Y fl-fr), Ribeiro et
al. 1293 (BG, INPA);rd. Torquato-Tapaj6s,km 38, 19 Feb
1968 (Y fl), Rodrigues et al. 8421 (F, INPA, US). MAR-
ANHAO: Mun. Monqho,Rio Turiaqu,nr. Urutuway,12 Feb
1985 (c), Balee et al. 809 (BG, MG, NY); Mun. Moncho,
Urutawy,Ka'aporIndianReserve,Rio Turiacu,28 Sep 1986
(Y fl-fr), Balee 2670 (NY); Sao Bento, 13 Nov 1923 (Y),
Snethlage 233 (F); Alto Alegria, 17 Nov 1923 (Y), Sneth-
lage 338 (F); Turiacu,2-3 Jan 1924 (6), Snethlage383 (F),
385 (F), 386 (F). MATO GROSSO:Nucleo Pioneirode Hum-
boldt, Rio Aripuana,21 Oct 1973 (Y fl), Berg et al. P.19817
(F, INPA, K, MO, NY, P, S, U), (6), Berg et al. P 19818 (F,
INPA, K, NY, U, US, S), 11 Oct 1973 (st), Berg s.n. (RB).
PARA: Sao Miguel do Guama,Oct 1906 (Y fl), Goeldi (MG)
7777 (BM, MG, G, P, S, U); Belem, 23 Aug 1906 (6),
Huber(MG) 7285 (BM, G); Tucurui,BR.230, km 50,5 Nov
1979 (? fl-fr), M. E F da Silva et al. 15 (INPA, MG) (6),
M. F E da Silva et al. 16 (INPA, MG).
BOLIVIA. BENI: Prov.Ballivian,Espiritu,RioYacuma,
9 Sep 1986 (6), Beck 5933 (LPB, MO), (Y fl-fr),Beck5934
(BG, LPB, MO); Prov.Yacuma,Rio Curiraba,4 Apr 1986
(9 fl-fr),M. MoraesR. 800 (BG, LPB, MO);Prov.Ballivian,
rd. Yucumo-Rurrenabeque,km 28, nr. Rio Hondo, 12 Mar
1989 (9 fl-fr), D. N. Smith et al. 12856 (BG, BOLV,LPB,
MO, USZ); Prov. Moxos, rd. San Borja-Trinidad,km 27, 6
Sep 1990 (6), D. N. Smith et al. 14336 (BG, MO, USZ);
Prov. Ballivian, nr. San Borja, 30 Sep 1976 (9 fl-fr), Ter-
ceros et al. 368 (BOLV,LPB, MO); Prov. Cercado,Ibiato,
7 Dec 1991 (st), Townsend6 (BG, LPB); Trinidad-Misiones
Guarayos,Sep 1926 (9 fl-fr), Werdermann2566 (K, LPB,
MO, S). COCHABAMBA:Nr. Chipiri, ca. 5 km N of Villa
Tunari,6 Dec 1955 (d), Berg et al. 1728 (BG, BOLV,COL,
LPB, USZ); Prov. Chapare,between Villa Tunari& Puerto
Villaroel,nr.Chimore,30 Dec 1982 (6), FerndndezC. 7913
(G, NY); Prov.Carrasco,Estaci6nValle de SajtaUMSS, 28
Oct 1991 (9 fl), Galarza et al. 2 (BOLV). LA PAZ: Prov.
Sud Yungas, nr. Sapecho, 26 Feb-2 Mar 1994 (6), Berg
1697 (BG, COL, LPB), (st, juv), Berg 1703A (BG), (st, juv),
Berg 1707A (BG), (6), Berg et al. 1708 (BG, LPB). PANDO:
Prov. Manupiri,rd. Altagracia-San Miquel-San Pedro, 23
Oct 1989 (9 fl), Beck et al. 19665 (BG, LPB); Prov.Nicolas
Suarez, nr. Porvenir, 10 Jan 1983 (6), FernandezC. et al.
8163 (G, NY); Prov. Nicolas Suarez, Campoana, 14 Jan
1983 (9 fl), Ferndndez C. et al. 8238 (G, NY). SANTA
CRUZ:Rio Pirai, Jan 1911 (9 fl-fr), Herzog 1508 (G, S);
Prov. Andres Ibafiz, 5.5 km NE of Cotoca, 1 km N of El
Campanero,28 Nov 1990 (9 fl-fr), Nee 40049 (BG, LPB,
MO, NY, USZ); JardinBotanico de Santa Cruz, rd. Santa
Cruz-Cotoca, km 12, 19 Dec 1991 (6), Nee 42151 (BG,
71
LPB, MO, NY); Prov. Ichilo, Rfo Ibabo, Reserva Forestal
Chore, 16-18 Aug 1990 (? fr), Neill et al. 9345 (MO, NY,
QCNE, USZ); Prov. 0. Santiestevan,15 km NNE of Mon-
tero, 9 Oct 1990 ( fl-fr), Saldias et al. 1236 (LPB, USZ);
Prov. Velasco, Parque Nacional Noel Kempff M., campa-
mento Los Fierros, 30 Oct 1993 (Y fl-fr), Saldias 3356
(USZ); Prov. Sara, Buena Vista, 3 Oct 1925 (d), Steinbach
7268 (A, BM, F, G, GH, K, MO, S, U), (Y fl-fr), Steinbach
7268bis (A, BM, E, F, G, K, MO, NY, S, U).
In subjuvenilematerial,the incisions of the lamina
are often not down to (near) the petiole, but up to 4
cm from it. This featureis occasionally retainedeven
at flowering state. This could be problematicin areas
where the ranges of Cecropiaconcolor and C. pachy-
stachya (could) overlap in the southernpart of the
Amazon basin or where the taxa may intergrade.To
be on the safe side, collections from Maranhaoand
Amazonaswith less deeply incised laminashave been
included in C. pachystachyain the presenttreatment.
The lamina segments are often ratherpronouncedly
lobate in (sub)juvenileplants. In this respectthe spe-
cies resembles C. metensis and C. pachystachya,but
it clearlydiffersfrom C. engleriana,in which lobation
of lamina segments in (sub)juvenilestates is less ap-
parent or common. The number of lateral veins in
juvenile materialcan be occasionally up to 30 pairs,
exceeding the maximumof 24 pairs of adultmaterial
and thus extendinginto the normalrangeof adultma-
terial of C. engleriana. The trees are small to
medium-sized (to ca. 15 m tall). In the middle and
lower Amazon basin, they may already start to pro-
duce flowers on trees of 1-1.5 m tall while still un-
branched.At the southernlimits of species range in
Bolivia (SantaCruz),the habitmay deviate.Here,the
trees are mostly quite robust with relatively thick
trunksand the leaves more crowded due to short in-
temodes. In AmazonianBrazil and adjacentAmazo-
nian Bolivia (Beni), the petiole and the uppersurface
of the lamina, the petiole, and the stipules are (ini-
tially) covered with ? dense arachnoidindumentum.
In other parts of Amazonian Bolivia (Cochabamba
and Santa Cruz) and southern Amazonian Peru
(Cuzco and Madre de Dios) the arachnoidindumen-
tum is sparseor lacking on these parts.The type with
sparsearachnoidindumentumon the petiole (etc.) has
often, or in some areasalways, subpersistentstipules.
In the conspicuous variationof the tree habit and the
regional presence of subpersistentstipules C. conco-
lor resembles C. engleriana. A collection made in
Colombia [Vichada:Mun. Puerto Carrenlo,Cerro El
Bita, aireadel JardinBotainico,30 Jan 1997 (2 fr), H.
Garcia et al. 60 (COL)], far outside the known range
of distribution,may have been introduced.
72
Vernacular names. Brazil: imbauiba branca
(Amazonas); ama'-y-puku (Ka'apor, Maranhao),
ama'-y-tuwir (Maranhao); imbautbabranca (Para).
Bolivia: ambaibonegro (SantaCruz).
9. Cecropia distachya Huber, Bol. Mus. Paraense
Hist. Nat. 6: 65. 1910; Berg, ActaAmaz6nica8(2):
173. 1978. Type. Brazil. Para: Santa Izabel, Oct
1906 (Y), Goeldi MG 7728 (holotype: MG; iso-
types: BM, G, L, P, S, U). Fig. 19
CecropiaripariaSnethlage,Notizbl.Bot.Gart.Berlin-
Dahlem8: 363. 1923.Type.Brazil.Acre:RioJurua,
Jurua-Mirim, Jun1901(d), Ule5587 (holotype:B,
destroyed;isotypes:G, K, LE,MG).
RevistaAcad.Colomb.
CecropiarichardiiCuatrecasas,
Ci. Exact.9(36/37):336. 1956. Type.Colombia.
Vaupes: RioPiraparand, Canio 4 Sep1952
Teemeenia,
(d), Schulteset al. 17191(holotype:GH;isotypes:
F, US).
Tree, to 30 m tall. Leafy twigs 1.5-6 cm thick,dark
green to red-brownto blackish,hispidulousto puber-
ulous to subhirtellous, partly with uncinate hairs.
Lamina (sub)coriaceous,ca. 20 X 20 cm to 85 X 85
cm, the segments (5-)7-10(-12), the free partsof up-
per segments obovate to subobovate, the incisions
down to 6/10-9/10; apices acute to acuminateor to
rounded;uppersurfacesmooth (to scabridulous),pu-
berulous (to hispidulous), initially often also sparse
with arachnoidindumentum;lower surface(minutely)
puberulous(with uncinatehairs) on the (main) veins,
sometimes also with sparse longer uncinate hairs,
with arachnoidindumentumin the areoles,sometimes
sparse (to nearly absent), sometimes (initially) also
on the main veins; lateralveins in the free partof the
midsegment 12-18(-25) pairs, usually marginallyor
sometimes submarginallyloop-connected, most (or
some) of them branched; petiole 15-60 cm long,
sparsely puberulous,sometimes (initially) also with
arachnoid indumentum, or subglabrous; trichilia
fused, the brown indumentumintermixedwith short
white (to brownishunicellular)hairs;stipules(6-) 10-
25 cm long, darkred-brownto brownor to brightred,
densely to sparselypuberulousor also with arachnoid
indumentumoutside, sparsely hairy to subglabrous
inside. Staminateinflorescencesin pairs,the peduncle
erect (or slightly deflexed), the spikes erect to +
spreading(or to pendulous);peduncle 2-12(-15) cm
long, often at the apex ? strongly broadened,
sparsely hispidulous to hirtellous; spathe (7-)10-18
cm long, dark red-brownto red, densely to sparsely
puberulousto hirtellousor also with sparse to rather
dense arachnoid indumentumoutside, glabrous in-
FLORANEOTROPICA
side; spikes (8-)15-25(-50), (0.5-)6-14 X 0.3-0.4 or
0.5-1 cm, ? angular,(sometimes green just before
anthesis), the stipes 0.3-1.5 cm long, puberulous,
sometimesonly with brownpluricellularhairs,or gla-
brous; rachis hairy or glabrous. Staminateflowers:
perianth tubular, ca. 1-1.5 mm long, with sparse
straighthairs and/orsparseto ratherdense arachnoid
indumentumbelow the apex, the apex slightly convex
to plane,muriculateor (also) with sparsebrownpluri-
cellular hairs;filamentsswollen; anthers0.6-0.7 mm
long, detachedat anthesis(?). Pistillate inflorescences
in pairs or solitary,erect or deflexed to pendulousin
fruit;peduncle (2-)5-13 cm long, hispidulousto hir-
tellous; spathe darkred-brownto red, 6-15 cm long,
densely to sparsely puberulousto hirtellous or also
with arachnoidindumentumoutside, glabrousinside;
spikes 2-4, (3-)8-15 X 0.6-0.8 or ca. 2.5 cm, to 22
X 1.5(-2) cm or to 38 X 4 cm in fruit,sessile or with
stipes to 0.5 cm long and subglabrous;rachisglabrous
or sparselyhairy.Pistillateflowers: perianthca. 1.5-
3.5 mm long, with arachnoidindumentumbelow the
apex, also in the lower part of style channel inside,
the apex plane with (a rim around)a circularaperture
or stronglyconvex with a slit-shapedaperture,muri-
culate to smooth; style long; stigma comose. Fruit
ellipsoid to oblongoid, 2.5-4.5 mm long, tuberculate
or smooth.
Distribution (Fig. 10.3). Throughoutthe Amazon
basin, extending to eastermVenezuela,and to French
Guiana;common in primaryforest, also in secondary
growth,in non-inundated(or inundated)places, at low
elevations.
Representative specimens examined. COLOMBIA.
15
17kmfromLaChorrera,
AMAZONAS:RioIgara-Parana,
Jul 1974 (st), Gasche et al. 222 (COL); Mun. Tarapaca,
Parque Nacional Amacayacu, Rio Catuh6, Lorena, 13 Jul
1992 (st), Rudas et al. 5268 (COL). CAQUETA: Nr. Arara-
cuara, 12 Jul 1986 (Y fl), Berg et al. 1553 (BG), 29 Feb
1992(Y fl-fr),Correaet al. 384 (COL). GUAVIARE:Sierra
de Chiribiquete,CampamentoNorte, 8 Dec 1990 (Y fl-fr),
Galeano et al. 2204 (BG, COL, MO); Mun. San Jose de
Guaviare,La Libertad,11 Aug 1989 (cd), Marulandaet al.
1204 (HUA); Sierrade Chiribiquete,26 Aug 1992 (st), Pa-
lacios et al. 2655 (COL).META: Nr.La Macarena,Rio Mor-
rocoy, 11 Feb 1995 (Y fl-fr), Francoet al. 4667 (BG, COL);
Alto de Bengala, Dec 1993 (? fl), Stevenson 779 (COL).
VAUPES: Type collection of C. richardii.
VENEZUELA. AMAZONAS: Depto. Rio Negro, Rio
Mawarinuma,Neblina Base Camp,2 Dec 1984 (Y fl), Boom
et al. 5252 (BG, VEN); Depto. Atabapo,Cucuritalde Can-
ame, Middle Caflo Caname, 30 Apr-I May 1979 (9 fl-fr),
Davidse et al. 16969 (MO, U, VEN); Depto. Atabapo,Rio
Ocamo, Santa Maria de Los Guaicas, 25 Feb 1981 (9 fr),
Gua'nchez786 (VEN); Hauchica, 11 km NE of San Carlos
SYSTEMATICTREATMENT 73

o 6] 1 19

199'3'~~~~~~~~~~~~~

FIG. 19. Cecropia distachya. 1. Lamina, reduced. 2. Apex of lamina and venation (Berg et al. 1626). 3. Stipules and
trichilium (Berg et al. 1627). 4. Staminate inflorescence with spathe and base of petiole with trichilium. 5. Staminate
inflorescence at anthesis and base of petiole with trichilium (Berg et al. 1626). 6. Staminate flower and stamen (Berg et al.
1597). 7. Pistillate inflorescence with spathe and trichilium. 8. Pistillate inflorescence at fruit and trichilium. 9. Pistillate
flower. 10. Fruitwith remnantof style (Berg et al. 1627).
74
de Rio Negro, 14 Nov 1977 (9 fl-fr), Liesner 3481 (MO,
NY, U, VEN); Rfo Caura,CanloLa Ceiba (CanloCumaca),
9-26 May 1988 (5), Stergios et al. 12888 (BG, NY); Rio
Paragua,Raudales de Maihia, 1 Jan 1962 (st), Steyermark
90526 (NY, US); San Carlos de Rfo Negro, 20 km S of
confluence of Rio Negro & Brazo Casiquiare,10 Nov 1979
(5), Uhl 51 (MO). BOLiVAR: Rio Tirica, 29 Aug 1954 (9
fl-fr),Bernardi1612 (NY); Rio Parguaza,just below Raudal
Maraca,ca. 110 km above river mouth, 29 Dec 1955 (9 fl-
fr) Wurdacket al. 40997 (BG, MO, NY).
FRENCH GUIANA. Regina region, Mt. Tortue,11 km
WNW of ApprouagueR., 18 Jun 1988 (5), Feuillet 10312
(BG); Basin de l'Approuague,rd. Regina-St.-Georges, 24
Nov 1995 (9 fl-fr), Granvilleet al. 13104 (BG, G); Bassin
de l'Oyapock, Cr. Gabaret, 16 Sep 1996 (9 fl-fr), Tostain
263, 270 (BG).
ECUADOR. NAPO: ParqueNacional Yasunf,nr. Heli-
puertode Amo Sur, 16-19 Jan 1988 (9 fl), Ceron3408 (BG,
QAME, QCNE). SUCUMBfOS: Cant6n Shushufindi, San
Roque, 14 Dec 1997 (st), E. Freire et al. 2784 (QCNE);
Reserva FaunisticaCuyabeno, nr. Laguna Grande, 19 Nov
1991 (5), Palacios et al. 9312 (BG, QCNE), 15 Mar 1990
(9 fr), Valencia et al. 375 (BG, QCA) & 67967 (AAU).
ZAMORA-CHINCHIPE:Cant6nNangaritza,Rio Nangaritza,
Miazi, 29 Jul 1993 (st), Gentry80664 (QCNE).
PERU. AMAZONAS:Rio Cenepa, SE of QuebradaKay-
amas, 28 Dec 1972 (9 fr), Berlin 736 (GH); Rio Santiago,
E of Caterpiza,ca. 65 km N of Pinglo, 7 Nov 1979 (9 fl-
fr), Huashikat 1189 (BG, MO), 17 Jan 1980 (9 fr), Hu-
ashikat 1828 (MO);Prov.Bagua,Dtto. Imaza,Rio Marafi6n,
Yamayakat,Feb 1995 (st), R. Vdsquezet al. 19938 & 19973
(MO); Prov.Bagua, Cascadasde Mayasito,5 Sep 1962 (d),
Wurdack1849 (F, NY, US, USM). Cuzco: Prov. Paucar-
tambo, rd. Pilcopata-Paucartambo,km 12, 2 Aug 1988 (5),
Berg et al. 1626 (BG, COL, Centro de Medicina Andina,
Cuzco), (9 fl-fr), Berg et al. 1627 (BG, COL, Centro de
MedicinaAndina,Cuzco); Prov.Paucartambo,nr.Fortaleza-
Patria, 18 Jan 1975 (9 fl-fr), Galeano S. 9 (US, USM).
HUANUCO: Prov.Pachitea,20-24 km SE of PuertoInca, 11
Sep 1988 (9 fr), Morawetzet al. 11-11988 (BG). LORETO:
Nr. Iquitos, 8 Nov 1940 (9 fl-fr),Asplund14413 (S); Prov.
Requena,rd. JenaroHerrera-Angamos,km 11, 18 Jul 1988
(5), Berg et al. 1597 (AAU, BG, COL), (9 fl), Berg et al.
1598 (AAU, BG, COL); Prov. Loreto, CampamentoPetro-
lero San Jacinto,Rio Tigre, 6 Sep 1979 (5), Diaz etal. 1470
(BG, MO); Prov. Maynas, nr. Sucusari,20 Feb 1991 (9 fl-
fr), Pipoly et al. 13115 (BG, MO), (st), Pipoly et al. 13141
(BG, MO, NY); Prov. Requena,JenaroHerrera,6 Sep 1983
(5), Spichigeret al. 3050 (G, NY), 7 Sep 1983 (9 fl), Spi-
chiger 3053 (G); Prov. Maynas, Allpahuayo,Nov 1990 (9
fl), R. Vasquez et al. 14725 (BG, MO, NY). MADRE DE
Dios: Tambopata,23 Feb 1984 (5), Gentry et al. 46069
(MO); TambopataNature Reserve, jct. of Rio La Torre &
Rio Tambopata,31 May 1987 (d), Gentry et al. 58001
(MO). SANMARTiN: Prov. San Martin,rd. Tarapoto-Yuri-
maguas, ca. km 35, 5 Dec 1997 (9 fl-fr), Berg et al. 1781
(BG, COL, MOL).
BRAZIL. ACRE:ReservaINCRA SantaLuzia,BR.364,
km 40, 5-19 Oct 1984 (9 fr), Campbellet al. 6806 (BG);
FLORA NEOTROPICA
Cruzeirodo Sul, EstradaAlemanha,7 May 1971 (6), Maas
et al. P12771 (INPA, K, MG, MO, NY, U, S, US). AMA-
ZONAS: Rd. Manaus-Caracari,km 65, 21 Sep 1973 (6),
Berg 282 (U); Rio Jurua,Carauari,32 km from P6rto Gav-
iao, 15-26 Oct 1980 (st), Lisboa et al. 1897 (MG); Mun.
Humaita,betweenRio Livramento& Rio Ipixuna,7-18 Nov
1934 (9 fl-fr), Krukoff7214 (A, BM, F, G, LE, K, MICH,
MO, NY, RB, S, U); Mun. Sao Paulo de Olivenqa,Belem
Cr., 26 Oct-i l Dec 1936 (9 fl-fr), Krukoff8921 (A, BM, F,
G, LE, K, MO, NY, S, U); Rio Negro, CommunidadeApa-
recida, opposite mouth of IgarapeTuari, 6 Nov 1987 (6),
Maas et al. 6935 (BG, INPA); Rio Cuieras, 2 km below
mouth of Rio Brancinho, 12 Sep 1973 (9 fl), Prance et al.
17827 (C, COL, F, INPA, K, MG, MICH, MO, NY, S, U,
US), (9 fl-fr), Prance et al. 17840 (COL, F, INPA, K, MO,
NY, P, S, U, US); Rio Cuieras, Jarada,16 Sep 1973 (d),
Prance et al. 18008 (C, COL, F, INPA, K, MICH,MG, MO,
NY, P, S, TEX, U); Mun. Manaus,rd. Manaus-Itacoatiara,
km 26, Reserva FlorestalDucke, 15 Sep 1994 (d), Ribeiro
et al. 1424A (BG, INPA).MATO GROSSO:Nucleo Pioneiro
de Humboldt,Rio Aripuana,22 Oct 1973 (9 fr), Berg et al.
P 19839 (COL, F, INPA, K, NY, U, US); rd. Vilhena-Jufna,
km 134, EstaqaoEcol6gica Ique-Juruena,24 Aug 1982 (d
fr), F A. M. dos Santoset al. 14228 (RB, UEC). PARA:Serra
do Cachimbo, nr. CachoeiraCurua, rd. Cuiaba-Santarem,
ca. km 875, 30 Oct 1977 (9 fl-fr), Berg et al. 773 (COL, F,
K, MG, MO, NY, U, US).
BOLIVIA. LA PAZ: Prov. Abel Iturralde,PuertoMos-
coso, Rio Heath, 25 Jul 1995 (9 fl-fr), Helme et al. 898
(LPB). SANTA CRUZ: Prov.Velasco, ParqueNacional Noel
KempffM., 13 Nov 1993 (9 fl-fr),Arroyoet al. 326 (USZ);
Prov.Velasco, ParqueNacional Noel KempffM., Meseta de
Huanchaca,18 Aug 1996 (9 fl-fr), Carri6net al. 133 (BG);
Prov. Velasco, Parque Nacional Noel Kempff M., campa-
mento HuanchacaI, 26 May 1994 (d), H. Gonzdles et al.
42 (BG); Prov. Guayaros,Rio Negro, 13 Jul 1993 (9 fl-fr),
I. G. Vargaset al. 2695 (USZ).
In Peru(Cuzco, Huainuco,Junin,and San Martin),
the spikes of the pistillate inflorescencesreach to 38
X 4 cm in fruit and the spikes of the staminateinflo-
rescences are also thicker than usually found else-
where. In otherpartsof the species' range,the spikes
are andremainmuch shorter.Two of these collections
with large spikes match the materialof Cecropiadi-
stachya in most features,but are clearly distinctin the
shape of the lamina. Moreover, they have unusual
arachnoidindumentumon the lamina above. They
may represent a distinct type of C. distachya (de-
scribed below) in which some (sub)juveniletraitsare
retainedor they may representa hybrid (with many
maternaland few paternalfeatures;see "Natureand
Variationof MorphologicalCharacters").
Lamina coriaceous, incisions in the upper part
down to ca. 5/10, the lower partslightly lobed; upper
surface with sparse to ratherdense arachnoidindu-
mentum;lateralveins in the free part of the midseg-
SYSTEMATICTREATMENT
ment 9-12 pairs, marginallyto submarginallyloop-
connected.Pistillate inflorescenceswith the peduncle
2-5.5 cm long, the upper part broadened in fruit;
spathe ca. 8 cm long; spikes 2, 5-6 (or more?) X 1-
1.3 cm, to 32 X 2 cm or to 35 X 3.5 cm in fruit.
Pistillateflowers: perianthca. 2.5 mm long. Fruitob-
longoid, 4-5 mm long.
Description based on the following collections:
PERU. HUANUCO: Prov. Leoncio Prado,Dtto. Rupa
Rupa, E of Tingo Maria, nr. Cerro Quemada,7 Sep
1978 (? fr), SchunkeV 10579 (BG, US). JUNIN: Pi-
chanaki, between Satipo and bridge over Rio Chan-
chamayo,900-1000 m, 28 Jun 1982 (9 fl-fr), Gentry
et al. 37251 (MO).
Collections with features transitionalto the more
typical type occur, as with regardto the inflorescence
(Berg et al. 1627) or the leaf shape (Morawetzet al.
11-11988).
The spikes of the staminateinflorescencescan be
green before anthesis(as in Berg et al. 1597). In most
specimens the apex of the perianth of the pistillate
flower is plane with a rim aroundthe roundaperture
and ? angularin circumference.In some collections
(e.g., Bernardi 1612 and Liesner 3481) the apex of
the perianthof the pistillate flower is (strongly)con-
vex, circularin outline, and with a slit-shapedaper-
ture. Some collections have up to 12 laminasegments
and/or submarginallyloop-connected lateral veins;
these featurescan be relatedto the (sub)juvenilestate.
Vernacular names. Colombia: juimekokai(Wi-
toto, Amazonas);jucoqui (Hui., Caqueta);yarumone-
gro (Guaviare).Venezuela:yagrumo morado (Ama-
zonas); kam'a-in-yek(Bolivar). Peru: toroc (Cuzco);
setico negro (Huanuco);shiari (Loreto). Brazil: im-
baubavermelha(Amazonas).
10. Cecropia elongata Rusby, Bull. New York Bot.
Gard.4: 446. 1907. Type.Bolivia. Withoutlocality
(probablyProv.Nor Yungas,Coripata),(6d), Bang
2260 (holotype:NY; isotypes: BM, G, GH, K, LE,
MO, US, WIS).
Tree,to 15 m tall. Leafy twigs 2-9(-15) cm thick,
purplish brown, puberulous to hispidulous, with
straighthairs and also dense brown pluricellulartri-
chomes. Laminachartaceousto subcoriaceous,ca. 30
x 30 cm to 90 x 90 cm (to 105 X 105 cm), the
segments (8-)10-16, oblanceolate, the incisions
down to 7/10-9/10; apices subacuteto acuminate;up-
per surface ? scabrous, hispidulous; lower surface
puberulous to subtomentellousto subtomentose on
the smaller veins to almost glabrous on the main
75
veins, with arachnoidindumentumin the areoles and
on the smaller veins; lateral veins in the free partof
the midsegment (18-)20-25(-40) pairs, submargin-
ally (in the lower part of the segment faintly) loop-
connected,unbranched;petiole 20-80(-105) cm long,
sparsely puberulous to hispidulous to subglabrous;
trichilia fused, the brown indumentum intermixed
with short white hairs; stipules ca. (10-)20-40 cm
long, red-brownto greenish, sparsely strigillose and
with dense brownpluricellulartrichomesor also with
arachnoidindumentumoutside, (sparsely) sericeous
inside. Staminateinflorescencesin pairs,the peduncle
erect, the spikes ? spreadingto pendulous;peduncle
ca. 7-25 cm long, hirtellousto subhirsute;spathe at
least 15 cm long, color unknown,densely brownish
to whitish subhirtellousto subsericeous outside, ?
sparselyhairyinside; spikes 13-25, 18-27 X ca. 0.4-
0.6 cm, the stipes 0.4-1 cm and glabrousor sparsely
hairy; rachis sparsely hairy. Staminateflowers: peri-
anth tubular, 1.5-2 mm long, with sparse to dense
shortarachnoidindumentumbelow the apex, the apex
slightly convex to plane, punctateto smooth, or ini-
tially also with brown pluricellulartrichomes; fila-
ments flat; anthersca. 0.6-0.8 mm long, without ap-
pendages, remaining attached to the filaments at
anthesis. Pistillate inflorescencesin pairs or solitary,
erect to spreadingto subpendulousin fruit;peduncle
(10-)20-25 cm long, sparsely to ratherdensely hir-
tellous to subhirsute;spathe at least 20 cm long, the
color unknown,the indumentumas in the staminate
inflorescenceor more densely hairy inside; spikes 4-
9, ca. 25-40 X 1 cm, to 47 X 1.5 cm in fruit, the
stipes 0.3-1.5 cm long and glabrous;rachisglabrous.
Pistillate flowers: perianthca. 1.5-2 mm long, with
arachnoidindumentumbelow the apex outside, ab-
sent inside, the apex convex, punctateto muriculate,
the apertureslit-shaped;style long, straight;stigma
comose. Fruitellipsoid to oblongoid, 2-2.5 mm long,
tuberculate,brown.
Distribution (Fig. 18.1). Bolivia (Cochabamba,
La Paz, and SantaCruz),in montaneforest, at 1600-
2400 m.
Specimens examined. BOLIVIA. COCHABAMBA:Rd.
Coachabamba-Villa Tunari,ca.km84, 2400m, 6 Dec 1995
( fl-fr),Berget al. 1722(BG,BOLV,COL,LPB),ca. km
102, 2000 m, 6 Dec 1995 (d), Berg et al. 1726 (BG, BOLV,
COL, LPB, USZ). LA PAZ: Prov. Nor Yungas,rd. Yolosa-
Chuspipata,2000, 11 Mar 1987 (6), Beck13549 (BG, LPB,
MO); Prov. Sud Yungas, rd. Huancane-San Isidro, km 7,
2250, 11 Feb 1991 (st), Beck19723 (BG); Chulumani,1800
m, Aug 1933 (6), M. Cdrdenas1165(NY); Prov.Inquisivi,
Khora-Mikilpirhua,24 Km N of Choquetanga,1900 m, 14
Jul 1994 (6), Salinas 3145 (BG); Prov.SudYungas,2.5 km
76
NW of Yanacachi,2120 m, 30 May 1987 (6), Seidel 994
(LPB); Prov. Nor Yungas, 13.5 km NW of Chuspipata,rd.
to Yolosa, 2100 m, 5 Oct 1984 ( Y fr), Solomonet al. 12459
(BG, LPB, MO); valley of Rfo Zongo, 1650 m, 29 Apr 1981
(9 fl-fr), VdzquezAvila 354 (LPB, RB), 18 Nov 1981 (9
fl-fr),Zuloagaet al. 1833(LPB).SANTA CRUZ: Prov.Flor-
ida, 5 km E of SantaRosa de Lima, 1470 m, 3 Jul 1996 (6),
Saldias et al. 4363 (NY); Prov. Caballero,K'arawasi, 1900
m, 20 Jul 1996 (st), Saldfas et al. 4442 (USZ).
This species certainlydoes not belong to the group
of montanespecies (includingthe white-leavedones)
found more northwardin the Andes. It might be re-
lated to Cecropia angustifolia. The stems and
branches are often conspicuously broadenedtoward
the apices. The apex of the terminalbud cover (the
stipules) is often torn off by elongation of the corre-
sponding leaf. Flowering (and fruiting) might be
largely seasonal. Populations examined in Bolivia
(Cochabamba)in December(1995) had only very few
trees with staminateinflorescences and the pistillate
inflorescences were largely in the same state of de-
velopment.The trees are inhabitedby ants.
11. Cecropia engleriana Snethlage, Notizbl. Bot.
Gart. Berlin-Dahlem 8: 365. 1923; Berg, Acta
Amazonica 8(2): 166. 1978. Cecropia concolor
Willdenow subsp. engleriana (Snethlage) C. C.
Berg & P. Franco,Fl. Ecuador48: 17. 1993. Type.
Brazil. Acre: Rio Acre, Seringal Sao Francisco,
Jun 1911, Ule 9313 (holotype: B (with staminate
inflorescences), destroyed;isotypes: G, K; under
the same numberin MG a specimen with pistillate
inflorescences). Fig. 17.3-10
Cecropiapacis Cuatrecasas,Revista Acad. Colomb. Ci.
Exact. 6(22/23): 288. 1945. Type. Colombia. Putu-
mayo: Rio Putumayo,between mouth of Rio Gua-
mues & Puerto Asis, Cuatrecasas11244 (holotype:
COL; isotype: F).
Cecropia yarinensis Cuatrecasas,Revista Acad. Col-
omb. Ci. Exact. 9(36/37): 339. 1956. Type. Peru.
Ucayali: Prov.CoronelPortillo,Pucallpa,4 km from
Yarina,rd. to Huanuco, 8 Apr 1953 (9), Ferreyra
9044 (holotype: US; isotype: UMS).
Tree,to 20(-30) m tall; trunkwith prominentstip-
ular scars. Leafy twigs 1.5-4 cm thick, green with
yellow spots to reddish to purplishbrown, hispidu-
lous, with curvedor uncinatehairs,or also with sparse
to ratherdense arachnoidindumentum.Lamina sub-
coriaceous to chartaceous,ca. 30 X 30 cm to 65 X
65 cm, the segments (7-)8-10(-1 1), the free partsof
the upper segments oblong to subobovate to oblan-
ceolate, entire or ? sinuate (to lobate), the incisions
FLORANEOTROPICA
down to (6/1-)8/10-9/10 (2-4 cm, or sometimes 8
cm from the petiole); apices short-acuminateto
(sub)acuteto obtuse;uppersurfacesmooth to scabri-
dulous, sparsely puberulousto subhispidulous,also
with sparse arachnoidindumentum,along the main
veins often with brownpluricellularhairs;lower sur-
face minutely puberulous on the main veins, with
arachnoidindumentumin the areoles, on the smaller
veins and/orat least initially also on the main veins;
lateralveins in the free partof the midsegment(21-)
24-32 pairs, marginally loop-connected, mostly
branched;petiole 25-50 cm long, sparselypuberulous
and/or also with sparse to (rather)dense arachnoid
indumentum;trichilia fused, the brown indumentum
intermixedwith (sparse)shortwhite hairs;stipules 8-
25 cm long, white to pale yellow or pinkish,caducous
or occasionally subpersistent,puberulousor sparsely
sericeous, often with ? dense arachnoidindumen-
tum, sometimes (also) with ? dense brownpluricel-
lular hairs outside, ? densely sericeous inside. Sta-
minate inflorescencesin pairs or solitary,pendulous;
peduncle (3.5-)10-19 cm long, sparsely puberulous
to hirtellousand with sparseto ratherdense arachnoid
indumentum;spathe 10-20 cm long, white to yellow-
ish or pinkisk,puberulousto hirtellousand often with
dense arachnoid indumentum outside, glabrous or
(densely) sericeous inside; spikes (8-)15-20, (1-) 10-
20 X 0.2-0.4 cm, the stipes 0.2-2 cm long, hairy or
subglabrous;rachishairy.Staminateflowers:perianth
tubular,0.8-1.5 mm long, glabrous,the apex slightly
convex; filamentsflat, anthers0.5-0.6 mm long, ap-
pendiculate, detached at anthesis, reattachedto the
marginsof the apertureby the appendages.Pistillate
inflorescencesin pairs or solitary,pendulous;pedun-
cle (3.5-)6-22 cm long, puberulousto hirtellousand
with sparse to ratherdense arachnoidindumentum;
spathe 10-17 cm long, the color and indumentumas
in the staminateinflorescence;spikes 4, (2-)5-18 X
0.4-1 cm, to 37 X 1-1.5 cm in fruit, sessile or with
stipes to 1(-1.5) cm long andhairy.Pistillateflowers:
perianthca. 2 mm long, with arachnoidindumentum
on andbelow the apex outside,absentinside, the apex
plane to slightly concave or slightly convex; style
short;stigma peltate (to subpelateor tongue-shaped).
Fruitellipsoid to oblongoidto ovoid, 1.5-2.5(-3) mm
long, ? tuberculateor smooth.
Distribution (Fig. 10.4). Upper Amazon basin,
from Colombiato Bolivia, often, at least in the north-
ern partof its range,associatedwith riverinehabitats,
at low elevations.
Representativespecimensexamined.COLOMBIA.
AMAZONAS: Mun. Letica, ParqueNacional Natural Ama-
SYSTEMATICTREATMENT
cayacu, Quebradade Agua Pudre, 19 Nov 1991 (st), Pipoly
et al. 16410 (COL, MO), 22 Mar 1992 (9 fr), Rudas et al.
3489 (COL, MO); Rio Caraparana,nr. El Encanto, 22-28
May 1942 (9 fl-fr), Schultes 3821 (COL, ECON, F); Dtto.
Puerto Narifno,Parque Nacional Amacayacu, 9 Oct 1989
(6), R. Vdsquezet al. 12685 (BG, COL, MO). CAQUETA:
Mun. Florencia,rd. to Gabinete,km 1, 18 Oct 1993 (9 fl-
fr), Franco et al. 4516 (BG), (6), Franco et al. 4517 (BG).
CAUCA: Rd. Mocoa-Pitalito, nr. Rio Caqueta,18 Feb 1995
(9 fl-fr), Franco et al. 4690 (COL). PUTUMAYO: Between
Mocoa & Urcusique,9 Jan 1945 (9 fl-fr), Ewan 16732 (F,
US); Mun. Mocoa, rd. Mocoa-Villa Garz6n,El Pepino, 18
Feb 1995 (d), Franco et al. 4687 (COL, HUA, LP,MO, NY,
US), (6), Franco et al. 4688 (COL, HUA); rd. Mocoa-
PuertoAsis, 3 May 1994 (9 fl-fr),Francoet al. 5517 (COL);
rd. Garz6n-PuertoAsis, 3 May 1994 (6), Francoet al. 5520
(BG, COL).
VENEZUELA. AMAZONAS: Depto. Atures, Serrania
Batata, 2 km NE of Salto Colorado, 55 km SE of Puerto
Ayacucho,Oct 1989 (9), Sanoja et al. 3329 (BG); Rio Ca-
siquiare,nr.Rio Orinoco, Jan-Feb 1969 (9 fl-fr), Velasquez
et al. 501 (US) = Fariniaset al. 501 (VEN), (d), Velasquez
et al. 506 (US) = Farifnaset al. 506 (VEN); Upper Rio
Orinoco, La Esmeralda, 19 May 1942 (9 fr), LI. Williams
15516 (US).
ECUADOR. MORONA-SANTIAGO: Centro Shuar Yu-
kutis, 22 Apr 1989 (st), Bennett et al. 3801 (QCNE); rd.
Zamora-Gualaquiza,Estaci6nExp. El Padmi,ca. 5 km N of
Los Encuentros, 4 Jan 1991 (6), Berg et al. 1657 (BG,
LOJA,QCA). NAPO:Nr. Tena,7 Jan 1940 (9 fl-fr),Asplund
10290 (S); Rio Napo, Descanso, 25 km from Coca, 1 Mar
1980 (9 fl-fr), Berg et al. 1104 (AAU, BG, COL, GB, MO,
QCA, U); ParqueNacionalYasuni,Pozo petroleroDaimi 2,
26 May-8 Jun 1988 (9 fr), Cer6n et al. 3847 (BG, MO,
QCNE); Afiangu, ParqueNacional Yasuni, 1-15 Feb 1986
(st), Korninget al. 47692 (AAU, BG, QCA); ReservaBiol-
6gica JatunSacha, 8 km E of Misahualli,25 Dec 1990 (6),
Neill et al. 9742 (BG, QCA, QCNE);nr. ReservaBiol6gica
JatunSacha, rd. Puerto Napo-Ahuano, 25 Dec 1990 (d),
Neill et al. 9744 (BG, QCA, QCNE). SUCUMBiOS: Rio
Aguarico, ca. 0?0', 76'40-45'W, 22 Feb 1980 (d), Berg et
al. 1075 (AAU, BG, COL, GB, K, MO, NY, QCA, TUR, U,
WIS); 17 km W of Lumbaqui,4 Nov 1974 (9 fl-fr), Gentry
12442 (MO, QCA, U).
PERU. AMAZONAS: QuebradaCikan-inci,W of Huam-
pami, 18 Dec 1972 (9 fl-fr), Berlin 592 (AAU, MO, NY,
USM); Prov.Bagua, Dtto. Imaza,Rio Marafi6n,Yamayakat,
6 Mar 1966 (9 fl-fr), N. Jaramillo et al. 579 (MO); Rio
Cenepa, QuebradaHuampami,29 May 1973 (9 fr), Kayap
832 (BG, F, NY); QuebradaSatik-entsa, 16 Jul 1974 (d),
Kayap 1099 (MO, U, USM); Rio Cenepa, nr. Huampami,
ca. 5 km E of ChavezValdivia,3 Aug 1978 (9 fl-fr),Kujikat
132 (BG, MO). Cuzco: Prov. Paucartambo,rd. Pilcopata-
Paucartambo,km 16, 2 Aug 1988 (9 fl-fr), Berg et al. 1629
(BG, COL,Centrode MedicinaAndina,Cuzco). HUANUCO:
Prov.Pachitea,ca. 26 km S of PuertoInca, 27-28 Nov 1988
(st), Johann 2/26 (BG), 2/35 (BG); Prov. PuertoInca, Dtto.
Yupapichis,DANTAS, 16-31 Mar 1991 (d), Tello1583 (G,
NY), 1-15 Apr 1991 (9 fl-fr), Tello 1731 (G, NY); Shapa-
77
jilla, 28 May 1939 (6), Woytkowski23 (F). LORETO:Prov.
Maynas, Dtto. Indiana,ExploramaInn, 21 Apr 1992 (Y fl-
fr), Grdndezet al. 4091 (MO);JeneroHerrera,25 Nov 1987
(Y fl), Lamotte325 (BG), (6), Lamotte326 (BG); Rfo Am-
piyacu, nr. Pebas, 24 Apr 1977 (Y fl), Plowman 7020
(ECON, F, US); mouthof Rio Santiago, s.d. (Y fl-fr), Tess-
mann4058 (G, NY, US); Iquitos,s.d. (Y fr), Tessmann5044
(NY, US); Prov.Alta Amazonas,Andoas, 20 Nov 1980 (6),
R. Va'squezet al. 776 (NY); Prov. Maynas, Pto. Alianza,
(Qda. Tamshiyacu),29 May 1981 (Y fl-fr), R. Vdsquezet
al. 1856 (BG, MO). MADRE DE DIos: Prov. Tambopata,
ComunidadeNativa Infierno,HermosaChica, 14 Feb 1989
(6), Alexiades et al. 290 (BG); Prov. Tambopata,30 km
SSW of Puerto Maldonado,confluence of Rio La Torre&
Rio Tambopata,26 Jun 1980 (Y fr), Barbour 5820 (BG,
MO); Prov. Manu, nr. Shintuya,30 Jul 1988 (Y fl-fr), Berg
et al. 1610 (BG, COL, Centrode MedicinaAndina,Cuzco);
Prov.Manu,ManuPark,Cocha Cashu, 8 Sep 1986 (st), Niu-
niez6044 (BG), Aug-Sep 1989 (st), Nuliiezet al. 11276 (MO).
SAN MARTiN: Prov. Lamas, rd. Lamas-Pamashto,4 Dec
1997 (Y fl-fr),Berg et al. 1765 (BG, COL,MOL), (d), Berg
et al. 1766 (BG, COL, MOL); Prov.Moyobamba,rd. Moy-
obamba-Rioja,ca. km 10, 8 Dec 1997 (Y fl-fr), Berg et al.
1792 (BG, COL, MOL);rd. Rioja-NuevaCajamarca,ca. km
5, 8 Dec 1997 (6), Berg et al. 1793 (BG, COL, MOL); rd.
Tarapoto-Juanjui,7 Oct 1984 (Y fl-fr), Maas et al. 5983
(U). UCAYALI: Nueva Requena, 11 km from CampoVerde
(km 30 on rd. Pucallpa-Lima),4 May 1988 (Y fl-fr),Arana
7 (K), 11 Aug 1988 (6), Berg et al. 1636 (BG, COL, K);
Prov. Coronel Portillo, nr. Pucallpa,4 km from Yarina,rd.
to Huanuco, 8 Apr 1953 (6), Ferreyra9045 (USM); Prov.
Coronel Portillo, nr. Pucallpa, Yarinacocha,19 Dec 1968
(d), Ferreyra17629 (US, USM).
BRAZIL. ACRE:Mun. MarechalTaumaturgo,Rio Ju-
rua, Fazenda Paraguay,3 Apr 1993 (6), Daly et al. 7726
(NY); Mun. Madureira,Rio Macaua,Col6nia Bom Futuro,
30 Mar 1994 (Y fl-fr), Lima et al. 529 (NY). AMAZONAS:
Mun. Sao Paulo de Oliven,a, nr. Palmares, 11 Sep-26 Oct
1936 (6), Krukoff8061 (A, BM, F, G, K, LE, MICH, MO,
NY, P, S, U, US), (Y fl), Krukoff8336 (A, BM, F, G, K, LE,
MO, NY, S, U). ROND6NIA: ParqueIndigenaAripuana,Jan
1982 (Y fl-fr), Coimbra 71B (MG); Estradade Ferro Ma-
deira, Mamore, 10 Sep 1923 (6), Kuhlmann407 (RB).
BOLIVIA. BENI: Prov. Ballividn, Misi6n Fatima, San
Borja, Rio Maniqui, 28 May 1988 (6), Beck et al. 16662
(BG, LPB, MO); Prov.Moxos, 130 km S of San Ignacio, 10
Sep 1991 (st), Del Aguila et al. 90 (LPB, MO); Prov. Bal-
livian, Triunfo,27 Jul 1976 (Y fl), Meneces et al. 45 (BOLV,
MO); Prov.Ballividn,rd.Yucumo-Rurrenabaque, 20-25 Jun
1989 (9 fr), D. N. Smith et al. 13564 (BG, BOLV, LPB,
MO, USZ); Prov.Ballivian, rd.Yucumo-Rurrenabaque, km
34, 7-14 Jul 1990 (6), D. N. Smithet al. 14126 (BG). Co-
CHABAMBA:Rd. Santa Cruz-Villa Tunari,km 240, 12-14
Jul 1989 (9 fr), D. N. Smith et al. 13713 (BG, LPB, MO,
USZ). LA PAZ: Prov. Iturralde,Luisita, 22 Feb 1984 (9 fl-
fr), Beck et al. 9931 (BG, LPB, MO, USZ); Prov. SudYun-
gas, rd. Sapecho-Yucumo,km 1.5, 28 Feb 1994 (9 fl-fr),
Berg 1700 (AAU, BG, COL, LPB, NY); Prov. Sud Yungas,
nr. Sapecho, 3 Mar 1994 (6), Berg 1710 (BG, COL, LPB,
78 FLORANEOTROPICA

NY); Prov. Sud Yungas, Serraniade Marimonos, Colonia Cecropia porvenirensis Cuatrecasas,Bot. Mus. Leafl.
Tarapaca',23 Jul 1993 (Y fl-fr),Seidel et al. 7316 (BG, LPB). HarvardUniv. 14: 25. 1949. Type. Colombia. Putu-
PANDO:54 km SW of Cobija, Triunfo, 23 Jul 1980 (Y fl- mayo: Puerto Porvenir,nr. Puerto Ospina, 20 May
fr), Penningtonet al. 31 (BG, F, K, LPB). SANTA CRUZ: 1942, Schultes 3697 (holotype: F, consisting of
Prov. Ichilo, ReservaForestalChore, Rio Itabo, 16-18 Aug leaves of C. ficifolia, here designated as the lecto-
1990 (Y fl-fr), Neill et al. 9345 (LPB, MO, QCNE). type; isolectotype: GH) and pistillate inflorescences
of C. membranacea;in COL only pistillateinflores-
The trees of the species are normally small to
cences of C. membranacea.
medium-sized but may sometimes, as in flood-plain
Cecropia mituana Cuatrecasas,Revista Acad. Colomb.
forest (in Ecuador),become up to 30 m tall. In areas Ci. Exact. 9(36/37): 337. 1956. Type. Colombia.
where Cecropia engleriana and C. concolor co-occur, Vaupes: Nr. Mitu, 16 Sep 1939 (d), Cuatrecasas
Bolivia and southem Peru (Cuzco and Madre de 6818 (holotype: US; isotype: COL).
Dios), more or less clearly transitionalfeaturescan be Cecropiaferreyrae Cuatrecasas,Revista Acad. Colomb.
found in C. engleriana, such as subpersistentstipules Ci. Exact. 9(36/37): 338. 1956. Type. Peru.Ucayali:
(Cuzco), some of the incisions of the laminadown to Nr. Pucallpa, Ferreyra9032 (holotype: US, mixed
nearthe petiole, and the numberof lateralveins in the collection, a leaf belonging to C. ficifolia, herewith
free partof the midsegmentin the rangenormalin C. designatedas the lectotype, and a staminateinflores-
concolor. In some specimenscollected in San Martin, cence belonging to C. membranacea).
the laminahave ca. 21 pairsof lateralveins in the free Tree,to 15(-20) m tall. Leafy twigs 2-6 cm thick,
partof the relativelyshort midsegment. green, subhispid to strigose (with uncinate hairs) to
Vernacular names. Ecuador: dondo (Quichua, subhirsute, occasionally glabrous. Lamina charta-
Napo); huarumo(Napo). Peru:suu, tseke, yawatseke, ceous, ca. 45 X 45 cm to 90 X 90 cm, the segments
yaw'-cfki (Amazonas); siari chal (Ucayali). Brazil: 6-8, the incisions down to ca. (3/10-)5/10-7/10(-8/
embadbabranca,imbauibatorem (Acre). Bolivia: am- 10), if the incisions beyond 5/10, then the free partof
baibo blanco (Beni). the midsegmentbroadlyspathulateto obovate;apices
obtuse to rounded,sometimes shortlyacuminate;up-
per surfacescabrous(or scabridulous),hispidulousto
12. Cecropia ficifolia Snethlage, Notizbl. Bot. Gart.
substrig(ill)ose (or to hirtellous); lower surface
Berlin-Dahlem8: 365. 1923; Berg, Acta Amazon-
sparselyto densely (sub)hirsute(to subvillous)on the
ica 8(2): 171. 1978. Type. Brazil. Acre: Rio Jurua'-
veins, with arachnoid indumentum in the areoles
Mirim,Jun 1901, Ule 5588 (holotype:B (with sta-
minate inflorescences),destroyed,in G, K, L, and (sometimes very sparse), often also on the smaller
MG specimens with pistillate inflorescences,the veins, sometimes extendingto the main veins; lateral
one in MG here designatedas the lectotype). veins in the free part of the midsegment 6-12, sub-
Fig. 20 marginallyloop-connected,mostly unbranched;pet-
iole 35-80 cm long, subhispidto subhirsute(to sub-
Cecropia standleyi Macbride, Publ. Field Mus. Nat.
villous) and with short arachnoid indumentum,
Hist., Bot. Ser. 11: 61. 1931. Type. Peru. Loreto:
occasionally glabrous;trichilia fused, the brown in-
Mishuyacu,nr.Iquitos,Oct-Nov 1929 (i), Klug399
(holotype:F; isotypes: NY, US). dumentum intermixed with dense ratherlong white
Cecropia hormigana Cuatrecasas,Revista Acad. Col- hairs; stipules (12-)18-25(-35) cm long, reddish or
omb. Ci. Exact. 6(22/23): 285. 1945. Type. Colom- greenish to yellowish (to orange) or whitish, subser-
bia. Putumayo:Rio San Miquel o Sucumbios, Que- iceous to subhirsute(to subvillous) and with sparse
bradade La Hormiga,17 Dec 1940 (Y), Cuatrecasas or sometimeswith very dense arachnoidindumentum
11141 (holotype: COL; isotype: F).
outside, densely (sub)sericeousinside. Staminatein-
Cecropia discolor Cuatrecasas,Revista Acad. Colomb.
florescences in pairs, the peduncle erect, the spikes
Ci. Exact. 6/22/23): 290. 1945. Type. Colombia.Pu-
tumayo:Rio Putumayo,PuertoOspina,25 Nov 1940 pendulous; peduncle 6-12(-18) cm long, hispidulous
(Y), Cuatrecasas 10787 (holotype: COL; isotype: to (sub)hirsute,occasionally glabrous;spathe 10-18
F). cm long, green to pale yellow or whitish, substrigose
Cecropia magnifolia Cuatrecasas,Revista Acad. Col- to subsericeousto subhirsute(to subvillous) and with
omb. Ci. Exact. 6(22/23): 291. 1945. Type. Colom- sparse arachnoidindumentumoutside, pilose to sub-
bia. Caqueta:Sucre, 1000- 1300 m, 4 Apr 1940 (Y), sericeous inside; spikes 8-13, 8-10 X 0.3-0.5 cm,
Cuatrecasas9092 (holotype:COL; isotype: F).
with stipes to 1.5 cm long and hispidulous; rachis
Cecropia mocoana Cuatrecasas,Bot. Mus. Leafl. Har-
vardUniv. 14: 24. 1949. Type.Colombia.Putumayo: hairy,the hairsrelativelylong. Staminateflowersses-
Nr. Mocoa, 3-7 Dec 1942 (ci), Schultes & C. E. sile or short-pedicellate;perianth tubular,ca. 1.5-2
Smith2003 (holotype:F). mm long, sometimes with sparseand shortarachnoid
SYSTEMATIC TREATMENT 79

FI.2.Cecropia ficifolia.1. Lamina, reduced (Francoet al. 4519). 2. Lamina, reduced. 3. Apex of lamina and
venation (Gentryet al. 39501).4. Stipules (Croat18177).5. Base of petiole with trichilium(Palacios5673).6. Staminate
inflorescencewith spathe. 7. Staminateinflorescenceat anthesis and base of petiole with trichilium.8. Staminateflower.9.
Stamen (Berget al. 1066). 10. Pistillate inflorescences with spathes and base of petiole with trichilium. 11. Pistillate
inflorescenceat anthesis. 12. Pistillate inflorescenceafter anthesis (Berg1224).13. Pistillate flowers. 14. Fruit(Berget al.
1637).
80
indumentumbelow the apex, the apex plane and gla-
brous; filaments flat; anthers ca. 0.6-0.8 mm long,
appendiculate,detachedat anthesis,reattachedto the
marginsof the apertureby the appandages.Pistillate
inflorescencesin pairs, erect, becoming pendulousin
fruit, occasionally subtendedby caducous bracts, to
6 cm long; peduncle (5-)9-21 cm long, hispidulous
to (sub)hirsute,occasionally glabrous; spathe 9-14
cm long, the color and indumentumas in the stami-
nate inflorescence;spikes 4-5(-8), (2-)6-12 X 0.6-
0.8 cm, to 25 X 1.7 cm in fruit, sessile or sometimes
with stipes to 0.8 cm long and puberulous;rachis
hairy,the hairsrelativelylong. Pistillateflowers: per-
ianth 2.5 mm long, with arachnoidindumentumbe-
low the apex outside, also in the lower part of and
below the style channelinside, the apex slightly con-
vex to plane or in fruit ? concave, muriculate,the
aperture circular, surroundedby a low rim; style
ratherlong, S-shaped or straight;stigma penicillate.
Fruit oblongoid, ca. 1.5-2.5 cm long, ? tuberculate.
Distribution (Fig. 10.2). Upper Amazon basin
and the llanos region of Colombia, in forest, in par-
ticularin the southempartof the rangeof distribution
common in secondary growth, in non-inundated
places, at elevations to 1200 m.
Representativespecimensexamined.COLOMBIA.
AMAZONAS: Leticia, Quebradade Arara,27 Jan 1969 (Y fl-
fr), Cuatrecasaset al. 27244 (COL, K, NY, US); Villa Bit-
tencourt, nr. Colombia-Brazil border, 5-6 Oct 1952 (st),
Garcia-Barriga 14722 (COL); between Leticia & Rio
Amaca-yacu,nr. Isla Santa Soffa, 15 Aug 1972 (Y fl), Id-
robo 6520 (AAU, COL); Mun. Leticia, Parque Nacional
NaturalAmacayacu, nr. Quebradade Agua Pudre, 21 Mar
1992 (Y fl), Rudas et al. 3353 (COL, MO); Rfo Igara-
parana,nr. Cgto. La Chorrera,7 Oct 1973 (d), Sastre 2404
(COL, G, U), 4 Jun 1974 (6), Sastre 3110 (COL, G, U).
AMAZONAS/VAUPES: Rfo Apaporis, Jino-Goje, between
Rfo Piraparana& Rfo Popeyaka, 3-11 Sep 1952 (Y fl-fr),
Garcia-Barriga 14442 (US); Rfo Apaporis, Soratama,be-
tween Rfo Pacoa & Rio Kananari,11 Jul 1951 (6d), Schultes
et al. 13028 (US); Rio Apaporis, Raudal de Jinjirimo,25
Nov 1951 (6), Schultes et al. 14555 (COL, GH, U, US).
CAQUETA: Nr. Araracuara,11 Jul1986( fl-fr),Berget al.
1550 (BG, COL); Florencia,29 Mar 1940 (Y fl-fr), Cuatre-
casas 8816 (COL, F); Rio Orteguaza,San Luis, Quebrada
de Miramar,15 Jan 1969 (Y fl-fr), Cuatrecasaset al. 27125
(COL, F, G, K, NY, US), (6), Cuatrecasas27126 (COL, F,
K, NY, US); Florencia,between Rfo Hacha & Rfo La Yuca,
25 Jan 1968 (Y fl-fr), Cuatrecasas et al. 27236 (COL, P,
US); Mun. Florencia,rd. to Gabinete,km 1.5, 18 Oct 1993
(9 fl-fr), Franco et al. 4519 (BG, TULV); Solano, 8 km SE
of Tres Esquinas, Rfo Caqueta,below mouth of Rfo Orte-
guaza, 2 Mar 1945 (9 fl-fr), Little et al. 9498 (COL, F, P,
US); Rfo Orteguaza,9 km S of Florencia, 25 Jan 1969 (9
fl-fr), Plowmanet al. 2272 (GH, K, S, US); Morelia,22 Nov
FLORANEOTROPICA
1941 (9 fl), SneidernA.1352 (A, K, LL, MICH, NY, US).
GUAINIA: Serraniade Naquen, Maimachi, 11 Apr 1993 (9
fl), Madrinidnet al. 982 (COL), (c), Madrinidnet al. 1000
(COL). META: Mun. Puerto L6pez, Alto Menagua, 10 Feb
1995 (9 fl-fr), Franco et al. 4658 (BG, COL, HUA, MO,
US); nr. La Macarena,Rio Moccoroy, 11 Feb 1995 (6),
Franco et al. 4664 (BG, COL, HUA, MO, US); Sabanasde
San Juande Arama,Rio Guejar,Los Micos, 22 Jan 1951 (9
fl-fr), Idroboet al. 1216 (COL, GH, U, US); SierraLa Ma-
carena, mouth of Rio Duda, 7 Oct 1975 (9 fl-fr), Idrobo
8394 (AAU, COL); Sierra de La Macarena,Rfo Guapaya,
29 Nov 1949 (9 fr), Philipson et al. 1623 (BM, COL, US).
PUTUMAYO:Rio Putumayo,PifiuniaNegra,20 Nov 1940 (d
fl), Cuatrecasas10702 (COL);Mun. PuertoAsis, rd.to Kan-
akas, 16 Feb 1995 (9 fl-fr), Franco et al. 4682 (COL);Mo-
coa, betweenLa Campucana& San Antonio, 1000 m, 2 May
1994 (9 fl-fr), Franco et al. 5511 (BG, COL). VAUPES: Rio
Unilla, Calamar,30 Oct 1939 (9 fl-fr), Cuatrecasas 7328
(COL, US); Rio Guaviare, San Jos6 del Guaviare,6 Nov
1939 (9 fl-fr), Cuatrecasas7479 (COL,US); nr.Miraflores,
6-29 Jan 1944 (9 fl-fr), GutierrezV et al. 542 (COL,F, GH,
MEDEL);nr. Miti, 17 Sep 1951 (i), Schulteset al. 13904
(COL, US); Rio Apaporis, Jinogoj6, 8 Mar 1952 (9 fl-fr),
Schultes et al. 15876 (US).
VENEZUELA. AMAZONAS: Depto. Rio Negro, Rio
Mawarinuma,7 km ESE of Puerto Chimo, 9-14 Jul 1984
(9 fl-fr),Davidse et al. 27161 (BG, NY); Depto. Rio Negro,
Neblina Base Camp,on Rio Mawarinuma,17 Jul 1984 (G),
Davidse et al. 27450 (BG, MO, NY); 8 km NE of San Carlos
de Rio Negro, 1 Dec 1977 (9 fl-fr), Liesner 4076 (MO, U,
VEN); 12 km NE of San Carlosde Rio Negro, 14 Apr 1979
(d), Liesner 6617 (MO, U, VEN); 20-25 km SE of Puerto
Ayacucho,4 Nov 1980 (i), Maas et al. 5105 (F, K, NY, U,
VEN); Rio Casiquiare,just above Capibara,2 Apr 1953 (9
fl), Maguireet al. 34787 (BG); Depto. Atures,rd. Gavilan-
Rfo Cataniapo(damsite), 11 May 1980 (i), Steyermarket
al. 122299 (MO, U, VEN); Depto. Atabapo,nr.Culebra,Rio
Cunucunuma, 22-29 Mar 1983 (6), Steyermark et al.
129060 (COL, U, VEN); Yavita,26 Jan 1942 (9 fl), LI. Wil-
liams 13983 (G, US, VEN); Solano, Rfo Casiquiare,12 Mar
1942 (9 fl-fr), Ll. Williams14760 (F, G, US); Rio Guainia,
Yavita-Pimichintrail,nr.Pimichin,2 Jul 1959 (6), Wurdack
et al. 43287 (BG, NY, US).
ECUADOR. MORONA-SANTIAGO: Nr. Lim6n (Gral.
Plaza), 1200 m, 27 Jan 1981 (9 fr), Berg 1224 (BG, COL,
QCA, U); rd. Lim6n (Gral. Plaza)-La Uni6n, km 7-10, 18
Feb 1987 (9 fl-fr), Bohlin et al. 1468 (BG, GB, QCA, S);
rd. Sucua-Macas, km 10, 24 Sep 1979 (6), Holm-Nielsen
et al. 20426 (AAU); 6 km SE of Gualaquiza,16 Sep 1975
(9 fl-fr), Little et al. 386 (COL, LOJA,Q, QAME, QCNE).
NAPO:Tena, 10 Oct 1939 (9 fl-fr), Asplund 9198 (S); rd.
Coca-Auca, 26 Feb 1980 (9 fl-fr), Berg et al. 1089 (AAU,
BG, COL, QCA, U); Rio Napo, 15 km W of Coca, 18-20
Apr 1985 (9 fl), Neill et al. 6336 (BG, MO, QCNE); Ani-
angu, ParqueNacional Yasunf,nr. Tiputinitrail, 14-22 Jan
1985 (9 fl-fr), 0llgaard et al. 57100 (AAU, BG, QCA);
ReservaBiol6gica JatunSacha,8 km E of PuertoMisahualli,
1-15 Sep 1987 (9 fl-fr), Palacios 1891 (BG, GB, K, MO,
NY, QAME, QCNE);Cant6nEl Chaco, Codo Sinclair, 16-
SYSTEMATICTREATMENT 81

20 Sep 1990 (9 fl-fr), Palacios 5673 (BG, QCNE). PAS- exanderde Humboldt,9 Sep 1980 (9 fl-fr),Maas etal. 4551
TAZA: Mera, ca. 1100 m, 13 Dec 1955 (9 fl-fr), Asplund (BG, USM).
18829 (S); nr. Puyo, 19 Dec 1955 (9 fl-fr), Asplund 18892 BRAZIL. ACRE:Nr. Cruzeirodo Sul, 21-28 Aug 1978
(S); nr. Mera, 1160 m, Harling 3809 (S). SucuMBios: Rio (61),Benson 8304 (INPA,UEC, US); Mun. Cruzeirodo Sul,
Cuyabeno, ca. 0?10'S, 75?55'W,20 Feb 1980 (d), Berg et Rio Jurua,Seringal Sao Joao, 14 Mar 1992 (9 fl-fr), Daly
al. 1066 (AAU, BG, COL, GB, MO, NY, QCA, TUR, U, et al. 7418 (BG, NY), 19 Mar 1992 (6), Daly et al. 7529
WIS), 20 Feb 1980 (9 fl-fr), Berg et al. 1071 (AAU, BG, (BG, NY); Rio Acre, Antimary, 30 Mar 1904 (9 fl-fr),
COL, NY, QCA, U); Lumbaqui,ca. 1000 m, 13 Aug 1975 HuberMG 4245 (BM, G, MG, U); Cruzeirodo Sul, Estrada
(9 fl-fr),Littleet al. 167 (Q, QAME,QCNE);ReservaFaun- Alemanha, 27 May 1971 (9 fl-fr), Maas et al. P13319
istica Cuyabeno, between Tarapoa& Tipishca, Bellavista, (INPA, K, MO, NY, S, U, US). AMAZONAS: Rio Negro,
14 Nov 1991 (9 fl-fr), Palacios et al. 8884 (BG, QCNE). Cucui, 4 May 1975 (6), Cavalcante3097 (MG), (9 fl-fr),
ZAMORA-CHINCHIPE: Rfo Nangaritza, nr. Nangaritza, 4 Cavalcante 3098 (MG); Codajas, Ig. do Engenho, 19 Apr
Dec 1990 (9 fl-fr), Neill et al. 9504 (BG, QCA, QCNE). 1952, (9 fr), E. Ferreira 58/239 (INPA, U); Mun. Tefe,
PERU. AMAZONAS: Rio Cenepa, 10 km E of Huam- Jauata,21 May 1933 (9 fr), Krukoff4517 (A, G, K, NY, S,
pami, 10 Oct 1972 (9 fl), Berlin 229 (USM); Rio Santiago, U, US); Mun. Humaita,nr.Tres Casas, 14 Sep-Il Oct 1934
nr.Caterpiza,29 Aug 1979 (6), Huashikat253 (MO), 6 Oct (9 fl), Krukoff6420 (NY); Mun. Sao Paulo de Olivenga,nr.
1979 (9 fl-fr), Huashikat793 (MO); QuebradaKayamas,3 Palmares, 11 Sep-26 Oct 1936 (d), Krukoff8333 (A, BM,
Mar 1973 (9 fl-fr), Kayap 442 (BG, MO). HUANUCO: F, K, LE, MICH,MO, NY, P, S, U, US); nr.Uaupes, 10 Feb
Puerto Bermddez, 14-17 Jul 1929 (st), Killip et al. 26614 1963 (9 fl-fr), Lanna Sobrinho451 = Castellanos 23756
(NY); Prov. Pachitea, Dtto. Honoria, Rio Pachitea, rd. to (GUA, RB, US); Rio Javari,PalmeirasArmy Post, 2 Aug
Ayamiria,27 Dec 1966 (9 fl-fr),SchunkeV 1417 (F, MOL); 1973 (9 fl-fr), Lleras P 17033 (COL,F, INPA, K, MG, MO,
Prov. Pachitea, Dtto. Honoria, Isla del Pacanasi, 14 Nov NY, U); Rio Javari,Angamo Garrison,between Angamo &
1967 (d), SchunkeV 2313 (F, US). LORETO:Rd. Iquitos- Natua, 5 Aug 1973 (d), Lleras P17161 (INPA, K, NY, U);
Nauta, km 2, 4 Jul 1988 (9 fl-fr), Berg et al. 1573 (BG, Rio Negro, Aperecida,IgarapeTuari,6 Nov 1987 (6), Maas
USM); 4 km SW of Iquitos, 15-16 Jul 1972 (6), Croat et al. 6934 (INPA, NY); Tapuruquara,17 Oct 1971 (9 fl-
18128 (BG, MO); Prov. Requena, Jenaro Herrera,9 Dec fr), Prance et al. 15375 (INPA); Manacapuru,4 Apr 1957
1977 (9 fl-fr), Gentry et al. 21300 (F, MO, USM), (d), (6'), Rodrigues 403 = INPA 5282 (INPA, U). MATO
Gentry et al. 21333 (MO, USM); Prov. Maynas, Rio Am- GROSSO:Rio Aripuana,nr. Igarapezinho,11 Oct 1973 (9
piyacu, Pebas, 30 Mar 1977 (6), Plowman et al. 6495 (F, fr), Berg et al. P 18419 (AAU, COL, EAP, F, GH, INPA, K,
US, USM); Prov.Maynas,Rio Yaguayacu,Brillo Nuevo, 12 MG, MICH, MO, NY, P, S, U, US); Nucleo Pioneiro de
Apr 1977 ( 9 fl-fr),Plowmanetal. 6805 (F, US, USM); Prov. Humboldt,Rio Aripuana, 14 Oct 1973 (9 fl), Berg et al.
Maynas,Rio Nanay,Mishana,27 Oct 1980 (9 fl-fr),R. Vds- P 18526 (F, INPA, K, MG, NY, P, S, U, US); Presidente
quez et al. 676 (MO, NY, TEX, USM); Prov. Alto Ama- Marques,Rio Mamore, 11 Oct 1923 (9 fl), Kuhlmann591
zonas, Yurimaguas,18 Sep 1981 (d), R. Vdsquezet al. 2568 = (RB) 19890 (RB); Nucleo Pioneiro de Humboldt, Rio
(F, MO. MADRE DE Dios: Parque Nacional Manu, Rio Aripuana,9 Oct 1973 (6), Prance et al. 18294 (F, K, NY,
Manu, above Rio Sotileja, 11 Oct 1986 (9 fl), Foster et al. S, U). ROND6NIA: Nr. Sao Loren9o mines, 26 Nov 1968
11777 (BG, LPB, US, USM); Prov. Tambopata,Reserva (61), Prance et al. 8910 (INPA,K, MG, MO, NY, S, U, US).
Tambopata,15 Aug 1990 (9 fl-fr), Reynel et al. 5144 (BG, BOLIVIA. BENI: Prov. Vaca Diez, nr. Alto Ivon, Dec
MO); Prov. Tambopata,Pampasde Heath, 25 Feb 1990 (9 1992 (9 fl-fr), Bergeronet al. 753 (BG), 25 Jan 1984 (9 fl-
fl-fr), Gentry et al. 69558 (BG, MO); Prov. Tambopata,nr. fr), Boom 4264 (NY); Prov.Vaca Diez, rd. Riberalta-Guay-
Sandoval, 15 Jan 1967, VargasC. 18572 (US). PASCO:Prov. aramerin,km 20, 20 Apr 1979 (d fl-fr), Krapovickaset al.
Oxapampa,Panjil, 12 km from Puerto Inca, 27 Sep 1982 35179 (RB); Prov.Vaca Diez, rd. Riberalta-Guayaramerfn,
7 1981 (9 fl-fr), Solomon6165 (LPB, MO, U).
(6), D. N. Smithet al. 2427 (MO). SANMARTiN:Prov.San km 3.3, Sep
PANDO: Prov.Manuripi, 12 km W of Conquista-Trampolin,
Martin,rd. Yurimaguas,ca. km 24, 5 Dec 1977 (6), Berg
et al. 1776 (BG, COL, MOL), ca. km 27, 5 Dec 1997 (9 fl- 6 Oct 1991 (st), Beck et al. 20168 (BG, LPB); prov Nicolas
nr. 10 Jan 1983 (9 fl), FerndndezC. et al.
fr), Berg et al. 1778 (BG, COL, MOL); Prov. Lamas, rd. Suarez, Porvenir,
Dec 8152 (G, NY); Cobija, Jan 1912 (9 fl-fr), Ule 9311 (F, G,
Tarapoto-Moyobamba,ca. km 70, 8 1997 (9 fl-fr),
MG).
Berg et al. 1789 (BG, COL, MOL); rd. Yurimaguas-Tara-
poto, km 50, 12 Oct 1985 ( 9 fl-fr), Gentryet al. 52282 (BG, This is a very variable species in the leaf apex, the
F, MO);rd.Tarapoto-Yurimaguas, km 68, trailPintoyacillo- leaf shape, and the indumentum of the lamina, the
Nuevo Lamas-San Miguel de Shanusi,2 Oct 1986 (9 fl-fr),
stipules, and the inflorescences. Arachnoid indumen-
Knapp et al. 8482 (BG, QCNE, USM); San Roque, 13 Jan
tum may be sparse or very dense. In Ecuador and
1930 (9), Ll. Williams7372 (F). UCAYALI: NuevaRequena,
Colombia, the lower leaf surface is often white by
11 km from CampoVerde(at km 33 rd. Pucallpa-Lima), 11
arachnoid indumentum. Moreover, the stipules
Aug 1988 (9 fl-fr), Berg et al. 1637 (BG, COL, K, MOL); dense
nr. Pucallpa, (9 fl-fr), Ferreyra9043 (US, USM); between and spathes are usually green to yellowish or whitish
Aguaytia & San Alejandro, 9 Nov 1964 (9 fr), Ferreyra because of the cover by dense arachnoid indumentum,
16102 (US, USM); rd. Pucallpa-Tingo Maria, Bosque Al- and other types of indumentum may be denser and
82 FLORANEOTROPICA

more conspicuous as well. In Colombia, the main CecropiamonostachyaC. C. Berg, Nord. J. Bot. 1: 485.
veins of the lamina beneathand the petiole are often 1981; Berg & FrancoRosselli, Fl. Ecuador48: 38, t.
densely hirsuteto subvillous. In the northernpart of 6. 1993. Type. Ecuador.Pichincha: Old rd. Quito-
the distributionrange, as in Meta and Caqueta(Co- Santo Domingo de los Colorados, km 45, ca. 1600
m, 10 Sep 1977 (Y), Berg et al. 416 (holotype: U;
lombia) and San Martin (Peru), the incisions of the
isotypes: AAU, NY, QCA).
laminaare <5/10 down to the petiole andthe number
of lateralveins in the free part of the midsegmentis Tree, to 15 m tall. Leafy twigs 1.5-3 cm thick,
often down to only 6 pairs. Moreover,the laminacan reddishto purpleor greenish,glabrousandthen often
be very large (to 95 cm long). These specimens are with a bluish (waxy) layer, or occasionally with ?
quite distinct from the more typical type, normally dense arachnoidindumentum;intemodes completely
with the characteristicbroadlyspathulatefree partof or largely filled with brownpith. Laminacoriaceous,
the midsegment. The type with dense indumentum ca. (20 X 20 cm to) 35 X 35 cm to 70 X 70 cm, the
and shallowly incised laminastendsto be ecologically segments 7-8, the free parts the upper segments ob-
differentas well, as it does not occur in dense stands ovate to elliptic, the incisions down to 5/10-8/10, in
in secondarygrowth,but ratheris dispersed,often in the upper part, less deeply in the lower part; apices
marginsof tree fall gaps. In areas (mainly in or close acute, acuminateto rounded;upper surface smooth,
to the foot hills of the Andes) where the species oc- subglabrousor puberulouswith sparsewhite hairson
curs as treelets in small gaps or in the peripheryof the main veins and "umbilicus,"occasionally with
largertree fall gaps, it is rarein man-madesecondary rather dense arachnoid indumentum;lower surface
growth, whereas elsewhere (in the Amazon basin) it glabrousor sparsely puberulousto hirtellousto sub-
is common. Therefore,the species may comprisetwo hirsute on the veins, with arachnoidindumentumin
subspecies. The leafy twigs and petioles are glabrous the areoles, also on the smaller veins, sometimes ex-
in the two collections from Guainia(Colombia).The tendingto the main veins; lateralveins in the free part
species shows clear morphological similarities to of the midsegment 12-18 pairs, submarginallyloop-
Cecropia obtusa. Prance et al. 15375, from Amazo- connected, sometimes branched;petiole (10-)20-50
nian Brazil, shows featurestransitionalto C. obtusa. cm long, glabrousor sometimes sparselyhirtellousto
Subjuvenileleaves of C. kavanayensisare quite sim- (sub)hirsutetoward the apex and the base or with
ilar to the relativelydeeply incised type of leaf of C.arachnoid indumentum;trichilia absent or separate
ficifolia. and poorly developed or fused and well-developed,
Vernacular names and use. Colombia: cetico the brown indumentumintermixed with very short
blanco, yarumode raton (Amazonas);jaibena, kYraY- white or sometimes long hairs;stipules ca. 15-30 cm
kai iikai (Witoto, Amazonas);udagomo (Mui.). Ven- long, pinkish to red or greenish, glabrous or some-
ezuela: toc'kori, yagrumo, yagrumo blanco (Ama- times (partly) sparsely subvillous and with brown
zonas). Ecuador: anduchi dundo, arduchinadurdu, pluricellularhairsor also arachnoidindumentumout-
dundu (Quichua, Napo); huarumo (Napo). Peru: side, glabrous inside. Staminate inflorescences soli-
tsake, suu (Amazonas);setico blanco (Loreto);sutiik tary or in pairs,erect;peduncle5-7.5 cm long, almost
(Jivaro,Loreto). Brazil: imbaubabranca(Amazonas; glabrous with very few long and appressed hairs;
Tukuna Indians use leaf petioles to make flutes for spathe9-19 cm long, pale brownto reddishor green-
pubertyrites). ish, (sub)glabrous(or subvillous) outside, glabrous
(or subvillous) inside; spikes 3-6, 6.5-15 X 0.5-1.5
cm, with stipes 0.4-0.5 cm long and glabrous;rachis
13. Cecropia gabrielis Cuatrecasas,Revista Acad. hairy.Staminateflowers with pedicels to 1 mm long;
Colomb.Ci. Exact. 9(36/37): 334. 1956. Type.Co- perianthtubular,2-2.4 mm long, with short straight
lombia. Antioquia:Nr. Angel6polis, 1950 m, 23 hairs or sometimes also with arachnoidindumentum
Nov 1947 (d), Barkley & Gutierrez17C675 = below the apex, the apex slightly convex, glabrousor
Gutierrez & Barkley 17C675 (holotype: F; iso- sparsely muriculate;filaments flat; anthers 0.6-0.8
types: COL, MEDEL,VALLE). Fig. 21 mm long, appendiculate,detachedat anthesis(?). Pis-
tillate inflorescencessolitaryor in pairs,erect;pedun-
CecropiadiguensisCuatrecasas formaalbicansCuatre-
cle 3.5-13 cm long, glabrous or sometimes with
casas,RevistaAcad.Colomb.Ci. Exact.9(36/37):
333. 1956.Type.Colombia.Valle:RfoSanJuan,be- arachnoidindumentum;spathe 13-26 cm long, the
low Queremal, 1350-1400 m, 19 Mar 1947 (d), color and indumentumas in the staminateinflores-
Cuatrecasas23884 (holotype: US; isotypes: F, US, cence; spikes 1 or 2, 8-15 X ca. 1 cm, to 25 X ca.
VALLE). 2.5 cm in fruit,subsessile;rachishairy.Pistillateflow-
SYSTEMATICTREATMENT 83

FIG. 21. Cecropiagabrielis. 1. Leafy twig (Berg et al. 416). 2. Base of petiole with trichilium(Websteret al. 27296).
3. Staminateinflorescencewith spathe and at anthesis (Berg 1291). 4. Pistillate inflorescencewith at anthesis. 5. Apices of
pistillate flowers (Berg et al. 416). 6. Staminateflower. 7. Stamen and filament after detachmentof anther(Franco et al.
5563). 8. Pistillate flower. 9. Fruit with persistent style (Franco et al. 4469). (By T. Schipper and Hendrieke Berg. From
Nord. J. Bot. 1: 486. 1981, modified.)
84 FLORANEOTROPICA

ers: perianth ca. 3-4(-6) mm long, with arachnoid part of the species range, whereas very often com-
hairs below the apex outside, sometimes also in the pletely absent in the southernpart of its range. The
style channelinside, the apex slightly convex to plane, leafy twigs and the upper surface of the lamina are
muriculateor smooth;apertureslit-shaped;style long, occasionally covered with white arachnoidindumen-
S-shaped to straight,hairy; stigma penicillate. Fruit tum (Webster et al. 27635). This species is very
ovoid, ca. 3 mm long, finely tuberculateto smooth. closely relatedto C. telealba, andprobably(moredis-
tantly) also related to C. bullata. It can be distin-
Distribution (Fig. 18.6). WesternColombia (An-
guished from the formerby usual absence of arach-
tioquiato Nariiio)and northwesternEcuador,in mon-
noid indumentumon the uppersurfaceof the lamina
tane forest, at 1200-2000 m.
and from the latter by the absence of villous indu-
Representativespecimensexamined.COLOMBIA. mentumon the leafy twigs and petioles. The trees are
ANTIOQUIA: Mun. Cocorna,Vrda. Viao, 1950 m, 26 Mar not inhabitedby ants.
1994 (Y fl-fr), Franco et al. 4589 (BG,HUA),(d), Franco
et al. 4590 (BG,HUA);nr.Yarumal, 1900m, 27 Mar 1994
(6), Franco et al. 4594 (BG, HUA).NARINO: Mun.Ri-
14. Cecropia garciae Standley,Publ. Field Mus. Nat.
caurte, rd. to Reserva NaturalLa Planada, 1800 m, 8 Oct
1993 (6), Franco et al. 4466 (BG,COL),(Y fl-fr), Franco
Hist., Bot. Ser. 22: 71. 1940. Type. Colombia.
et al. 4469 (BG,COL), (6), Franco et al. 4471 (BG,COL), Cauca: Gorgona Is., 11 Feb 1939 (Y), Killip &
22 Feb 1995 (Y fl-fr), Franco et al. 5542 (BG,COL),(6), Garcia 33221 (holotype:F; isotype: COL, US).
Franco et al. 5543 (BG, COL,HUA,LP,MO, US); Mun. Fig. 22
Barbacoas,Cgto. Altaquer,Vrda. El Barro, 1325 m, 6 Dec
Tree,to 12 m tall, often with a short trunk.Leafy
1993 (st), Franco et al. 4959 (COL), 10 Dec 1993 (6),
twigs 3-5 cm thick, dark green to dark purple;his-
Franco et al. 5107 (COL), 11 Dec 1993 (9 fl), Franco et
al. 5142 (COL);ReservaNaturalLa Planada,aboveChu-
pidulous to hispid to setose with irritatingbristles.
cunes, 1700 m, 3 Jan 1989 (9 fl-fr), Gentry et al. 64399 Lamina (sub)coriaceous,ca. (55 X 55 cm to) 70 X
(PSO); Reserva NaturalLa Planada, 1800 m, 14 Jan 1989 70 cm to 11O X 110 cm, the segments 14-19, lan-
(9 fr), Restrepo 440 (MO, PSO); Ricaurte,1300 m, 12 Apr ceolate, the incisions down to the petiole; apices acu-
1941 (9 fl-fr),Sneidern A.563 (A, COL, K, LL, MICH,NY, minate;marginin dry material ? revolute;uppersur-
S). VALLE: Mun. Dagua, rd. to Queremal,ca. 76 km from face scabridulous (to scabrous or to smooth), very
Buenaventura,1300 m, 17 Mar 1994 (9 fl-fr),Franco et al. sparsely hispidulous, the "umbilicus" (sub)setose;
4556 (BG, TULV);Rio Bitaco, 1 km E of Bitaco, ca. 1500 lower surface with arachnoidindumentumin the ar-
m, 16 Nov 1963 (9 fl-fr), Hutchinson et al. 3039 (COL, F).
eoles and on the smaller veins, extending to the pri-
ECUADOR.CARCHI: Nr. Maldonado, ca. 1500 m, 5 mary and lateral veins, these also with brown pluri-
Sep 1981 (9 fl-fr), Balslev 1984 (QCA); Rio Blanco, abovecellular hairs; lateral veins ca. 35-50 pairs,
Chical, 1300-1500 m, 25 Sep 1979 (6), Gentryet al. 26527
submarginallyloop-connected, unbranched;petiole
(MO). PICHINCHA: Rd. Empalme-Chiriboga,ca. 1350 m,
ca. 50-100 cm long, with ? dense arachnoidindu-
21 Feb 1981 (6), Berg 1291 (AAU, QCA, U, US), 9 Feb
1994 (9 fl-fr),Berg 1687 (AAU, BG, COL,QCNE);Cant6n
mentum and setose, often only adaxially at the base
Quito, Chirobiga,ReservaForestalLa Favorita,1600-1800 and in the uppermostpart;trichilia fused, the (pale)
m, 8 Feb 1990 (st), Ceron et al. 8593 (MO); Maquipucuna, brownindumentumintermixedwith shortbrown(uni-
5 km E of Nanegal, 1550 m, 9 May 1990 (6), Gentryet al. cellular)hairsor sometimesalso sparselysetose; stip-
69901 (QCNE);Nanegal, ReservaEcol6gica Maquipucuna, ules 18-30 cm long, darkred-brownor darkpurple-
1250 m, 9 Jan 1992 (9 fl-fr), Quelal 65 (BG, NY, QCNE); red, sparsely to ratherdensely setose with appressed
rd. Quito-Manabf,Sep 1874 (st), Sodiro s.n. (Q); Nanegal,or patent irritatingbristles outside, glabrous inside.
Sep 1902 (st), Sodiro s.n. (Q); between Hda. El Carmen Staminateinflorescencesin pairs, erect; peduncle 4-
(Maquipucuna)& Marianitas,Rio Umacacha, 1200-1500 12.5 cm long, hispidulous or partly (short-)setose;
m, 2 Sep 1989 (9 fl-fr), Websteret al. 27296 (BG, MO, spathe 10-15 cm long, darkred, sparselysetose with
QCA); Reserva Maquipucuna,trail Rio Umacacha-Hda.El
appressedto patent bristles outside, glabrousinside;
Carmen,4.5 km SE of Nanegal, ca. 1250 m, 16 Sep 1989
spikes ca. 15-35, 4-13 X ca. 0.3 cm, (sub)sessile;
(d), Websteret al. 27635 (BG, QCA), 17 Sep 1989 (9 fl-
rachis hairy. Staminateflowers: perianthtubular,ca.
fr), Websteret al. 27689 (BG, QCA).
1 mm long, with sparse shortarachnoidindumentum
In Colombia, this species has sometimes two below the apex, the apex plane, muriculateto hispi-
spikes in the pistillate inflorescence.All states of de- dulousor smooth;filamentsflat;anthersca. 0.7-1 mm
velopment of the trichilia, from absent to well- long, appendiculate,detached at anthesis (?). Pistil-
developed,can be found, even on the same plant.Tri- late inflorescencesin pairs, erect; peduncle (3-)6-13
chilia appearto be consistentlypresentin the northern cm long, sparselyto ratherdensely hispidulousto pu-
SYSTEMATICTREATMENT 85

.~~~~~~~~~~~~~~~
~ ~~~~~~~~~~~~~A

/ nT I

floer FIG.
11 FriIFac ta.49)
22. Cecropiagarciae. 1. Lamina,reduced.2. Apex of lamina and venation (Jaramilloet al. 13740). 3. Stipules
and young leaf (Franco et al. 4535). 4. Twig with opened prostomaand base of petiole with trichilium(Tipazet al. 1258).
5. Pair of staminateinflorescences with spathes and base of petiole with trichilium.6. Staminateinflorescence at anthesis
(Franco et-al. 5176). 7. Staminateflower and stamen (Franco et al. 4535). 8. Pair of pistillate inflorescences with spathes
and base of petiole with trichilium(Franco et al. 5177). 9. Pistillateinflorescenceat anthesis(Tipazet al. 1258). 10. Pistillate
flower. 11. Fruit(Franco et al. 4693).
86 FLORA NEOTROPICA

berulous and/orpartly (short-)setose,often also with 1965 (9 fl-fr), Little et al. 21097 (MO, NY, Q, QAME,
sparse to dense arachnoidindumentum;spathe 8-12 QCNE, US); Cant6n San Lorenzo, Reserva IndfgenaAwi,
cm long, the color and indumentumas in the stami- Rfo Mira, 10 km W of Alto Tambo, 16-26 Mar 1991 (d),
nate inflorescence;spikes 5-21, 5-18 X 0.3-0.5 cm, Rubio et al. 1263 (MO, QCNE); Reserva Ecol6gica Cota-
cachi Cayapas, Rio Santiago, Angostura, 17-26 Jul 1994
to 21 X ca. 1.5 cm in fruit, (sub)sessile; rachis
(st), Tiradoet al. 1104 (QCNE).
(sub)glabrous.Pistillateflowers: perianthca. 1.5-2.5
mm long, with arachnoid indumentum below the This species is in most charactersmore or less
apex, also below the style channel inside, the apex similar to Cecropia hispidissima, but they are clearly
convex, muriculate,the apertureslightly slit-shaped; distinctin the trichilia.In C. garciae, the trichiliaare
style short;stigma comose. Fruit ellipsoid to obovoid well-defined, only sometimes with sparse irritating
or to ovoid, 1.2-1.7 mm long, tuberculate. bristles, and the Miullerianbodies are small and whit-
ish, whereas in C. hispidissimathe Muillerianbodies
Distribution (Fig. 8.7). From westernEcuadorto are large and pinkish being produced in ? loosely
eastern Panama, in forest and secondary growth, at defined trichilia with numerousirritatingbristles. In
elevations to 1050(-1800) m. C. garciae, the numberof segments of the lamina is
Representative specimens examined. PANAMA. usually 15-19, whereas it is usually 12-15 in C. his-
DARIEN: Withoutlocality,1966(juv?), Bristan 121 (MO). pidissima. Moreover,the lamina is usually scabridu-
COLOMBIA.ANTIOQUIA:Mun.Frontino, ParqueNa- lous and plane in C. garciae, but smooth and often
cional NaturalLas Orquideas,20 Jun 1982 (6), Bernal et ? plicatein C. hispidissima.Cecropiagarciae is nor-
al. 359 (COL);Mun.Frontino,Cgto. Nutibara,rd.Nutibara- mally found at elevations below 1000 m. The single
La Blanquita,Murriregion, 4 Mar 1995 (6), Franco et al. collection from a much higher elevation in Carchi
5559 (BG, COL, HUA). CAUCA: Mun. GuapI,ParqueNa-
(Ecuador)was made in an area where other lowland
cional Naturalde Gorgona, 1 May 1986 (6), Lozano C. et
species (of moraceousgenera) have also been found
al. 5082 (COL); Mun. Guapi, ParqueNacional Naturalde
Gorgona,rd. to Playa Blanca, 5 Sep 1987 (9 fl-fr), Lozano
at elevations above the normalupperlimit of lowland
C. et al. 5572 (COL, MO). NARINO:Mun. Barbacoas,nr. species. The trees of this species are normallyinhab-
El Diviso, 7 Oct 1993 (st), Franco et al. 4464 (BG, COL, ited by ants of the genus Azteca. The identityof Bris-
HUA, LP, MO, NY, P), 21 Feb 1995 (9 fl-fr), Franco et al. tan 121, the only collection fromPanama,is not quite
4693 (COL, HUA, LP, MO, NY, P); Mun. Barbacoas,rd. certain.
Tumaco-Junin,Cgto. Junin, 12 Dec 1993 (6), Franco et al.
5176 (BG), (9 fl), Franco et al. 5177 (BG); rd. Tumaco- Vernacular names. Ecuador:cosedera,yuarumbo
Tuquerres,W of Junin, 25 Nov 1981 (d), Gentry et al. (Esmeraldas).
34941 (BG, MO); 2-3 km S of Barbacoas,on rd. to Junfn,
7 Jan 1989 ( 9 fl), Gentryet al. 64550 (PSO);rd.Barbacoas-
Junin, 1050 m, 7 Aug 1962 (6), Mora 0. 2311 (COL, US); 15. Cecropia glaziovii Snethlage,Notizbl. Bot. Gart.
RR Tumaco-El Divisio, km 86, 27 Jul 1952 (9 fl-fr), Berlin-Dahlem8: 358. 1923; Berg & Carauta,Al-
Romero-Castanieda 3324 (COL). RISARALDA:Mun. bertoa 1(1): 9. 1986. Syntypes. Brazil. Rio de Ja-
Mistrat6,Cgto. Puertode Oro, Chirrincha,28 Sep 1991 (9 neiro: Between Petropolis & Sao Antonio, 1879
fl-fr), Franco et al. 3607 (BG, COL). VALLE: Rio Calima, (6), Glaziou 11559 (B, destroyed; isotypes: C,
La Trojita,19 Feb 1944 (9 fl-fr), Cuatrecasas16467 (COL,
GH, LE, P); and Mun. Rio de Janeiro,Florestada
F, VALLE);Rio Cajambre,Quebradade Ord6tiez, 11 May
Tijuca, nr. Pies, 1891 (9), Glaziou 18499 (B, de-
1944 (9 fl-fr), Cuatrecasas 17272 (F, US, VALLE);Rio
Anchicaya,Sabaletas,13 Nov 1945 (6), Cuatrecasas19804
stroyed;isotypes: A, C, G, H, K, LE, P); Glaziou
(F, US); nr. Buenaventura,15 Mar 1994 (d), Franco et al. 18499 (9) at P here designatedhere as the lecto-
4535 (BG, TULV);Mun.Buenaventura,rd. to Sabaletas,km type).
12, 17 Mar 1994 (9 fl-fr), Franco et al. 4551 (BG); Color- Cecropia macrantheraWarburgex Snethlage, Notizbl.
ada, BuenaventuraBay, 3 Jun 1944 (d), Killip et al. 38713 Bot. Gart. Berlin-Dahlem8: 357. 1923; Warburgin
(COL, F, US, VALLE). Glaziou, Bull. Soc. Bot. France 59(Mem. 3): 645.
ECUADOR.CARCHI: Reserva Indigena Awa, Gualpi 1913, nomen.Type. Brazil. Rio de Janeiro,Serrados
Alto, 1800 m, 15-28 Jun 1991 (9 fr), Rubio et al. 1535 Orgaos, 1879 (6), Glaziou 12173 (holotype:B, de-
(BG, QCNE); TobarDonoso, 19-28 Jun 1992 (d fl-fr), Ti- stroyed; isotype: P, comprising vegetative parts of
paz et al. 1258 (MO, QCNE). ESMERALDAS:Between Ma- Pouroumaguianensis Aublet subsp. guianensis and
taje & Molina, 5 Sep 1991 (9 fr), Jaramillo et al. 13740 a staminateinflorescenceof C. glaziovii).
(GB, NY, QCA); nr. TobarDonoso, jct. of Rfo Juan& Rio
Camumbi,28 Jul 1966 (9 fr), Jdtiva et al. 1151 (MO, NY, Tree, to 20 m tall. Leafy twigs 2-5 cm thick, green,
S, US); nr. Borb6n, 2 Aug 1967 (6 + 9 fl-fr), Jdtiva 2105 grayishgreen,or sometimespartlybluish,hispidulous
(MO, NY, S, U); Borb6n, 3 km S of Rio Cayapas, 19 Sep to hirtellous, with curved to uncinate hairs. Lamina
SYSTEMATICTREATMENT 87

subcoriaceousto chartaceous,ca. 25 X 25 cm to 70 Distribution (Fig. 11.1). Eastern Brazil, from Ba-

X 70 cm, the segments 8-12, the free parts of the hia to Rio Grande do Sul, in moist forest, common in
uppersegments subobovate(to subpandurate),the in- secondary growth, at elevations to 1300 m.
cisions down to 5/10-8/10(-9/10); apices roundedto Representative specimens examined. BRAZIL. BA-
acuminate; upper surface scabrous, hispidulous to HIA: Rd. Mi9anga-Ciu, 15 Feb 1980 (? fl-fr), Arauijo224
strigose; lower surface minutely puberulous(to sub- (RB); between rd. Sao Jose-Buerarema& Una, km 29, 1
tomentellous)on the main veins, subtomentellous(to Apr 1980 ( fr), Berg et al. 1146 (BG, K, NY, U); nr.Bahia,
subtomentose)on the smaller veins, with arachnoid 1839 (d), Blanchet s.n. (BM, F, G); Ilheus, 12 May 1981
indumentumin the areolesor also on the smallerveins (Y), Hage et al. 662 (CEPEC, GUA); Mun. Marau, rd.
and concentratedalong the margin;lateralveins in the BR.030, between Ubaitaba& Marad,km 33, 5 Feb 1979 (Y
free part of the midsegment 12-16(-23) pairs, sub- fl), Mori et al. 11357 (K, NY, U); Mun. Jussari, Serra do
Teimoso, 7.5 km N of Jussari,9 Feb 1998 (Y fl-fr), Thomas
marginally loop-connected, often branched;petiole
et al. 11774 (NY). ESPiRITO SANTO: VilaVelha,30-31 Aug
20-55 cm long, (minutely) puberulous(mainly with 1980 (Y), Andrade118 (GUA); SantaTeresa, 16 Nov 1988
curved to uncinatehairs) to subhispidulousor subgl- (d), BoudetFernandeset al. 2623 (GUA), (Y fl-fr),Boudet
abrous;trichilia fused, the brown indumentuminter- Fernandeset al. 2624 (GUA); Mun. Conceicao do Castelo,
mixed with ? dense rathershort white to brownish rd. BR.262, 5 Dec 1984 (Y fl-fr), Hatschbachet al. 48643
(unicellular)hairs;stipules (7-)15-27 cm long, bright (BG, US); betweenVila Verde& Domingo Martins,29 May
to darkred to purplish,sometimes greenish, sparsely 1993 (d), Simoes et al. 73 (GUA); Reserva Florestal Lin-
puberulous, sparsely to ratherdensely hirtellous or hares, 21 May 1993 (Y fl-fr), Sposito 82 (CRVD). MINAS
sometimes ? densely pale yellow- to white-hirsute, GERAIS: Mun. Vicosa, between the centre of Vicosa & the
University, 10 Sep 1967 (d), Carauta414A (GUA, RB, U,
or also with sparse arachnoidindumentumoutside,
US), (Y fl-fr), Carauta414B (GUA, RB, U, US); Mun. Ar-
glabrousor sparselyto densely hairy inside; terminal axa, between Araxd & Barreira,23 Mar 2001 (5), Ernani
buds slightly inflated. Staminate infiorescences in Diaz 446 (GUA); Juiz de Fora, 3 Sep 1970 (Y fr), Krieger
pairs, the peduncle erect to pendulous,the spikes + 9054 (GUA);Vi,osa, AgriculturalCollege, 3 Dec 1958 (st),
spreadingto pendulous;peduncle 2-13 cm long, his- Irwin 2199 (MICH,NY, US); 15 Feb 1939 (Y fl-fr), Mexia
pidulous to subglabrous;spathe 9-22 cm long, dark 4349 (A, BM, F, GH, K, MICH, MO, NY, S, U, US); Car-
to pale red or greenish, hirtellous and with sparse angola, Oct 1991 (d), Leoni 1699 (GUA). PARANA: Volta
arachnoid indumentum outside, glabrous inside; Grande, 1 Feb 1904 (d), Dusen 3542 (GH, S); Therezina,
spikes (4-)8-12(-25), (5-)8-22 X (0.3-)0.5-l.6(-2) 26 Jan 1911 (st), Dusen 11217 (GH, S); Volta Grande,31
Mar 1912 (d), Duse'n 14043 (GH, K, MICH, NY, P, S);
cm, yellow, sometimes pink, with stipes 0.5-2 cm
Mun. Morretes,Est. Marumbi,8 Feb 1985 (9 fl-fr),Hatsch-
long, glabrousand often with a bluish waxy layer or bach 48892 (BG, US); Mun. Morretes, Passa Sete, 5 Jul
hispidulous, rarely sessile; rachis hairy or glabrous. 1988 (9 fr), Hatschbach et al. 52158 (BG); Mun. Paran-
Staminate flowers: perianth tubular, (1.5-)2-2.5 mm angud,PR.53, 4 km N of Rio Guaraguacu,5 Jul 1988 (d),
long, with dense arachnoidindumentumbelow the Hatschbachet al. 52159 (BG); Paranagud,Ilha das Cabras,
apex, the apex ? convex to almost plane, muriculate 14 Jun 1986 (9 fl-fr), W S. Souza et al. s.n. (UEC). RiODE
or smooth; anthers 0.8-2 mm long, at anthesis de- JANEIRO:Nr. Cabo Frio, 16 Mar 1980 (d), Berg etal. 1134
tached, remaining attached to the filament by (BG, K, RB, U), (9 fl-fr), Berg et al. 1136 (U); Mun. Rio
stretched spiral thickenings (or not detached, or de- de Janeiro,SerraCarioca,Estradada Vista Chinesa, 19 Dec
1963 (9 fl-fr), Carauta208 (GUA, K, NY, US); Mun. Ter-
tached and reattachedto the marginsof the aperture
es6polis, GranjaComari, 10 Feb 1964 (9 fr), Carauta211B
by the appendages?).Pistillate inflorescencesin pairs, (GUA, U, US); Mun. Resende, Vila de Itatiaia, 1 Jan 1968
erect, becoming pendulous in fruit; peduncle 1.5- (d), Carauta553 (GUA); Mun. Rio de Janeiro,Estradada
20(-28) cm long, hispidulous to puberulousto sub- Vista Chinesa,km 2, 21 Oct 1968 (9 fl-fr), Carauta645 (E,
glabrous; spathe ca. 10-15 cm long, the color and F, GUA, MICH, SP, U); Mun. Petr6polis, Represa do Rio
indumentumas in the staminateinflorescence;spikes CaxambuPequeno, 19 Jul 1978 (), Carauta2976 (F, GUA,
4-9, 7-12 X 0.4-0.5 cm, to 30 x 1.2 cm in fruit, RB); rd. Barrado Pirai-Valen,a, 4 Mar 1980 (d), Carauta
sessile or with stipes to 0.5(-1.5) cm long and et al. 3425 (GUA); Mun. Rio de Janeiro,Grutade Imprensa,
sparsely puberulousto subglabrous;rachis hairy (or Sao Gabriel,3 Nov 1962 (9 fl-fr), Cuatrecasaset al. 26626
(US), (d), Cuatrecasaset al. 26627 (P, US); nr. Rio de Ja-
subglabrous).Pistillateflowers: perianthca. 1.5 mm
neiro, GCvea,19 Aug 1890 (5), Glaziou 18500 (C, G, K, P,
long, with arachnoid indumentumbelow the apex, US), (5), Glaziou 18501 (BG, C, G, K); Mun. Mage, Par-
also in the lower partof the style channel inside, the aiso, 9 Nov 1985 (d), Gomes 4 (RB); Angra dos Reis, 22
apex plane to slightly convex, muriculate;style rather Mar 1951 (d), M. Kuhlmann2673 (SP), (9 fl), M. Kuhl-
short;stigmacomose. Fruitellipsoid to oblongoid,ca. mann 2674 (SP). Rio GRANDE DO SUL: Picada Verao, 20
2-2.5 mm long, smooth, pale or darkbrown. Jan 1991 (9 fl-fr), Quadrosetal. 1679 (GUA), (5), Quadros
88
et al. 1680 (GUA);Os6rio,Maquine,22 Dec 1979 (d), Wae-
chter et al. 1516 (GUA).SANTA CATARINA:Mun.Brusque,
Mata do Hoffmann, 8 May 1951 (9 fr), Klein 113 (US);
Mun. Itapoa,ReservaVoltaVelha, 17 Feb 1993 (9 fr), Ne-
grelle et al. A.743 (GUA); Mun. Palhoqa,Morrodo Cavalo,
11 Mar 1953 (9 fl-fr), Reitz et al. 319 (GUA, US); Mun.
Itajaf,Morroda Fazenda,28 Apr 1954 (9 fl-fr), Reitz et al.
1797 (NY, US); Itajaf,Morro da Ressacada, 29 Mar 1956
(6), Reitz et al. 2902 (NY, US); Mun. Sao Franciscodo Sul,
Garuva, Tres Barras, 19 Dec 1957 (6), Reitz et al. 5736
(GUA); nr. Tubarao,May 1889 (9 fl-fr), Ule 1196 (P); Rio
Novo, Orleans, 14 Jan 1993 (9 fl-fr), Zanette et al. 1438
(GUA). SAOPAULO: Mun. Ubatuba,Rio Escuro, Rio Es-
curo, PraiaDura, 18 Dec 1979 (6), Benson 10837 (K, NY,
UEC); Campinas, 10 Jul 1968 (6), Carauta et al. 623
(GUA), (9 fl-fr), Carautaet al. 624 (C, GUA, U, US); Mun.
Iguap6,Pinheiro, 11 Jun 1982 (9 fl-fr), Hatschbach44969
(U, US); Sao Paulo, Botanical Garden,2 Sep 1949 (6), W
Hoehne et al. 3325 (GUA, US, SP, SPF) = M. Kuhlmann
3326 (SP); Jacarei,30 Sep 1949 (6), W Hoehne et al. (SPF)
12787 (SPF); Mun. Ubatuba,PraiaDura,26 Mar 1978 (6),
Joly 7349 (GUA, RB, UEC); Reserva Florestal das Fontes
do Ipiranga,7 Jun 1988 (9 fl-fr), Romaniuc 760, 15 Jan
1989 (6), Romaniuc787 (SP); Cubatao,BR.207, km 68, 14
Dec 1990 (d), Romaniucet al. 1015 (SP); Tapiraf,Serrade
Paranapicaba,10 Feb 1992 (9 fl-fr), Romaniucet al. 1273
(SP), (d), Romaniucet at. 1275 (SP); Santos, Mata da La-
goa, 2 May 1994 (9 fl-fr),Romaniucet al. 1396 (SP); Tim-
buri, Fazenda Dominiciana, 14 Jun 1995 (9 fl-fr), Tama-
shiro et al. 1255 (SP).
Cecropia glaziovii is a rather variable species in
terms of the length of the peduncle and (even on the
same tree) the numberand diameterof the spikes of
the staminateinflorescence.The peduncle of the pis-
tillate inflorescenceis mostly to ca. 10 cm long, some-
times to ca. 20 cm long, or occasionallyto 30 cm long
in fruit. The outer surface of the stipule is normally
rather sparsely puberulousto hirtellous, sometimes
densely hirtellous, or occasionally densely hirsute
(e.g., Berg et al. 1146 and Tamashiroet al. 1255).
The uncommonfeaturestend to occur in the periph-
ery of the known distributionrange of the species.
The leaves on the stem are often large (to ca. 100 X
100 cm) before branching and the lamina is often
more deeply incised (down to ca. 9/10, even down to
1.5-3 cm from the petiole); the free partof the mid-
segment has ca. 20-23 pairs of lateralveins, and the
arachnoidindumentis often sparseand (almost)con-
finedto the marginbeneath.Cecropiaglaziovii is very
closely relatedto C. palmata, showing distinct simi-
larities in the shape and indumentumof the leaves, in
both the pistillate and staminateinflorescences, and
in the staminateflowers. The two species appearto
be allopatric. More or less clear differences are as
follows: (1) The white arachnoid indumentum is
FLORA NEOTROPICA
sparse on the spathes in C. glaziovii, but dense on
those of C. palmata; (2) the stipules are mostly 15-
25 cm long and caducousin C. glaziovii, but 7-15 cm
long and mostly subpersistentin C. palmata; and (3)
the staminateinflorescencesof C. glaziovii often have
>4 spreadingto pendulous spikes, but those of C.
palmata normallyhave only 4 pendulousones. In ma-
terialwith thick spikes, long flowers,and stamens> 1
mm long, the detachedanthersremainattachedto the
filamentby stretchedspiral thickenings.However,it
is not clear what happensto the smaller anthers(>1
mm long) of short flowers (in more slender spikes);
they may remainattachedto the (for the greaterpart
? swollen) filament or become detached and re-
attachedby the appendages;see under C. palmata.
The collections Glaziou 18500-18507 are partly
mixed, comprising material of C. glaziovii and C.
pachystachya.
16. Cecropia goudotiana Trecul,Ann. Sci. Nat., ser.
3, 8: 83. 1847. Type. Colombia. Tolima:Piedras,
Jul 1844 (d), Goudot I (holotype:P). Fig. 23
RevistaAcad.Colomb.
CecropiaarbelaeziiCuatrecasas,
Ci. Exact.6(21): 65. 1944;Not. Fl. Colomb.VI.
(Trab.Col. Bot. etc. Colomb.)44. 1944.Type.Co-
lombia.Tolima:Curvasde Gualanday, betweenIba-
gue & Girardot,22 Jul 1939 (d), Cuatrecasas &
Perez Arbeldez 6503 (holotype: COL; isotypes: F,
US).
Tree, to 5(-10) m tall. Leafy twigs 1.5-2.5 cm
thick, darkgreen to brownish,hirtellousto subhispi-
dulous with curvedhairs.Laminachartaceous,ca. 25
X 25 cm to 35 X 35 cm (to 60 X 60 cm), the segments
6-7, the upper segments elliptic, the incisions in the
upper part of the lamina down to 5/10-6/10; apices
rounded; upper surface scabridulous (or scabrous),
somewhat bullate, densely puberulous to subhispi-
dulous;lower surfacedensely hirtellousto subtomen-
tose on the veins, the arachnoidindumentumnot in
the areoles, sparsely to ratherdensely covering the
hairs on the veins, largely disappearing, remain-
ing along the margin;lateralveins in the free partof
the midsegment 7-10 pairs, submarginally loop-
connected, branched;petiole 15-30(-50) cm long,
puberulousto hirtellous;trichiliafused, the brownin-
dumentum intermixed with rather long grayish to
white hairs; stipules 4-6 cm long, green, densely
grayish hirtellousto subtomentoseoutside, sericeous
inside. Staminate inflorescencesin pairs or solitary,
pendulous;peduncle6-11 cm long, hirtellousto sub-
setulose; spathe 10-16 cm long, green, densely hir-
tellous outside, sparselyhairy inside; spikes 4, 10-17
SYSTEMATIC TREATMENT 89

-%N~~~~~~~~~~~~~~~~~~~~~N

3~~~~~~~~~~~~

I mn ~ ~ 1mn

FIG. 23. Cecropiagoudotiana. 1. Leafy twig with stipules, lamina, and trichilium.2. Apex of lamina and venation.
3. Pair of staminateinflorescenceswith spathes and base of petiole with trichilium.4. Staminateinflorescencesat anthesis
and bases of petioles with trichilia. 5. Staminateflower and stamen (Franco et al. 4652). 6. Pairof pistillate inflorescences
with spathes and base of petiole with trichilum. 7. Pair of pistillate inflorescences after anthesis and base of petiole with
trichilium(Franco et al. 4650). 8. Pistillate flower. 9. Fruit (Franco et al. 4651).
90 FLORA NEOTROPICA

X 0.4-0.5 cm, with stipes 0.3-0.5 cm long and
densely puberulous;rachis sparsely hairy. Staminate
flowers: perianthtubular,1-1.5 mm long, glabrous,
the apex plane; filaments ? swollen; anthers0.5-0.6
mm long, short-appendiculate,remainingattachedto
the filament at anthesis (?). Pistillate inflorescences
in pairs or solitary,erect to deflexed, soon pendulous;
peduncle 4-12 cm long, hirtellous to subsetulose;
spathe 7-11 cm long, the color and indumentumas
in the staminateinflorescence;spikes (2-)4(-6), often
two by two fused at the base, 7-10 X 0.4-0.5 cm, to
15 X 0.7-0.8 cm in fruit, sessile; rachis (sparsely)
hairy, sometimes partly with arachnoid hairs, or
(sub)glabrous.Pistillate flowers: perianth 1-1.5 mm
long, with arachnoidhairs below the apex, absentin-
side, the apex ? convex to plane, glabrous and
smooth; style ratherlong; stigma comose. Fruit ob-
longoid, ca. 1.2-1.5 mm long, tuberculate.
Distribution (Fig. 11.3). Colombia,probablycon-
fined to Tolima, on rock escarpmentsin areas with
dry forest, at low elevations.
Specimensexamined.COLOMBIA.TOLIMA: Mun.
Piedras, rd. Piedras-Doima, km 5, 7 Feb 1995 (d fl-fr),
Franco et al. 4650 (BG, COL, HUA, LP, US); Mun. Vena-
dillo, rd. to Merida,7 Feb 1995 (Y fl-fr), Franco et al. 4651
(BG, COL, HUA, LP, US), (d), Franco et al. 4652 (BG,
COL, HUA, LP, US), (Y fl-fr), Franco et al. 4653 (BG,
COL, HUA, LP, US); Mun. Venadillo, Vrda. La Sierrita,
Finca El Cedro, 12 Aug 1980 ( fl-fr), Idroboet al. 10988
(COL); Mariquita,7 Jan 1918 (Y fl), Pennell 3635 (A).
Specimensexamined.FRENCHGUIANA:Montagne
The species is relativelyrarein its rangeof distri- de Trinite, 12 Feb 1984 (9 fl-fr), Granvilleet al. 6611(BG,
bution, the northernpartof the dry upperMagdalena CAY,G, P, U); HauteMarouini,RocherKoutou,16 Aug
valley; it is confinedto rocks and rocky escarpments. 1987 (st), Granvilleet al. 9319 (US), 17 Aug 1987 (6),
in the free part of the midsegment 9-10 pairs, mar-
ginally to submarginallyloop-connected,unbranched
or branched;petiole ca. 15-35 cm long, minutelypu-
berulous;trichilia fused, the brown indumentumin-
termixedwith 5-9 mm long white hairs;stipules 3-4
cm long, pink to dark red, puberulousto hirtellous
and with sparse arachnoid indumentum outside,
(sub)glabrous inside. Staminate inflorescences in
pairs, erect; peduncle 8-9 cm long, sparselypuberu-
lous; spathe 4-6 cm long, pale pink to yellow,
sparsely puberulousto hirtellous and with sparse to
ratherdense arachnoidindumentumoutside, glabrous
inside; spikes 6-9, 2-4.5 X 0.3-0.4 cm, with stipes
0.3-0.4 cm long and sparselyhairy;rachishairy.Sta-
minateflowers:perianthtubular,1-1.5 mm long, with
arachnoid indumentum below the apex, the apex
slightly convex to plane, glabrous;filamentsflat; an-
thers ca. 0.5-0.8 mm long, detachedat anthesis,reat-
tached to the marginsof the apertureby the append-
ages. Pistillate inflorescencesin pairs,erect;peduncle
6-10 cm long, hispidulous;spathe ca. 5-8 cm long,
the color and indumentumas in the staminateinflo-
rescence; spikes 4(-6), (at anthesis) 4-7 X 0.5-0.8
cm, sessile; rachis glabrous.Pistillate flowers: peri-
anth ca. 1.5 mm long, with arachnoidindumentum
below the apex, the apex muriculateto sparselymin-
utely puberulous;stigma subpeltate.Fruit unknown.
Distribution (Fig. 8.6). FrenchGuiana,on steep,
wet slopes of graniterocks (inselbergs).
Granvilleet al. 9392 (BG, INPA, P, U, US).

This species is knownfrom only two localities and

17. Cecropia granvilleana C. C. Berg, Bull. Mus. is found in small patches at bases of inselbergs. It
Hist. Nat. (Paris),ser.4, sect. B, Adansonia7: 255, shows affinitiesto Cecropiapeltata.
t. 1. 1985; Berg in Gorts-vanRijn, Fl. Guianas,
ser.A., fasc. 11: 97, t. 20. 1992. Type.FrenchGui-
ana. Montagnede Trinit6,18 Jan 1984 (6), Gran-
18. Cecropia herthae Diels, Notizbl. Bot. Gart.
ville et al. 6000 (holotype: P; isotypes: BG, BR,
Berlin-Dahlem15: 367. 1941. Type.Ecuador.Pas-
CAY,G, NY, U). Fig. 24
taza:Mera,ca. 1100 m, 3 Dec 1938 (9) Schultze-
Tree,to 10 m tall, the trunkwith prominentstip- Rhonhof3072 (holotype:B, destroyed,duplicates
ular scars. Leafy twigs 1.5-2 cm thick, green, puber- not traced);here replacedby: Ecuador.Napo: Re-
ulous to hispidulous,partlywith uncinatehairs.Lam- servaBiologica JatunSacha, 8 km E of Mishualli,
ina chartaceousto subcoriaceous,ca. 25 X 25 cm to 25 Dec 1990 (9), Neill & Berg 9743 (neotype:
40 X 40 cm, the segments9-10, oblong to lanceolate, QCNE; isoneotypes:BG, GB, MO, NY). Fig. 25
the incisions down to ca. 6/10-8/10; apices acute to Cecropia congesta Cuatrecasas, RevistaAcad.Colomb.
subacuminate;upper surface scabrous to scabridu- Ci. Exact. 6(22/23): 299. 1945. Type. Colombia.Pu-
lous, hispidulous;lower surface hirtellous to puber- tumayo:Rio San Miguel, betweenRio Conejo & Rio
ulous on the veins, with arachnoidindumentumin Hormiga, 15 Dec 1940 (D fr), Cuatrecasas 11069
the areoles and on the smaller veins; lateral veins (holotype:COL; isotype: F).
SYSTEMATICTREATMENT 91

0.5mm
I 9 3 f GS0mm

FIG. 24. Cecropia granvilleana. 1. Leafy twig with stipules, lamina, trichilia, and staminateinflorescences. 2. Sta-
minate flower and stamen (Granvilleet al. 6000). 3. Pistillate flower and stigma. 4. Fruitwith persistentstyle (Granvilleet
al. 6611). (By H. Rypkema.From Bull. Mus. Natl. Hist. Nat., Paris, s6r. 4, 7[Sect. B, Adansonia 3]: 256. 1985.)
92 FLORA NEOTROPICA

I~~~~~~~4

SS; X

2;!~~~~~~~~~~I

FIG. 25. Cecropia herthae. 1. Lamina, reduced. 2. Apex of lamina and venation. 3. Stipules and bases of petioles
with trichilia (Neill et al. 9743). 4. Staminateinflorescence with spathe. 5. Staminateinflorescence at anthesis (Zak et al.
4404). 6. Staminate flower and stamen (Neill et al. 9743A). 7. Pistillate inflorescence with spathe (Zak et al. 4471). 8.
Pistillate inflorescenceat anthesis and bases of petioles with trichilia. 9. Pistillate flower (Neill et al. 9743).
SYSTEMATICTREATMENT
Tree,to 30 m tall. Leafy twigs 2-8 cm thick, ini-
tially darkgreen with largepinkishlenticels, the older
parts often blackish, hispidulous.Laminacoriaceous
to subcoriaceous,ca. 60 X 60 cm to 80 X 80 cm (to
100 X 100 cm), the segments (10-)14-16, oblance-
olate, the incisions down to 1-2 cm from the petiole
or down to the petiole (and subpetiolulate);apices
(sub)acuminateto acute; upper surface scabrous to
scabridulousand hispidulous or smooth and subgla-
brous to sparsely puberulous (to hirtellous); lower
surfacepuberulous(with uncinateand straighthairs)
or partly subhirtellous,with arachnoidindumentum
confined to the areoles; lateral veins in the free part
of the midsegmentca. 30-40 pairs, marginallyloop-
connected, unbranched(or faintly branched);petiole
ca. 50-75(-l 10) cm long, puberulous (or adaxially
partly hirtellous) or also with sparse to ratherdense
arachnoid indumentum; trichilia fused (slightly
sunkeninto the base of the petiole), only with brown
pluricellularhairs; stipules 20-30 cm long, pink to
pale dull red, caducous or subpersistent,puberulous
and with sparseshortarachnoidindumentumoutside,
glabrousinside. Staminateinflorescencesin pairs,the
peduncle erect, the spikes erect to ? deflexed; pe-
duncle 3.5-9 cm long, hispidulous and with sparse
arachnoidindumentum;spathe (not mature)6-7 cm
long, sparsely minutely puberulousoutside, glabrous
inside; spikes ca. 50-100, (4-)8-14 X 0.2-0.4 cm,
with stipes 0.3-1 cm long andglabrousor puberulous;
rachis hairy. Staminateflowers: perianthtubular,ca.
1-1.2 mm long, glabrous, the apex slightly convex,
sparselymuriculate;filamentsflat;anthersca. 0.4 mm
long, appendiculate,detached at anthesis (?). Pistil-
late inflorescencesin pairs, erect (to ? spreading);
peduncle 3-4 cm long, hispidulousto hirtellous and
with sparsearachnoidindumentum;spatheca. 7-8 cm
long, the indumentumas in the staminate inflores-
cence; spikes (7-)9-12, (in fruit) 5-12 X 1.5-2 cm,
sessile or with stipes to 0.5 cm long and puberulous;
rachis glabrous.Pistillateflowers: perianthca. 2 mm
long, with arachnoid indumentumbelow the apex,
also below the style channel inside, the apex convex,
dome-shaped,smooth or muriculate,the apertureslit-
shaped; style short;stigma comose. Fruit oblongoid,
ca. 4 mm long, almost smooth, pink.
Distribution (Fig. 10.6). NorthernAmazonianEc-
uador and adjacentColombia (Putumayo)and Peru
(Loreto),in forest, at low elevations.
Specimens examined. COLOMBIA. AMAZONAS:
Mun. Leticia, Parque Nacional Amacayacu, Quebradade
Agua Pudre, 15 Nov 1991 (Y fl), Pipoly et al. 16272 (COL).
PUTUMAYO:Mun. Orito,Vrda.El Parafso,17 Feb 1995 (Y
fr), Franco et al. 5574 (BG, COL).
93
ECUADOR. MORONA-SANTIAGO: Pozo petrolero
Garza,ca. 35 km NE of Montalvo, 2-12 Jul 1985 (6), Zak
et al. 4404 (MO, QCNE), (Y fl), Zak et al. 4471 (BG, MO,
QCNE). NAPO: Rio Napo, nr. Santa Rosa, Rio Huambuno,
4 Mar 1990 (st), Berg et al. 1644 (BG, COL, QCA); Parque
Nacional Yasuni, Pozo petrolero Daimi 2, 26 May-8 Jun
1988 (Y fl-fr), Ceronet al. 4021 (BG, QCNE);Cant6nFran-
cisco Orellana, Huashito, 3-21 Nov 1989 (Y fr), Gudino
179 (BG, MO, QCNE); ParqueNacional Yasuni, rd. Pom-
peya Sur-Iro, km 38.7, 20 Oct 1994 (st), Jaramillo et al.
17110 (QCA); Cascadade San Rafael, 1400 m, 1 Aug 1975
(st), Little et al. 234A (Q); Reserva Biol6gica JatunSacha,
8 km E of Misahualli,26 Dec 1990 (d), Neill et al. 9743A
(BG), 8 Aug 1987 (st), Palacios 1774 (QAME, QCNE), 14
Aug 1987 (Y fl-fr), Palacios 1888 (BG, MO, QAME,
QCNE), 22 Sep 1989 (st), Palacios 4494 (BG, MO, QAME,
QCNE), 21 Oct 1989 (st), Palacios et al. 4684 (BG, QCNE).
PASTAZA: Cant6n Pastaza,Rio Lliquino, 1 km N of Pozo
Villano 2, 29 Aug 1997 (d), A. Alvarez et al. 2269 (BG).
SUCUMBiOS: Shushufindi,3 Aug 1975 (st), Little et al. 35
(Q, QCNE), 28 Dec 1974 (d), Vickers92 (F).
PERU. LORETO:Dtto. Indiana,ExploramaInn, 21-22
Apr 1992 (st), Grdndezet al. 4107 (BG), 4156 (BG).
The present description of the vegetative parts
largely matches the description of this species by
Diels (1941) and is in accordancewith his comment
that Cecropia herthae resembles C. sciadophylla.
However,there is a discrepancybetween the present
descriptionof the pistillate inflorescenceand Diels's
description of the peduncle as 12-15 cm long, his-
pidulous to hirtellous;spikes 5, 10-13 cm long; per-
ianth with arachnoidindumentumbelow the apex;
stigmapenicillate.At least the differencein the length
of the peduncle cannot be easily explained, if one
does not assume that the type collection was a mixed
collection. This species is, at first sight, very similar
to C. sciadophylla, in particularto the type with non-
petiolate leaf segments. However, C. herthae is
clearly distinct in the presence of trichilia.Both spe-
cies are also ecologically similar; they can become
tall trees and can be found side by side.
Vernacular name. Ecuador:guarumonegro (Pas-
taza).
19. C. heterochroma C. C. Berg & P. Franco,Novon
6: 246, t. 11. 1996. Type. Panama.Veraguas:Rd.
Escuela Agricola Alto Piedra-Rio Dos Bocas, km
10, 26 Jul 1974 (9), Croat 25880 (holotype:MO;
isotype: BG). Fig. 26
Tree, to 8 m tall. Leafy twigs 2-5 cm thick, green
or purplish, densely hirtellous with uncinate hairs.
Lamina chartaceous,ca. 35 X 35 cm to 75 x 75 cm,
green or purplishbeneath,the segments 6-8, the in-
94 FLORA NEOTROPICA

/ ~~~~~~~1W

I /0

FIG. 26. Cecropia heterochroma.1. Lamina, reduced. 2. Apex of lamina and venation (Nevers 4953). 3. Stipules
and base of petiole with trichilium (Berg 299). 4. Pair of staminateinflorescence at anthesis (Nevers 3752). 5. Staminate
flower. 6. Stamen (Berg et al. 393). 7. Pistillate inflorescence at anthesis and base of petiole with trichilium. 8. Pistillate
inflorescencewith spathe (Nevers 4953). 9. Pistillate flower (Herreraet al. 1763). 10. Fruitwith persistentstyle (Nee et al.
11004). (From Novon 6: 247. 1996, modified.)
SYSTEMATICTREATMENT
cisions down to 3/10-5/10; apices subacuminateto
rounded;uppersurface minutely hispidulous,scabri-
dulous; lower surface minutely puberulous with
curved hairs on the veins, the arachnoidindumentum
very sparse,mainly on the main veins andthe margin,
not in the areoles, soon disappearing;lateralveins in
the free partof the midsegment 10-12 pairs, submar-
ginally loop-connected,some of them branched;pet-
iole 30-55 cm long, green or purplish, puberulous,
partlyhirtellousto subhispidwith uncinatehairs;tri-
chilia fused, the brown indumentumintermixedwith
short white hairs; stipules 5-10 cm long, green or
reddish,subhirtellousto subhispidwith uncinatehairs
outside, glabrous inside. Staminateinflorescencesin
pairs, erect; peduncle 3-6.5 cm long, reddishor pur-
plish, hirtellousto subhirsuteto subhispid;spathe 8-
14 cm long, reddish, purplish,or greenish, sparsely
hirtellous and often with dense brown pluricellular
hairs outside, glabrous inside; spikes 4-5, 3-6.5 X
0.3-0.4 cm, with stipes to 0.5 cm long and sparsely
puberulous to glabrous; rachis glabrous. Staminate
flowers: perianthtubular,2-3 mm long, glabrous,the
apex 5-angularin circumference,adheringto apices
of adjacentperianths,slightly concave to plane, but
the margins of the aperturebent upward, glabrous;
filaments flat; anthers 0.6-0.8 mm long, oblong to
lanceolatein outline, appendiculateor not, remaining
attachedat anthesis.Pistillate inflorescencessolitary,
erect; peduncle 8-12 cm, red to purplish,puberulous
to hirtellous;spathe ca. 10-15 cm, the color and in-
dumentumas in the staminateones; spikes (l-)2-4,
3-10 X 0.4-0.6 cm, to 15 X ca. 1 cm in fruit,
(sub)sessile; rachis glabrous. Pistillate flowers:
perianth 1.5-2 mm long, with arachnoidhairs below
the apex, also below the style channelinside, the apex
slightly convex, glabrous, the aperturecircular,sur-
roundedwith a low rim; style ratherlong; stigma co-
mose. Fruit ellipsoid, ca. 2-3 mm long, smooth,
brown.
Distribution (Fig. 8.2). From eastern Panamato
Colombia,in forest marginsandin secondarygrowth,
at elevations to 1000 m.
Representative specimens examined. PANAMA. Bo-
CAS DEL TORO:Isla Bastimentos, 22 Mar1993 (9 fl-fr),
Foster et al. 14728 (SCZ). COLON:Santa Rita Ridge rd.,
between TransisthmianHwy. & Agua Clara, 11 Dec 1973
(9 fl-fr), Berg 299 (BG); SantaRita Ridge, 1 Mar 1971 (9
fl), Croat 13887 (BG, MO); Santa Rita Ridge rd., 4-6 km
from TransisthmianHwy., 13 Apr 1976 (6), Croat 34288 and on the smallerveins, often (initially) extendedto
(MO); SantaRita Ridge rd., 21-26 km from Transisthmian the main veins; lateral veins in the free part of the
Hwy., 4 Jul 1982 (9 fr), Knapp5844 (BG, MO). DARIEN:
ParqueNacional Darien, nr. Cruce de Mono, 5 Nov 1989 connected,unbranched;petiole 30-85(-100) cm long,
(juv), Fisher 52 (BG). PANAMA: Cerro Jefe, 30 Aug 1977 with sparseto dense arachnoidindumentumand fili-
95
(Y fl-fr),Berg et al. 393 (BG), (juv), Berg et al. 393A (BG),
(d), Berg et al. 394 (BG); El Llano-Cartird., km 7, 3 Sep
1977 (d), Berg et al. 401 (BG); Campo Tres, 5 km NE of
Altos de Pacora, 9 Mar 1973 (6), Busey 822 (BG, MO);
Cerro Jefe, 25 Aug 1972 (Y fr), Correa et al. 1806 (MO,
PMA); 3 mi N of Cerro Azul, 26 Jul 1970 (Y fl-fr), Croat
11587 (MO); El Llano-Cartfrd., km 8.7, 31 Oct 1977 (Y
fl), Folsom et al. 6161 (BG, MO); CerroJefe, 13 Sep 1970
( fl-fr), Foster et al. 1896 (PMA, US), 1 Jan 1972 (9 fl-
fr), Gentry et al. 3447 (MO, NY); Cerro Jefe, 29 Jul 1967
(9 fl-fr + 6), Kirkbrideet al. 21 (MO, NY); El Llano-Cartf
rd., km 18, 2 Mar 1975 (9 fl-fr), Mori et al. 5122 (BG,
MO); El Llano-Cartird., km 14, 28 Mar 1974 (9 fl), Nee
et al. 11004 (MO, NY, PMA). SAN BLAS:Nusagandi, 16
Aug 1989 (9 fl), Fisher 19 & 20 (BG); Rio Play6n, Chico,
27 Aug 1994 (9 fl), H. Herreraet al. 1763 (BG); Nusagandi,
NW of Punta Mama, 13 Aug 1984 (9 fl-fr), Nevers et al.
3731 (MO, PMA); El Llano-Cartird., continentaldivide, 25
Aug 1984 (6), Nevers 3752 (MO); El Llano-Cartfrd., km
19.1, 4 Mar 1985 (9 fl), Nevers et al. 4953 (MO, PMA); El
Llano-Cartfrd., 13 Mar 1986 (6), Nevers et al. 7378 (BG,
MO, PMA). VERAGUAS: Rio Dos Bocas, 15.6 km NW of
SantaFe, 31 Aug 1974 (6), Croat27627 (MO).
COLOMBIA. VALLE: Mun. Buenaventura,Bajo Cal-
ima region, rd. Buenaventura-Malaga,km 51.3, 9 Feb 1990
(6), Croat et al. 70406 (BG, MO).
This species has a morphwith the laminapurplish
underneathand a morph with the lamina pale green
underneath;they are found side by side. The species
is probably related to Cecropia obtusifolia.
20. Cecropia hispidissima Cuatrecasas, Revista
Acad. Colomb. Ci. Exact. 9(36/37): 325. 1956.
Type. Colombia.Valle:Rio San Juan,below Quer-
emal, 27 Mar 1947 (9), Cuatrecasas 23986 (ho-
lotype: US; isotype: F). Fig. 27
Tree, to 10(-25) m tall, often with a short trunk.
Leafy twigs 3-9 cm thick, green, setose with irritating
hairs, sometimes subsetoseto subvillous,or also with
arachnoidindumentum.Lamina coriaceous, ca. 40 X
40 cm to 85 X 85 cm (to 1 5 X 1 5 cm), the segments
(9-)10-13(-15), lanceolate,the incisions down to 2-5
cm from the petiole or to the petiole and then often
almost petiolulate; apices (sub)acuminateto acute;
uppersurfacesmooth or nearlyso, (sub)glabrous,the
"umbilicus"usually (sub)setose; lower surface with
brownpluricellularhairsor often also with arachnoid
indumentumand sometimes ? sparselysetose on the
main veins, with arachnoidindumentumin the areoles
midsegment 25-40(-44) pairs, submarginallyloop-
96 FLORA NEOTROPICA

\ 0.I,44.

4.1 C

;EgAt

venation1(Croat
1. Lamina,reduced.2. Apexof laminaand
FIG.27. Cecropia"hispidissima. 11588).3. Stipules,

1
FIGnoeta.278).CecrpiaPistiissteimaoesec amia reduesiDdso
2.3)Apex
ta. oflaia insvntionat (Croatr1158.3.Fr Stipurncoes,

al. 4552). (From Fl. Ecuador48: 26. 1993, modified.)

SYSTEMATICTREATMENT 97

form brown pluricellularhairs, usually also (partly) Cartird., km 5, 11 Nov 1973 (Y fl-fr),Nee 7949 (AAU, GH,
setose (with irritatinghairs) to hirsute, glabrescent; MO, PMA); above El Valle, rd. to La Mesa, 12 Feb 1974
trichilia fused, their margins often not well-defined, (st), Tyson6951 (PMA). SANBLAS:Rd. Nusagandi-Cartf,
17 Jul 1986 (5), McDonagh et al. 79 (BM); El Llano-Carti
sometimes extending to the base of the stipules or
rd., continental divide, 25 Aug 1984 (Y fl), Nevers 3754
almost lacking, the brown indumentumintermixed
(MO).
with irritatingbristles,the Muillerianbodies relatively COLOMBIA. ANTIOQUIA: Mun.Anorf, 12-16 km NW
large (ca. 3 mm) and pink; stipules 12-40 cm long, of Anorf, rd. to Dos Bocas, ca. 1000 m, 18 Nov 1989 (5),
darkred-brown,darkpurple-red(to blackish), green, Callejas et al. 8753 (HUA, NY); Mun. San Rafael, nr.Cam-
or sometimes whitish, densely subhispidto hirsuteto pamentoVersalles, 1350 m, 23 Mar 1994 (5), Franco et al.
substrigose (to subvillous) and with sparse to dense 4576 (BG, HUA); Mun. San Luis, Rio Claro, 25 Mar 1994
arachnoidindumentumoutside, glabrousinside. Sta- (Y fl-fr), Franco et al. 4586 (BG, HUA); Valdivia, 1250 m,
minate inflorescencesin pairs, erect; peduncle 8-12 27 Mar 1994 (Y fr), Franco et al. 4598 (BG, HUA); Mun.
Anorf,Vrda. El Roble, 1400 m, 8 Mar 1995 (5), Franco et
cm long, very densely hispid; spathe 5-14 cm long,
al. 5565 (BG, COL, HUA, LP); Mun. San Rafael,Vrda. La
yellowish green or reddishbrown to darkred to dark
Luz, Rio Nare, 1300 m, 10 Jun 1993 (5), D. Sdnchez S.
purple (to blackish), densely hispid to setose or to 1972 (MEDEL);valley of Rio Anorf,between Dos Bocas &
hirsute, sometimes subvillous and with dense arach- Anorf, 23 km W of Zaragoza,27 Aug 1976 (9 fl-fr), Shep-
noid indumentumoutside, glabrousinside; spikes ca. herd 596 (COL, MO); Mun. San Carlos, rd. Juanes-Alto
15-65, (3-)7-13 X (0.3-)0.7-1 cm, with stipes 0.3- Samana, km 12.6, 20 May 1988 (9 fl-fr), Zarucchi et al.
0.4 cm long and hispid; rachis hairy.Staminateflow- 6756 (BG, MO). CHOc6: Mun. San Jose del Palmar,ca. 1
ers sessile or short-pedicellate;perianthtubular,ca. km from San Jos6 del Palmar,1200 m, 19 Mar 1994 (9 fl-
2-2.5 mm long, glabrousor with arachnoidhairsbe- fr), Franco et al. 4569 (BG, HUA); 10 km W of Istmo de
San Pablo, W of Las Animas, 12 Jan 1979 (9 fl-fr), Gentry
low the apex, the apex slightly convex to plane;
et al. 24088 (BG, COL, HUA, MO); rd. Medellin-Quibd6,
filaments slightly swollen; anthers ca. 0.7-0.8 mm
ca. 20 km W from top of divide of Cord. Occidental,900-
long, appendiculate,detached at anthesis, reattached 1000 m, 3 Jan 1981 (9 fl-fr), Gentry et al. 30051 (COL,
to the marginsof the apertureby the appendages(?). MO); nr. San Jos6 del Palmar,1380 m, 11 Nov 1985 (9 fl-
Pistillate inflorescences in pairs, erect; peduncle 6- fr), Lozano C. et al. 4907 (COL, MO). NARINO:Nr. Alta-
18 cm long, densely setose to hirsute (to subvillous) quer, 1150 m, 21 Feb 1995 (9 fl-fr), Franco et al. 4697
and with dense arachnoidindumentum,glabrescent; (COL, HUA). RISARALDA: Mun. Mistrato,rd. to Puertode
spathe ca. 4-13 cm long, the color and indumentum Oro, Rio Currumai,29 May 1990 (9 fr), Orozcoet al. 2181
as in the staminateinflorescence;spikes 6-20, 3-12 (COL, MO). VALLE:Rio San Juan,below Queremal,1300-
1500 m, 27 Mar 1947 (5), Cuatrecasas23987 (K, US); nr.
x ca. 0.4 cm, to 19 X 1-1.5 cm in fruit,(sub)sessile;
Buenaventura,15 Mar 1994 (5), Franco et al. 4535 (BG);
rachis(sparsely)hairy.Pistillateflowers:perianth,ca. Mun. Buenaventura,rd. to Sabaletas,km 22, 17 Mar 1994
2.5 mm long, with arachnoidindumentumbelow the (9 fl-fr), Franco et al. 4552 (BG, HUA), km 24, (9 fl-fr),
apex outside, absentinside, the apex slightly convex, Franco et al. 4554 (BG, HUA); Rio Anchiyaca, nr. CVC
muriculate or glabrous; style long, straight;stigma hydroelectricplant, 15 Dec 1981 (9 fl-fr), Gentry 35671
penicillate. Fruit ellipsoid to ovoid, ca. 1.2-1.5 mm (BG, COL, MO).
long, ? tuberculate. ECUADOR. AZUAY:Rd. La Troncal-Zhud, Manta
Real, Rio Patul, 1200 m, 11-12 Jul 1991 (9 fl-fr), Yanezet
Distribution (Fig. 10.5). FromeasternPanamato al. 300 (QCA). CANAR:Rd. La Troncal-Suscal,ca. 20 km,
western Ecuador, in forest or secondary growth, at 3 Feb 1981 (5), Berg 1239 (AAU, BG, COL, GB, MO,
elevations to 1500 m. QCA, U). COTOPAXI: Rd. Quevedo-Latacunga,Tenefuerte,
Rio Pilal6, 750-1300 m, 21 Feb 1982 (5), Dodson et al.
Representative specimens examined. PANAMA. 12769 (MO, QCNE). EL ORO:Rd. Pifias-SantaRosa, km
COLON:Rio Escandaloso,nr.Mina Boquer6nNumero2, 14 11, 8 Nov 1979 (st), Dodson et al. 9005 (MO). ESMERAL-
Jul 1979 (d), Antonio 1309 (MO); Santa Rita Ridge rd., DAS:Cant6nQuininde,Bilsa Biological Station,40 km NW
between TransisthmianHwy. & Agua Clara, 11 Dec 1973 of Quininde, 12 Mar 1997 (st), Clarket al. 4089 (BG). Los
( Y fl-fr),Berg 298 (BG); nr.Rio Piedras,rd. to PuertoBello, Rios/PICHINCHA: El Centinela,at crest of Montafiasde Ila,
16 Jul 1966 (d), Blum 2530 (MO); Salud, 2 Aug 1971 (Y rd. PatriciaPilar-24 de Mayo, km 12, 27 Nov 1978 (juv),
fr), Lao 181 (MO, PMA);Pipelinerd., 8 km NW of Gamboa, Dodson 7303 (BG, MO, QCNE, SELBY). PICHINCHA: Rd.
26 Aug 1973 (D fl-fr), Nee 6643 (AAU, MO, NY, PMA). Quito-PuertoQuito, ca. 93 km fromNanegalito,7 Feb 1982
PANAMA: El Llano-Cartird., 3 Sep 1977 (Y fl-fr), Berg et (5), Berg et al. 1682 (AAU, BG, COL, QCNE); rd. Santo
al. 402 (BG); Cerro Jefe, 27 Jan 1966 (Y fl), Blum et al. Domingo de los Colorados-Quevedo,km 21, 30 Jan 1979
2099 (MO); CerroAzul, 3 mi N of CerroAzul (vill.), 26 Jul (9 fl-fr), Dodson et al. 7413 (MO, QCNE, U); ReservaFlo-
1970 (5), Croat 11588 (MO, NY); CerroJefe, ca. 1000 m, restal ENDESA, rd. Quito-Puerto Quito, km 113, 10 km
22 Sep 1972 (D fl-fr), Gentry 6149 (GH, MO); El Llano- from rd., 22-23 Nov 1984 (juv), Jaramillo7436 (AAU, GB,
98
MO, NY, QCA); Cant6nPedroVicenteMaldonado,Reserva
Rio Silanche, Nov 1996 (st), Penningtonet al. 15230 (BG).
The species is very closely related to Cecropia
garciae, as discussed under the latter. This species
shows a variationin the trichilia from almost absent
to even extended to the base of the stipules. In Valle
(Colombia),a lowland and a submontanetype appear
to be present, probably morphologicallydistinct by
different color of the stipules (green vs. dark red to
[dark]purplish).The stipulesareoccasionallywhitish
due to dense arachnoidindumentum.The character-
istic stiff (setose) hairs are absentin some collections
made in Choco and Valle, being replacedby weaker
hairs providing a subvillous indumentum.Ants are
often absent, or if present, then a species of Pachy-
condyla. These ants perforatethe node bearing the
stipules formingthe apicalbud cover andthey harvest
the (large and pink) Muillerianbodies when still en-
closed in the bud.
21. Cecropia hololeuca Miquel, in Martius,Fl. Bras.
4(1): 148. 1853; Berg & Carauta,Albertoa 1(1):
6. 1986. Ambaibahololeuca (Miquel)Kuntze,Re-
vis. Gen. P1. 2: 624. 1891. Type. Brazil. Without
locality, Pohl s.n. (holotype: W, destroyed),neo-
type here designated:Brazil. Rio de Janeiro:Mun.
Rio de Janeiro,Corcovado, Aug 1890 (Y fl-fr),
Glaziou 18497 (IAN; isoneotypes: A, BM, C, E,
G, K, NY, P).
Cecropiacandida Snethlage,Notizbl. Bot. Gart.Berlin-
Dahlem 8: 367. 1923. Type. Brazil. Rio de Janeiro,
Serra de Macae, Ule 4863 (holotype: B, destroyed,
photographex B in MO).
Tree,to 25 m tall. Leafy twigs 2-8 cm thick, green
or slightly bluish to brown, glabrous, hirsute to vil-
lous, and/or with arachnoidindumentumor some-
times with only brown pluricellularhairs. Lamina
coriaceous, ca. 40 X 40 cm to 120 X 120 cm, the
segments 8-10, the free parts of the upper segments
oblong to subobovate to oblanceolate, the incisions
down to 7/10-9/10; apices roundedto obtuseto short-
acuminate; upper surface glabrous, with ? dense
arachnoidindumentum;lower surfacewith arachnoid
indumentumin the areoles and on the smaller veins,
or also on the main veins; lateralveins in the free part
of the midsegment 12-15 pairs, submarginallyloop-
connected,branchedor unbranched;petiole 30-95 cm
long, glabrous, with arachnoidindumentumor also
hirsuteto subvillous at the base, or only with brown
pluricellularhairs;trichiliaabsent;stipules 10-40 cm
long, yellowish white to brownish,(partly)pale yel-
FLORANEOTROPICA
low to white-sericeous or -villous or also (or only)
with dense brown pluricellular hairs and/or with
(sometimes dense) arachnoid indumentumoutside,
densely sericeous inside. Staminate inflorescences in
pairs, erect; peduncle 5-12 cm long, red to purplish,
glabrousor (sub)villousat the base or also at the apex,
with dense brown pluricellularhairs and/or sparse
arachnoidindumentum;spathe lacking or with only
bractson the upperpartof the peduncle,these to 0.5
cm long, caducous; spikes 9-17, 5-15 X 0.5-1 cm,
often ? moniliform, wine-red to dark purple to al-
most black, with stipes 0.8-2.5 cm long andglabrous;
rachis hairy, with straight hairs or also arachnoid
ones. Staminateflowers: perianthtubular,ca. 2 mm
long, with short and stiff hairs in the ribes below
the apex or glabrous, the apex almost plane; fil-
aments flat; anthers ca. 0.6-0.8 mm long, short-
appendiculate,detachedand reattachedto marginsof
the apertureat anthesis (?). Pistillate inforescences
in pairs, erect, pendulousin fruit;peduncle 5-17 cm
long, glabrous or (sub)villous, red(dish); spathe ab-
sent, but with 1 or 2 bracts on the upper part or the
apex of the peduncle, these to 1.5(-8) cm long, ca-
ducous;spikes 1 or 2 (rarely3), 5-12 X 0.5-1 cm, to
35 x 3 cm in fruit, initially red to purplish,turning
blackish, sessile or with stipes to 1.5 cm long and
glabrousor hairy;rachishairy.Pistillateflowers: per-
ianthca. 2-2.5 mm long, with arachnoidindumentum
below the apex outside, also in the lower part of the
style channelinside, the apex ? convex, muriculate;
style ratherlong, straight;stigmapenicillate.Fruitel-
lipsoid to oblongoid,ca. 3-4 mm long, ? tuberculate,
darkbrown.
Distribution (Fig. 8.6). Eastern Brazil, in moist
forest, in mountainousand hilly regions, at elevations
to 1300 m.
Representativespecimensexamined.BRAZIL.BA-
HIA: Rd.fromRd.Sao Jose-Buerarema
to Una, km 6, 1 Apr
1980 (Y fl-fr),Berg et al. 1142 (BG,K, NY, U); rd. Itabela-
Guaratinga,km 20-25, 31 Mar 1989 (st), MattosSilva et al.
2674 (G); Santa Cruz de Cabralia,Reserva Biol6gica do
Pau-brasil, 16 Sep 1971 (Y fl-fr), T S. Santos 1950 (U).
ESPiRITO SANTO: SantaTeresa,21 Nov 1988 (6), Bausen
s.n. (GUA); Linhares,ReservaFlorestalde CVRD, Est. Fla-
mengo, 10 Feb 1981 (6), Folli 310 (BG,CRVD,GUA, MG).
MINAs GERAIS: Juiz de Fora, Oct 1934 (6), Brade14130
(RB); Viqosa, FazendaCriciuma,9 Sep 1967 (6 + Y fl-fr),
Carauta 404 (GUA, K, RB, US); rd. BR.262, Mun. Man-
huacu, 4 Dec 1984 (9 fl-fr), Hatschbachet al. 48598 (BG,
C, MO); 5-10 km N of Teofilo Otoni, 15 Jul 1988 (9 fr),
Hatschbach et al. 52174 (BG); Esta,co ExperimentalCo-
ronel Pacheco, 11 Dec 1944 (st), Heringer 1685 (US); Mun.
Marliera,ParqueEstadualdo Rio Doce, 18 Sep 1975 (6),
Heringeret al. 15054(US), 19 Sep 1975(9 fr),Heringer
SYSTEMATICTREATMENT
et al. 15057 (BG, MO, US); Carangola, 1 Sep 1990 (5),
Leoni 1218 (GUA), (9 fl-fr), Leoni 1219 (GUA);Viqosa,24
Feb 1930 (9 fl-fr),Mexia 4389 (BM, F, G, GB, K, MO, NY,
S, U); Caldas, Sep 1867 (9 fl-fr), Regnell III.1100 (S); Ri-
berao, Rio Novo, Sep 1894 (d), in herb. Schwacke 11128
(RB). RiO DE JANEIRO: Mun. Rio de Janeiro,Estradada
Vista Chinesa, 13 Mar 1963 (5), Angeli 350 (GUA, MG,
RB, US); Cachoeiras do Macacu, Morro do Reservat6rio,
28 Feb 1968 (5), Carauta 581 (GUA, U, US); Rio de Ja-
neiro, Estradado Sumare, 15 May 1968 (9 fl-fr), Carauta
601 (GUA); Rio de Janeiro,Pedra de Gavea, 26 Feb 1977
(5), Carauta et al. 2328 (RB); Campos, Morro do Coco,
Cruz da Serra,Morrodo Bai, 13 Dec 1982 (d), Carautaet
al. 4423 (GUA); Duas Barras, 13 Dec 1088 (d), Carautaet
al. 5683 (GUA); Barra Mansa, Ribeirao de Sao Joaquim,
Fazenda Campo Alegre, 18 Apr 1989 (5), Carauta et al.
5816 (GUA); Resende, Rio Palmital,22 Feb 1966 (st), Cas-
tellanos 25695 (GUA); Corcovado,Tijuca,21 Jan 1891 (5),
Glaziou 18498 (A, C, G, IAN, K, P); Larangeiras,7 Sep
1890 (9 fr), Glaziou s.n. (P); Rio de Janeiro,Corcovado,
Sumare, 20 Apr 1922 (5), Kuhlmann(RB) 3664 (RB, U);
Rio de Janeiro,Horto Florestal, 20 Feb 1929 (9 fl), Kuhl-
mann (RB) 111679 (GUA, RB, U); Rio de Janeiro,Estrada
do Redentor,Tijuca, 1939 (9 fl-fr), Kuhlmann(RB) 136990
(RB, U); Rio de Janeiro,Corcovado,Sumare,Jun 1928 (5),
Kuhlmann(RB) 141261 (RB), (9 fl-fr), Kuhlmann(RB)
141262 (RB); Mun. Mage, Paraiso, 2 Oct 1984 (9 fl-fr),
Martinelli et al. 9908 (GUA, K, NY, RB). SAO PAULO:
Mun. SerraNegra, Alto da Serra,ca. 1000 m, 22 Nov 1991
(9 fl-fr), Barros et al. 2385 (SP); Represa do Jaguari,31
Dec 1979 (9 fl-fr),Benson 10878 (UEC);Aruja,12 Jul 1981
(9 fl-fr), Custodio Filho 654 (SP); Sao Jose dos Campos,
30 Aug 1949 (d), W Hoehne et al. 2004 (SP, SPF, US), (9
fl-fr), W Hoehne et al. 2005 (SP, SPF); Ibiti,27 Jul 1943 (9
fl-fr), M. Kuhlmann1326 (SP); Mun. Santa Izabel, Igarata,
27 Sep 1950 (9 fl-fr), M. Kuhlmann2567 (NY, SP); Cam-
pinas, 8 Jul 1988 (st), Matthes2 (GUA); BraganqaPaulista,
FazendaSan Antonio, 21 Sep 1991 (9 fl-fr), Mello-Silva et
al. 547 (SP, SPF); Mogi das Cruzes, Rodovia dos Trabal-
hadores,22 Apr 1992 (5), Romaniucet al. 1300 (SP); Mun.
Amparo, rd. Amparo-Pedreira,5 Apr 1993 (st), Romaniuc
et al. 1366 (SP).
Cecropia hololeuca is the only species of the ge-
nus lacking the characteristic spathe, fully enclosing
the spikes before anthesis. Insteadof the spathe, one
or sometimes two bractsare found at the apex or just
below the apex of the peduncle. These bracts are
mostly to 1.5 cm long, sometimes to 8 cm long. The
inflorescence remains enclosed in the apical bud until
anthesis. This species resembles several Andean mon-
tane species in the dense white arachnoid indumen-
tum on the upper surface of the lamina and some of
them in the absence of trichilia and/or the presence
of either smooth or densely hairy leafy twigs.
Vernacular names. Brazil: imbauibabranca(Ba-
hia, Espirito Santo, Minas Gerais); imbauibavermelha
99
(Minas Gerais); embaubaSui,imbaubuqu(Rio de Ja-
neiro); embautbabranca(Sao Paulo).
22. Cecropia idroboi Cuatrecasas, Revista Acad.
Colomb.Ci. Exact.9(36/37): 335. 1956. Type.Co-
lombia.Vaupes:Mesa La Lindosa, 15-20 km S of
San Jose del Guaviare, 13-15 Dec 1950 (Y), Id-
robo & Schultes662 (holotype:US; isotypes:BM,
COL, GH, MO, NY, U, US).
Tree,to 10 m tall. Leafy twigs 1.5-2.5 cm thick,
hispidulous to puberulous.Lamina (sub)coriaceous,
ca. 15 X 15 cm to 40 X 40 cm, the segments 7-8,
the free partsof uppersegments (broadly)obovateto
elliptic, the incisions down to 3/10-5/10; apices
roundedor apiculate;uppersurfacesmoothto scabri-
dulous, sparselyhispidulousto strigillose; lower sur-
face puberulous on the main veins or also on the
smaller veins, with arachnoidindumentumin the ar-
eoles or also sparsely to densely on the lateralveins
or also on the main veins, sometimes the whole sur-
face with dense arachnoidindumentum;lateralveins
in the free part of the midsegment 8-10(-12) pairs,
marginallyloop-connected, most of them branched;
petiole ca. 10-40 cm long, sparselypuberulousor also
with dense arachnoidindumentum;trichiliafused, the
brown indumentumintermixedwith sparse to rather
dense short brownish or whitish unicellular hairs,
sometimes with dense arachnoidindumentum;stip-
ules 3-7 cm long, dark red (?), sparsely puberulous
to hirtellous, sometimes with dense arachnoidindu-
mentum outside, hairy inside. Staminate inflores-
cences in pairs or solitary,erect (?); peduncle6-8 cm
long, hispidulous; spathe 5-6 cm long, dark red,
sparsely puberulous (to subglabrous) outside, gla-
brousinside; spikes ca. 15, 1.5-5 X 0.2-0.3 cm, with
stipes to 0.3 cm long and densely puberulous;rachis
hairy. Staminateflowers: perianthtubular,0.8-1 mm
long, with sparse arachnoidindumentumbelow the
apex, the apex almost plane, glabrous;anthers0.5-
0.6 cm long, detachedat anthesis (?). Pistillate info-
rescences in pairs, erect (or deflexed in fruit?);pe-
duncle 3-6.5 cm long, hirtellousto puberulousor also
with ? dense arachnoidindumentum;spathe4-5 cm
long, the color and indumentumas in the staminate
inflorescence;spikes 4-5, 3-6 X ca. 0.8 cm, to 9 x
ca. 1.3 cm in fruit,(sub)sessile.Pistillateflowers:per-
ianth 1.5-2 mm long, with arachnoidindumentumbe-
low the apex, also in the lower partof the style chan-
nel inside, the apex convex, muriculateto granulate,
the apertureslit-shaped;style long, S-shaped;stigma
comose. Fruit oblongoid, ca. 2 mm long, smooth,
darkbrown.
100
Distribution (Fig. 10.6). Colombia (Caqueta,
Meta, and Vaupes), on sandstone rocks, at low ele-
vations.
Specimensexamined.COLOMBIA. CAQUETA:Sierra
de Chiribiquete,7 Dec 1990 (Y), Castroviejoet al. 11978
(COL).META: LaMacarena, RioGuayabero,Jan-Mar1959
(? fl-fr), Garcia-Barrigaet al. 17053 (COL,NY, US).
VAUPES: Rio Kuduyari, CerroYapoboda,5 Oct 1951(Y fl-
fr),Schulteset al. 14240(COL,US), 5-6 Oct 1951(Y fl-
fr),Schultes14370(COL,GH,US), 17 Nov 1952(Y fl-fr),
Schultes et al. 18500 (BG, K, MO, NY), 9 Oct 1966 (5),
Schultes 24272 (COL).
This species shows strongmorphologicalaffinities
to Cecropiadistachya;these two taxa might prove to
be conspecific. Cecropia idroboi differs from C. dis-
tachya in the less deeply incised lamina, the smaller
numberof lateralveins in the free partof the midseg-
ment, and the shorterspikes of the pistillate inflores-
cence. The trees remainsmallerthanon averagein C.
distachya. Cecropia idroboi is probablyecologically
distinct, as a species growing on rocks or in rocky
places. However, specimens which could be referred
to C. distachyawithoutdoubt,have been collected in
the same areaandin (almost)similarhabitats.Garcfa-
Barriga 17053 matches C. idroboi in most features,
but it is quite distinct in the dense arachnoidindu-
mentum on the petiole, lower surface of the lamina,
and the peduncle, and in this respect similar to C.
metensis, found in the same locality.
23. Cecropia insignis Liebmann,Kongel. DanskeVi-
dens. Selsk. Naturvidensk.Math Afh., ser. 5, 2:
281. 1851; Burger,Fieldiana,Bot. 40: 125, t. 23.
1977; Croat,Fl. BarroColoradoIsland346, t. 194.
1978; Berg & FrancoRosselli, Fl. Ecuador48: 29.
1993. Type. Nicaragua.Rio San Juan,Oersteds.n.
(type material not yet traced. Oersted 5806 (C),
also from Rio San Juan does not match Lieb-
mann's description and belongs to C. peltata),
herewith replaced by: Nicaragua. Rio San Juan:
Nr. Caiio Chontaleno,20 km NE of El Castillo, 7-
9 Mar 1978 (9), Neill 3392 (neotype: MO; iso-
neotype: BG). Fig. 28
CecropiaeximiaCuatrecasas,RevistaAcad. Colomb.Ci.
Exact. 6(22/23): 287. 1945. Type. Colombia. Valle:
Rio Calima, La Trojita, 19 Feb-10 Mar 1944 (Y),
Cuatrecasas 16830 (holotype: COL; isotypes: F,
US).
Cecropia sandersonianaP. H. Allen, The rain forest ofthe upper part of the style channel inside, the apex
Golfo Dulce 162 & 409 (as C. standleyana). 1956. convex or almost plane, (sub)muriculate,? angular
Nr.
Type.CostaRica.Puntarenas: Golfito, 4 Jan 1953 in circumference,the apical part long, the aperture
(9), Allen 6688 (holotype:EAP; isotype: US).
FLORANEOTROPICA
Cecropia polyandrophoraCuatrecasas,Phytologia 20:
465. 1971. Type. Colombia. Antioquia: between
Villa Artega & Chigorod6,La Pradera,3 Oct 1961
(i), Cuatrecasas & Willard26187 (holotype: US;
isotypes: COL, US).
Tree,to 25(-40) m tall. Leafy twigs 2-5 cm thick,
reddish-brownto blackish, (rather)sparsely hispidu-
lous. Lamina coriaceous (and sometimes ? plicate)
to subcoriaceous,ca. 50 X 35 cm to 100 X 85 cm (to
120 X 100 cm), the segments 7-8(-9), the free parts
of uppersegments elliptic to (sub)obovateto oblong,
the incisions down to ca. 7/10; apices obtuse; upper
surface smooth, initially with sparse to ratherdense
arachnoidindumentum;lower surface with sparseto
dense brown pluricellularhairs on the main veins,
with arachnoidindumentumin the areoles, on the
smallerveins, and (at least initially) also on the main
veins; lateral veins in the free part of the midseg-
ment, 12-18(-24) pairs, submarginally loop-
connected, unbranched;petiole 35-80 cm long, red-
brown to grayish, with dense brown pluricellular
hairs and dense arachnoid indumentum; trichilia
fused or separate,the brownindumentumintermixed
with short stiff white to brownish(unicellular)hairs;
stipules 22-50 cm long, pinkish to red to red-brown,
caducous (or subpersistent),subsericeous to hirtel-
lous to hispidulous and with dense brown pluricel-
lular hairs outside, + densely sericeous inside. Sta-
minate inflorescences in pairs, erect; peduncle 3-7
cm long, hispidulousto puberulous;spathe 12-16(-
22) cm long, pinkish to red to red-brown,puberulous
to hispidulous or to strigillose or also brown pluri-
cellular hairs and/or (rather)sparse arachnoidindu-
mentum outside, sparsely to densely hairy or gla-
brous inside; spikes 5-9 and 8-10 X 0.6-1.2 cm or
ca. 15-35 and 6.5-14.5 X 0.3-0.5 cm, with stipes
0.5-2 cm long and glabrous;rachis hairy. Staminate
flowers: perianth tubular,ca. 1.5-2 mm long, with
short and stiff hairs below the apex, the apex almost
plane; filamentsflat;anthersca. 0.7 mm long, appen-
diculate, detached at anthesis (?). Pistillate inflores-
cences in pairs or solitary, initially erect, pendulous
in fruit; peduncle 9-15 cm long, green, hispid or
subglabrous;spathe 10-22 cm long, the color and in-
dumentumas in the staminateinflorescence;spikes
(2-)4-5(-7), 8-18 X ca. 0.8 cm, to 26 X ca. 1 cm in
fruit, with stipes 0.5-1 cm long and glabrous;rachis
hairy. Pistillate flowers: perianth ca. 2 mm long,
with arachnoidindumentumbelow the apex, also in
slit-shapedto circular,borderedby a rim; style long,
SYSTEMATICTREATMENT 101

gi~~~~~~~~~ Tn .+

5crn.

FIG. 28. Cecropia insignis. 1. Lamina, reduced. 2. Apex of lamina and venation (Berg 1240). 3. Stipules, young
leaf, and base of petiole with trichilium(Berg 1260). 4. Staminateinflorescence with spathe. 5. Staminateinflorescenceat
anthesis. 6. Staminateflower and stamen (Berg 1240). 7. Pistillate inflorescences, one with spathe, the other at anthesis,
and base of petiole with trichilium(Berg 1260). 8. Pistillate flower.9. Fruit(Franco et al. 5560).
102 FLORA NEOTROPICA

curved; stigma truncate,comose. Fruit oblongoid to Blanquita,Murriregion, 4 Mar 1995 (9 fl-fr), Franco et al.
ellipsoid, 2-2.5 mm long, ? tuberculate. 5560 (BG, COL, HUA, LP, MO); Mun. San Luis, Quebrada
La Cristalina, 26 Feb 1987 (9 fl-fr), Ramirez et al. 691
Distribution (Fig. 10.6). From Hondurasto the (COL);valley of Rio Anorf,23 km W of Zaragoza,between
Pacific coastal region of Ecuador,and the Magdalena Dos Bocas & Anorn,31 Aug 1976 (st), Shepherd606 (COL,
valley (up to Tolima)and the valley of Rio Porce and HUA); Mun. Zaragoza,Cgto. Providencia,9 Feb 1971 (9
Rio Medellin to near Medellin, in (treefall gaps in) fl-fr), Soejarto et al. 2661 (GH, HUA); Mun. San Carlos,
primaryuplandforest, riverineforest, and in second- Cgto. El Jordan,TrochaLos Planes, 9 Apr 1991 (9 fr), Ma.
ary growth,at elevations to 1000 m, in Costa Rica to Veldsquezet al. 293 (HUA). BOYACA:Mun. PuertoBoyaca,
1400 m. Vrda. Puerto Pinz6n, 17 Jan 1998 (st), Cortes et al. 1546
(UDBC). CHOCO:Colombia-Panama border, Alto del
Representative specimens examined. HONDURAS. Lim6, 23 Sep 1979 (9 fl-fr), Barbosa 1191 (COL), (d),
OLANCHO: Between Campamento& borderOlancho/Mor- Barbosa 1194 (COL);Mun. Riosucio, Urabaregion, Cerros
azan, rd. Tegucigalpaca-Juticalpa,ca. km 100-101, 27 Oct del Cuchillo, 18 Nov 1987 (9 fl), Cdrdenas897 (JAUM,
1996 (Y), Berg 1809 (EAP). MO); nr. Bahia Solano, 4 Apr 1990 (9 fl-fr), Franco et al.
NICARAGUA. BLUEFIELDS:Base of CerroSan Isidro, 3006 (COL);Mun. San Jose del Palmar,1-2 km W of San
Rfo Kama-RioEscondido, 25 Mar 1966 (d), Proctor et al. Jos6 del Palmar,19 Mar 1994 (d), Franco et al. 4567 (BG,
27245 (NY). JINOTEGA: Las Brisas, ComarcaKilambe,930 COL, HUA), (9 fl), Franco et al. 4568 (BG, COL, HUA);
m, 13 Jul 1980 (Y fr),Sandino 168 (BG,MO).ZELAYA: S Nuquf, Alto de Buey, 1000 m, 23 Jun 1940 (9 fl-fr), Snei-
of Rio Wawa,60 km NW of PuertoCabezas, 17 Mar 1972 dern A.61 (A, COL, K, LL, MEDEL, MICH, NY, S, US);
(? fl), Little25201 (MO, US); nr.Rfo Wawa,60 km NW of Rio Jurad6,4 Oct 1940 (9 fl-fr), SneidernA.245 (A, COL,
PuertoCabezas, 27 Mar 1971 (5), Little 25340 (MO, US); NY); ParqueNacional NaturalLos Katios, Alto del Lim6n,
20 km W of Awas Tingni, S of Rio Wawa, 1 Apr 1971 (Y 9 May 1983 (9 fl-fr), Zuluaga R. 682 (COL). TOLIMA:
fl), Little et al. 25356 (MO); 20 km NW of Alamicamba,16 Mariquita,17 May 1990 (9 fl-fr),Francoet al. 2965 (COL),
Apr 1971 (st), Little 25382 (MO, US); new rd. Siuna-Ma- 7 Feb 1995 (9 fl-fr), Franco et al. 4654 (BG, COL, HUA).
tagalpa,ca. 10.1 km beyond Rio Uli, nr.Wani, 6 May 1978 VALLE: Between El Aguacate & Quebradade la Yuca, 8
(Y fl-fr),Stevens 8813 (MO). Feb 1944 (9 fl), Cuatrecasas16096 (F, US, VALLE);Rio
COSTA RICA. CARTAGO:Turrialba,( Y fl), Blum2218
Calima,Aguaclara,23 May 1946 (9 fr), Cuatrecasas21262
(MO); 0.5 km NE of Tapanti,1360 m, 11 Jun 1967 (Y fl),
(F, US, VALLE);Rio Calima, La Brea, 25 May 1946 (st),
Lent 1031 (MO, NY); 0.5 km S of PeniasBlancas de Cachi,
Cuatrecasas 21315 (US); Rio Anchicaya, Rio Blanco, 29
1400 m, 5 Sep 1971 (Y fr), Lent2119 (MO, NY); Turrialba,
Mar 1949 (9 fl-fr), Cuatrecasas24009 (US), (d), Cuatre-
1100 m, 7 Jul 1992 (Y fl-fr), Rivera 1933 (K). PUNTAR-
casas 24010 (F); Mun. Cisneros, 15 Mar 1994 (d), Franco
ENAS:Finca Loma Linda, 1 mi SW of CafnasGordas, 1150
et al. 4532 (BG, COL, HUA).
m, 26-27 Feb 1973 (Y fl), Croat22273 (MO); ParqueNa-
ECUADOR. ESMERALDAS:Rd. Santo Domingo de los
cional Corcovado,SirenaWoods,30 Apr 1989 (? fl-fr),Ker-
Colorados-Quininde,km 171-188, Rio Cocola, 17 Feb 1950
nan 1064 (BG); San Vito, Las Cruces Botanical Garden,
(st), Acosta S. 16221 (F); Fila Bilsa, 7 km E of San Jose de
1250-1275 m, 4 Jul 1994 ( fl-fr),Kresset al. 94-422 (US);
Bilsa, ca. 80 km S of Esmeraldas,30 Jan 1991 (st), Gentry
Osa Peninsula, Aguabuena, 3.5 km W of Rinc6n, 19 Jan
et al. 72939A (MO); between Rio Blanco & Rio Chipo, 18
1993 (9 fl), Thomsen541 (BG), 17 Apr 1993 (st), Thomsen
873 (BG). Aug 1967 (9 fl-fr), Ja'tivaet al. 2233 (NY); 3 km E of
PANAMA. COCLE: Cerro Pil6n, 1968 (st), Lallathin Quinind6, 10 Apr 1943 (st), Little6220 (Q, US); Rio Guayl-
5065 (MO). COLON:10 mi N of Gamboa,14 May 1973 (st), labamba,10 km E of Quinind6,4 Oct 1965 (d), Littleet al.
Gentry 7451 (BG); Salud,4 Aug 1971 (9 fl), Lao et al. 211 21224 (NY, Q, QAME, QCNE, US); Reserva Ecol6gica
(H, PMA). Los SANTOS: 12 mi S of Macaracas,22 Feb Cotacachi-Cayapas,CharcoVicente, Rio San Miguel, 6-9
1966 (5), Blum et al. 2176 (MO); 16 mi S of Macaracas, Sep 1993 (st), Palacios et al. 11164 (QCNE); Anchayacu,
QuebradaBejuco, 22 Jan 1966 (d), Tysonet al. 2927 (MO). Eloy Alfaro, Mayronga,21 Oct 1993 (9 fr), Penningtonet
PANAMA: Nr.El Llano,3 Sep 1977 (st), Berg et al. 410A al. 14176 (BG); Cant6nEloy Alfaro, ReservaEcol6gica Co-
(BG); BarroColoradoIs., 9 Jan 1969 (5), Croat7023 (MO), tacahi Cayapas,Rio Santiago, Angostura, 19-24 Sep 1994
2 Feb 1971 (5), Croat 13211 (MO), 29 Dec 1970 (5), Fos- (st), Tiradoet al. 1443 (QCNE). IMBABURA:Nr. Lita, 7 Feb
ter 2052 (PMA); J.W.T.C.Headquarters,1 Apr 1956 (9), 1981 (5), Berg 1240 (BG, COL, QCA, U), 9 Feb 1981 (9
Johnston 1758 (A, MO). SAN BLAS: Nusagandi, 18 Aug fl-fr), Berg 1260 (AAU, BG, COL, MO, NY, QCA, U). Los
1989 (st), Fisher 33 (BG). Rios: Rfo PalenqueBiological Station,rd. Quevedo-Santo
COLOMBIA. ANTIOQUIA:Rio Le6n(orBacubd), be- Domingo de los Colorados, km 56, 8 Jun 1974 (9 fl-fr),
tweenVillaArteaga& Chigorod6, 3 Oct 1961 Dodson 5579 (US), 11 Aug 1977 (9 fl-fr), Dodson et al.
La Pedrara,
(9 fl-fr), Cuatrecasas et al. 26186 (COL, US); Mun. San 6834 (AAU, QCA, US). PICHINCHA:Cant6nPedroVicente
Luis, Rio Claro, 25 Mar 1994 (d), Franco et al. 4585 (BG, Maldonado,Reserva Rio Silanche, Nov 1996 (9 fl), Pen-
COL, HUA); Mun. Taraza,Cgto. El Doce, Finca Las Mer- nington et al. 15246 (BG); Cant6n Pedro Vicente Maldon-
cedes, 28 Mar 1994 (d), Franco et al. 4600 (BG, COL, ado, ReservaForestalRio Pitzara,Nov 1997 (st), Pennington
HUA); Mun. Frontino, Cgto. Nutibara, rd. Nutibara-La et al. 16112 (BG).
SYSTEMATICTREATMENT
The staminate inflorescences have only 5-9
spikes, 0.6-1.2 cm diam.,in the northernandsouthern
partof the distributionrange. However,in Colombia
(Antioquia,Choc6, and Valle) the staminateinflores-
cences have ca. 15-35 spikes, 0.2-0.5 cm diam. This
very differenttype of staminateinflorescenceis rep-
resentedby the collections Cuatrecasaset al. 26187
and Franco et al. 4532, 4585, and 4600, which are in
their vegetative parts similar to the other collections.
Materialwith pistillate inflorescencesfrom the same
region are not different.This species shows morpho-
logical and ecological similarities to the Amazonian
Cecropia distachya, both becoming tall forest trees
with coriaceous and mostly smooth laminas. Cecro-
pia insignis also shows close morphologicalaffinities
to the montaneC. plicata; the latteris distinctby the
presenceof arachnoidindumentumon the leafy twigs
and more numeroussegments of the lamina.
24. Cecropia kavanayensis Cuatrecasas,Fieldiana,
Bot. 28: 210. 1951; Velasquez, Acta Bot. Venez.
6: 41, t. 6. 1971; Berg in Gorts-van Rijn, Fl.
Guianas, ser. A, fasc. 11: 97. 1992. Type. Vene-
zuela, Bolivar:Between SantaTeresitade Kavan-
ayen & Rio Pacairao,20-21 Nov 1944 (6), Stey-
ermark62754 (holotype:US; isotype: F).
CecropiaauyantepuianaCuatrecasas,ActaBot.Venez.
ActaBot.Venez.6: 41, t.
2: 203. 1967;Veldsquez,
5. 1971. Type.Venezuela.Bolfvar:CerroAuyan-
tepui,RioLomitacamp,1800m, 10May1964(6),
93643(holotype:US).
Steyermark
Bol. Soc.Venez.Ci.
Cuatrecasas,
Cecropiasteyennarkii
Nat.32: 321. 1976.Type.Venezuela.Bolivar:Mes- Distribution (Fig. 10.1). EasternVenezuela and
eta del Jaua,CerroJaua,1810-1880m, 28 Feb-5 adjacentBrazil (Roraima),in forest, at 1000-1900 m.
Mar 1974 (6), Steyermark109842(holotype:US;
isotypes:K, U). Specimens examined. VENEZUELA. AMAZONAS:
Tree,to 15(-30) m tall. Leafy twigs 4-15 cm thick,
puberulous,sometimes also with sparsearachnoidin-
dumentum, uncinate hairs lacking. Lamina coria-
ceous, ca. 40 X 40 cm to 90 X 90 cm, the segments
7-8, the free parts of the upper segments elliptic to
oblong to (sub)obovateor to ovate, the incisions down
to 5/10-6/10(-8/10?); apices roundedto obtuse; up-
per surface smooth to scabrous, sparsely to rather
densely hispidulous (or if juvenile, then densely hir-
tellous); lower surface hirtellous to tomentellous to
puberulous(withoutuncinatehairs)on the veins, with
arachnoid indumentumin the areoles, also on the
smaller veins and along the margin, sometimes ex-
tendingto the main veins; lateralveins in the free part
of the midsegment 12-16 pairs, submarginallyloop-
connected, (the lower ones) branched;petiole 40-85
103
cm long, dark purple, sparsely puberulousto hirtel-
lous or to hirsute and with (rather)dense arachnoid
indumentum;trichilia fused, the brown indumentum
intermixedwith short brownishto whitish (unicellu-
lar) hairs;stipules 15-35 cm long, red-brown,densely
to rathersparsely hirtellous to subhirsuteand often
also rather sparse arachnoid indumentum outside,
sparselyto densely subsericeousinside. Staminatein-
florescences in pairs,the peduncleerect (?); peduncle
4-10 cm long, sparsely puberulous;spathe 9-12 cm
long, red-brown,puberulousto hirtellous to subhir-
sute, also with ? dense brownpluricellularhairs,and
often with sparseto dense arachnoidindumentumout-
side, glabrousinside; spikes 4-15, 3-7 X 0.3-0.7 cm,
with stipes to 1.2 cm long and subglabrous;rachis
hairy. Staminateflowers: perianth tubular, 1.5-2.5
mm long, with sparse straighthairs or arachnoidin-
dumentumbelow the apex, the apex almost plane,
smooth; filamentsflat; anthers0.5-0.7 mm long, ap-
pendiculate, detached at anthesis, reattachedto the
marginsof the apertureby the appendages.Pistillate
inflorescencesin pairs,pendulous;peduncle9-12 cm
long, glabrous to sparsely puberulousto hirtellous;
spathe 9-11 cm long, the color and indumentumas
in the staminateinflorescence;spikes 4, (4-)8-10 X
ca. 1.2 cm, to 22 X 1.8 cm in fruit, (sub)sessile or
with spikes to 0.4 cm long and puberulous;rachis
hairy. Pistillate flowers: perianth 4-4.5 mm long,
with arachnoidindumentumbelow the apex outside,
also below the style channel inside, the apex convex,
smoothor muriculate;style long, ? S-shaped;stigma
penicillate. Fruit oblongoid, ca. 3 mm long, tubercu-
late.
Depto. Rio Negro, Cerro de La Neblina Camp V, 1250 m,
21-24 Mar 1984 (d), Liesneret al. 16945(BG), (Y fl-fr),
Liesneret al. 16951(BG).BOLiVAR: Apacara,RioCaroni,
1500 m, 12 Nov 1946 (st), Cardona1951 (US, VEN); Mun.
Cedetio, Serranfade Maigualida,20 km E of San Jose de
Kayama, 1250 m, Apr 1989 (Y fl-fr), Ferndndez5505 (BG,
MO); Mun. Roscio, N of San Juande Kamoirin, ca. 10 km
S of La Ciudadela,1300 m, 6 Mar (Y fr), Huber9235(BG,
NY, VEN); rd. Kavanayen-SantaElena, km 13, 1200 m,
Maas et al. 5384 (K, MO, U); Depto. Sifontes, Parupa,To-
ron Meru, 1300 m, 25 Apr 1986 (d), Pic6n 1108 (BG);
Cerro Uananapan,S of Uei-tepui, 1330-1450 m, 25 Apr
1960 (st), Steyernark et al. 758 (NY, US); Mt. Roraima,
between QuebradaKa-hua-parn& Glycon swamp, 1220-
1980 m, 1 Oct 1944 (st), Steyermark59016 (F, NY); Cerro
Venamo,Rfo Venamo,nr.Guyanaborder,900-1000 m, Stey-
ermark et al. 92863 (US, VEN); drainageof Rfo Cuyuni,
130-131 km S of El Dorado, ca. 1300 m, 19-26 Dec 1970
104
(Y fl-fr), Steyermarket al. 104065 (K, NY, S, U, VEN); indumentumoutside, glabrousinside; spikes (6-)15-
Meseta del Jaua,Cerro Jaua, 1810-1880 m, 28 Feb-5 Mar
1974 ($ fl-fr), Steyennarket al. 109843 (K, U, US); Mt. cm long and hairy; rachis hairy. Staminateflowers:
Roraima, Arabapu,ca. 1400 m, 1 Jan 1928 (st, juv), Tate
250 (NY).
BRAZIL. RORAIMA: MaitaMtns., 3?15'N,63?38'W,12
Feb 1971 (d), Prance et al. 10473 (INPA, NY, U, US).
This species shows morphological affinities to mm long, remainingattachedto the filamentby 2 fili-
both Cecropia obtusa and C. ficifolia. It is myrme-
cophytic.
Vernacular name. Venezuela: camai-yin-yek rescences in pairs, deflexed to pendulous;peduncle
(Arekuna,Bolivar).
25. Cecropia latiloba Miquel, in Martius,Fl. Bras.
4(1): 147. 1853; Berg, Acta Amazonica8(2): 175.
1978. Ambaiba latiloba (Miquel) Kuntze, Revis.
Gen. P1. 2: 624. 1891. Type. Brazil. Amazonas:
Rio Japura,Martius s.n. (holotype: M, photo-
graphsin F, k). Fig. 29
Cecropiaparaensis Huber,Bol. Mus. ParaenseHist. Nat. long, curved; stigma comose. Fruit oblongoid, ca. 3
6: 64 1910. Type. Brazil. Para:Rio Amazonas, Ob- mm long, finely tuberculateto (almost) smooth.
idos, Cacaoal Imperial,6 Mar 1909 (Y), Ducke s.n.
(holotype:MG 10199; isotype: RB 19456).
Cecropia orinocensis Standley, Publ. Field Mus. Nat.
Hist., Bot. Ser. 22: 14. 1940; Velasquez, Acta Bot.
Venez.6: 47, t. 9. 1971. Type.Venezuela.Amazonas:
Rio Orinoco,mouthof CafioCarida,12-24 Jan 1930
(Y), Holt & Gehriger 248 (holotype: US; isotype:
NY).
Tree, to 15(-20) m tall, the trunkwith ratherprom-
inent stipularscars.Leafy twigs 1.5-3 cm thick, green
to red-brownto purplishblack, hispidulous.Lamina
chartaceous,ca. 20 X 20 cm to 55 X 55 cm (to 70 X
70 cm), the segments 8-9, the free partsof upperseg-
ments elliptic to oblong, the incisions (in the upper
part of the lamina) down to 5/10-7/10, the lamina
attachedto the petiole ratherclose to its base; apices
obtuse; upper surface ? scabrous,hirtellous to his-
pidulous; lower surface puberulouson the veins and
with dense or sparse arachnoidindumentumin the
areoles; lateral veins in the free part of the midseg-
ment 12-14 pairs, marginallyloop-connected,often
branched,running(almost) straightinto the margin;
petiole 25-60 cm long, puberulous;trichilia fused,
with only brown indumentum;stipules 8-13(-22),
bright to dark red to brownish (or green), caducous
or subpersistent,puberulousor sparselysubsericeous
outside, sparsely villous inside. Staminate inflores-
cences in pairs,the peduncleerect and the spikes pen-
dulous; peduncle 4-10 cm, hispidulous to hirtellous
to subhirsute;spathe (5-)9-12(-18) cm long, pink to
red, puberulousor also with (rathersparse)arachnoid
FLORA NEOTROPICA
20, (3-)14-20 X 0.3 cm, with stipes (0.2-)0.4-0.6
perianthtubular,0.6-0.8 mm long, sparsely puberu-
lous just below the apex, the apex slightly convex to
plane, sparselypuberulouson the margin,the aperture
surroundedby a rim; filaments flat; anthers0.3-0.5
form connectionsbetween the connectiveand the up-
per marginof the filamentat anthesis.Pistillate inflo-
5-17 cm long, hispidulousto hirtellousto subhirsute;
spathe 7-15 cm long, the color and indumentumas
in the staminate inflorescence; spikes 4-6, 3-12 X
0.5-0.8 cm, to 25 X 1.5-2 cm in fruit,sessile (or with
stipes to 0.5 cm long and glabrous;rachis glabrous.
Pistillate flowers: perianthtubular,4 mm long, with
arachnoidindumentumbelow the apex outside, also
below the style channel (or absent) inside, the apex
slightly convex to slightly concave, smooth or punc-
tate, the apertureslit-shaped,borderedby a rim; style
Distribution (Fig. 8.1). Amazon basin, the basin
of Rio Orinoco, and in the Guianas, in periodically
inundatedplaces, especially along black-waterrivers
(on nutrient-poorand not well-aerated soil) and on
sand savannas,at low elevations.
Representativespecimensexamined.COLOMBIA.
AMAZONAS:Mun.Leticia,Quebrada
de Arara,27 Jan1969
(d), Cuatrecasaset al. 27250 (COL, F, G, K, NY, US); Rio
Nov 1945(d), Duque-Jaramillo
Loretoyacu, 2030 (COL);
Rio Yari,nr. mouthof QuebradaEl Mochilero,23 Apr 1986
(st), Galeano et al. 1093 (COL);Rio Caquetd,Isla El Tigre,
23 Sep 1988 (st), M. Sdnchez S. et al. 1473 (BG, U); Rio
Putumayo,between Rio IgaraParand& Rio Yaguas,20 Jun
1942 (st), Schultes 3996 (ECON, F); TrapecioAmaz6nico,
Rfo Loretoyacu, Oct 1945 (d), Schultes 6720 (COL, F);
AmacayacuNational Park, Matamata,21 Jan 1989 (Y fr),
TyeC.92 (BG, K); PuertoNarifno,
ParqueNacionalAma-
cayacu, 2 Aug 1989 (st), R. Vdsquezet al. 12449 (COL,
MO).AMAZONAS/VAUPES:Rio Apaporis,Jinogoj6,20 Jun
1952 (Y fl-fr), Schultes et al. 16811 (ECON, US). CA-
QUETA: Rio Yari, nr. Araracuara,15 Jul 1986 (Y fr), Berg
et al. 1562 (BG); Rio Yari, nr. mouth of QuebradaEl Mo-
chilero, 25 Apr 1986 (? fl-fr), Galeano et al. 1093 (COL);
Rio Orteguaza,betweenPeneya & Getucha,28 Jul 1926 (st),
Woronowet al. 6302 (LE). PUTUMAYO:Rio Caucaya, be-
tween Vivianacocha& La Peinilla, 10-14 May 1993 (Y fr),
Bernal et al. 2039 (COL).
VENEZUELA. AMAZONAS: Depto. Atabapo,Rio Parn,
Sep 1989 (Y fr), Delgado 720 (BG, NY); Rio Casiquiare,
Jan-Feb 1969 (Y fl-fr), Farinas et al. 502 (US, VEN);
Depto. Atabapo,CafnoJayuwapuey,affluentof Rio Ocamo,
Jan 1990 (2 fl), Ferndndez6775 (BG, NY); Brazo Casi-
SYSTEMATIC TREATMENT 105

VW

7 .\~~~~~~~~~T

I --I TI T~ .-.---

'~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~

S~~~~~/;~
At )m t o/C

FIG. 29. Cecropia latiloba. 1. Leaf, stipules, and trichilia. 2. Apex of lamina and venation. 3. Pair of staminate
inflorescences and base of petiole with trichilium. 4. Staminate inflorescence at anthesis and base of petiole with trichilium
(Berg et al. 1057). 5. Staminateflowerand stamenafterdetachmentof anther(Prance et al. 20925). 6. Pistillateinflorescence
at anthesis and base of petiole with trichilium(Berg et al. 1032). 7. Pistillate inflorescenceafter anthesisand base of petiole
with trichilium(Berg et al. 1575). 8. Pistillate flower.9. Fruit(Berg et al. 1562).
106
quiare, 1-20 Feb 1931, Holt et al. 657 (NY, US); Rio Siapa,
between mouth & campamento de los Yanomani-
Emonitheri, 17-20 Sep 1986 (Y fl-fr), Stergios et al. 9401
(BG, MO, NY, VEN); LowerRio Casiquiare,Solano, 10 Mar
1942 (? fr), LI. Williams 14733 (F, G); Upper Rio Casi-
quiare,Capihuara,3 Jun 1942 (? fr), Ll. Williams15765 (G,
US); Rio Casiquiare,between mouth of Rio Siapa & mouth
of Rio Paciba, 26 Jul 1959 (Y fr), Wurdacket al. 43634
(BG, NY). APURE: Dtto. San Fernando,between Rfo Ori-
noco, opposite Isla Peladura& lagunasca. 1 km inland,25-
26 Apr 1977 (6), Davidse et al. 12211A (MO, U, VEN);
Dtto. San Fernando,mouthof Rio Arauca, 14-15 May 1977
(Y fl), Davidse et al. 13258 (MO, VEN); Dtto. Pedro Ca-
mejo, Rio Orinoco, opposite Isla Pantallo,24-25 Feb 1978
(d), Davidse et al. 14500 (MO, U, VEN); Dtto. Pedro Ca-
mejo, Rio Orinoco, Isla El Fraile, 3 Nov 1989 (Y fr), Duno
et al. 174 (BG). BOLiVAR: Mun. Raul Leoni, 25 km N of
Macizo Ichun, Alto Rio Paragua,Apr 1988 (Y fl-fr), Fer-
ndndez4399 (MO); Mun. Raul Leoni, 50 km SW of Macizo
Guaiquinima,72 km W of Karnn,Nov 1988 (Y fr), Fer-
ndndez et al. 4819 (MO); Rio Nichare, between mouth &
CanioSarrapio, 3-10 Aug 1985 (Y fr), Homer et al. 73
(MO); Rfo Oris, nr. CampamentoTurumban,13 May-13
Jun 1987 (6), Stergios 10800 (BG, NY); La Paragua,25
Mar 1940 (6), Ll. Williams 12689 (F, K, S, US, VEN).
DELTA AMACURO:Dtto. AntonioDiaz, Rio Acure,between
Las Margaritas& Curiapo,31 Jan 1980 (6), A. Femnandez
et al. 3527 (U); Sacupana,Apr 1986 (Y fl), Rusbyet al. 273
(A, BM, E, F, G, GH, K, MICH, MO, NY, VEN); Rio Cu-
yubini, between la Paloma & Moron, 20 Nov 1960 (6),
Steyermark87693 (BG, NY); between La Margarita &
Puerto Miranda,Rio Acure, 26 Nov 1960 (d), Steyermark
87808 (BG, NY, VEN); Depto. Antonio Diaz, Rio Cuyubini,
between mouths of Rio El Toro & Rio Mecoro-uabanoco,3
Nov 1980 (Y fr), Trujilloet al. 17307 (NY, U, VEN).
GUYANA. MazaruniR., PuruniR., 21 Apr 1953 (Y fl-
fr), "J.B."85 (= FD 7769) (K, NY, U); U. Takutu-U. Es-
sequiboregion, Rewa R., 0-5 km S of confluencewith Kwi-
taro R., 25 Feb 1997 (6), Clarke 3937 (BG, US); Cuyuni-
Mazaruni region, Cuyuni R., between Aurora & 4 km
upstream,8 Oct 1989 (Y fl-fr), Gillespie et al. 2195 (BG,
COL);EssequiboR., Rockstone,31 Jul 1921 (Y fr), Gleason
864 (GH, NY); Demerara-Mahaicaregion, 23 km E & W of
Linden Hwy., within 3 km of second tollgate, 16 Aug 1993
(6), Henkel et al. 2584 (BG); Upper Takutu-UpperEsse-
quibo region, S of RupununiSavanna,KidekperdanaCr., 8
km SE of Aishalton, 26 Nov 1993 (Y fl-fr), Henkel et al.
3519 (BG); Upper Takutu-UpperEssequibo region, Kassa-
kaityu, 24 Mar 1994 (6), Henkel et al. 5318 (BG); Kakunu
Mtns., Rupununi R., Crabwood Cr., 27 Jun 1995 (6),
Jansen-Jacobset al. 4231 (BG, U); RupununiR., SW of Mt.
Makarapan,10 Sep 1988 (Y fl-fr), Maas et al. 7339 (COL,
K, MO, NY, U); Berbice-Corantyneregion, Potoko Cr., 24
Apr 1990 (Y fr), McDowell 2455 (NY).
SURINAME. Kwattaweg, W of Paramaribo,21 Dec
1948 (6), BBS 223 (NY, U); CoppenameR., nr. Kaaiman-
stone, 7 Sep 1933 (9 fl-fr),Lanjouw720 (U); Distr.Nickeri,
KabaleboDam projectarea, 8 Sep 1980 (9 fl-fr),Lindeman
& Girts-van Rijnet al. 288 (NY, RB, U); WilhelminaMtns.,
FLORA NEOTROPICA
confluence of Lucie R. & Oost R., 12 Jul 1963 (2 fr), Ma-
guire et al. 54150 (BG, NY, US); CorantijnR., 3 Aug 1935
(6), Rombouts64 (U); CoppenameR., nr. Heidoti, 24 Apr
1956 (6), Schulz(LBB) 7616 (F, U); 1 km NW of Voltzberg,
10 Feb 1965 (6), Schulzet al. (LBB) 10621 (U, US); Litanie
R., Dec 1903 (6), Versteeg418 (U); Oelemari R., 19 Mar
1963 (6), WesselsBoer 971 (U), (2 fl-fr), WesselsBoer 972
(U); confluence of Paloemeu R. & TapanahoniR., 13 Apr
1963 (2 fl-fr), WesselsBoer 1256 (U).
FRENCH GUIANA. Comte R., Apr 1961 (2 fl-fr),Au-
bre'ville354 (P, U); Inini Basin, Saut Sonnelle, 23 Jan 1994
(6), Cremers13269 (BG), (st), Cremers13270 (BG); Ap-
prouageR., TortueCr., 25 Jan 1967 (2), Oldeman2321 (P,
U, US); Maripasoula,25 Jan 1984 (2 fl), Prevost 1468 (U);
Belingui, S of Maroni,Beiman Cr., 21 Nov 1984 (2 fl-fr),
Prevost 1693 (INPA, P, S, U), (6), Prevost 1700 (INPA, P,
S, U).
ECUADOR. NAPO: ParqueNacional Yasuni,rd. Pom-
peya Sur-Iro, km 38.4, 17 May 1994 (st), Jaramillo et al.
16518 (QCA), 16525 (QCA). SUCUMBiOS: Reserva Faun-
istica Cuyabeno, Rio Cuyabeno, upstream from Laguna
Grande, 11 Mar 1990 (2 fr), Balslev et al. 97162 (AAU,
BG, QCA, QCNE); Rfo Cuyabeno,ca. 0'20'S, 75?55'W,15
Feb 1980 (2 fl-fr), Berg et al. 1032 (BG, COL, QCA, U),
18 Feb 1980 (2 fl-fr), Berg et al. 1056 (AAU, BG, COL,
GB, MO, NY, QCA, TUR, U), (6), Berg et al. 1057 (AAU,
BG, COL, GB, MO, QCA, TUR, U).
PERU. LORETO: Nr. Iquitos, 6 Nov 1940 (2 fl-fr),As-
plund 14350 (S), 14 Nov 1940 (6), Asplund14499 (S); Prov.
Maynas, Caballo Cocha, 24 Apr 1982 (2 fr), Ayala et al.
3310 (BG, MO, NY); Prov. Requena,Rio Ucayali, Huarmi
Isla Cocha, 14 Dec 1987 (2 fr), Ayala et al. 5930 (F, MO);
rd. Iquitos-Maynas,km 2, 4 Jul 1988 (6), Berg et al. 1570
(BG), (2 fr), Berg et al. 1575 (BG, COL, USM); Prov.Re-
quena, Reserva Nacional Pacaya Samiria, 5 May 1983 (2
fr), Del Carpio 2087 (USM); Prov. Maynas, Rfo Yaguasy-
acu, 9 Nov 1977 (2 fl), Gentry et al. 20514 (F, MO, NY,
USM); Prov.Maynas,Rio Curaray,nr. BuenaVista, 16 Nov
1986 (2 fl), Kalliola et al. P5-4 (BG, USM) Prov.Maynas,
Rio Amazonas,Isla Rondifia,25 Mar 1977 (2 fr), Plowman
et al. 6433 (F, GH, US, USM); Prov. Alta Amazonas, An-
doas, 20 Nov 1980 (d), R. Va'squezet al. 776 (USM). MA-
DRE DE DIos: Prov.Tambopata,15 km ENE of PuertoMal-
donado, 1991 (2 fr), Fisher 179 (BG); Parque Nacional
Manu, Rio Manu, Rio Cumerjali,21 Oct 1986 (6), Foster
et al. 11960 (BG, LPB, MOL, US, USM); Manu, Pakitza,
19 Jan 1987 (2 fl-fr), Nuiniez6896 (BG); Prov. Tambopata,
Las Piedras, QuebradaGamitana,26 Jan 1991 (2 fr), Ti-
mand et al. 1406 (MO). UCAYALI: Prov. Coronel Portillo,
Yarinacocha,Dec 1968 (2 fr), Ferreyra17625 (US, USM);
Prov. Coronel Portillo, Boquer6ndel PadreAbad, 50 km E
of Aguaytia, 9 Aug 1988 (2 fl), Gentryet al. 29531 (BG);
Prov.CoronelPortillo,Lago Yarinacocha,12 Sep 1980 (6),
Maas et al. 4603 (BG, USM); Rio Ucayali, Masea, Nov
1923 (2 fl-fr), Tessmann3307 (NY).
BRAZIL. AcRE: Rio Jurua,nr. Colonia RodriguezAl-
vez, 29 Sep-10 Oct 1986 (6), Campbellet al. 11161 (BG);
Mun. Rio Branco, Lago Amapa, 3 Dec 1982 (2 fl-fr), L.
Coelho et al. 1908 (INPA, NY); Mun. Bujari, Rio Branco,
SYSTEMATICTREATMENT
Riozinho do Andira,24 Mar 1995 (Y fl-fr), Pardo et al. 75
(BG, NY); Mun. Mancio Lima, Rio Moa, ParqueNacional
da Serra do Divisor, 8 May 196 (Y fr), M. Silveira et al.
1310 (NY); Rio Jurua,Fortaleza,Nov 1901 (Y fl-fr), Ule
5938 (G, L, MG). AMAPA: Rio Jarn,IgarapeGrande,ca. 1
km N of Arumanduba,24 Dec 1961 (Y fr), Egler 45948
(BG, MG, NY, U, US). AMAZONAS: Rio Solimoes, 1-2 km
E of Boca de Januaca,2 Oct 1973 ( ? fr), Berg et al. P17586
(F, INPA, K, MG, MO, S, U, US); Paranddo Autaz-Mirim,
Lago do Caioe, 26 Aug 1973 (Y fr), Berg et al. P 19769 (C,
EAP, F, INPA, K, MG, MO, NY, S, U, US); Mun. Novo
Ayroa,Rio Negro, Arquipelagode Anavilhanas,24 Jan 1992
(d), L. V Ferreira 109 (BG, K, NY); Manacapurui,6 Mar
1904 (? fl-fr), Huber (MG) 4179 (BM, G, MG, U); Lower
Rio Purus, 18 Mar 1904 (Y fl-fr), Huber (MG) 4195 (BM,
G, MG, U); Mun. Humaita,nr. Livramento,on Rio Livra-
mento, 12 Oct-6 Nov 1934 (d), Krukoff6718 (A, BM, F,
G, K, MICH, MO, NY, RB, S, U, US); Mun. Sao Paulo de
Oliven,a, nr. Palmares, 11 Sep-26 Oct 1936 (d), Krukoff
8407 (A, BM, F, G, K, MICH, MO, NY, P, S, U, US); Rio
Negro, AnavilhanasIs., 2?34'S,60?19'W,7 Aug 1991 (Y fr),
Mori et al. 21959 (BG, NY, S); Mun. Maraa,Rio Japua,nr.
liha Nova, 3 Dec 1982 (d), Plowman et al. 12160 (INPA,
MG, NY, RB); nr. Tototobi, 27 Feb 1969 (Y fl-fr), Prance
et al. 10289 (INPA, K, MG, NY, S, U, US); nr.Manaus,Rio
Solimoes, Furo de Xiborena,4 Apr 1974 (6), Prance et al.
20925 (GH, INPA, K, MG, MO, NY, P, S, U, US), (Y fl-fr),
Prance et al. 20927 (INPA, K, MG, MO, NY, S, U, US),
(d), Prance et al. 20929 (C, F, INPA, K, MG, MO, NY, P,
U, US). PARA:Rio Trombetas,1 km S of CachoeiraPorteira,
2 Jun 1974 (Y fl-fr), Campbell et al. P22441 (INPA, NY,
U); Mun. Oriximina,Rio Trombetas,Minera,co Santa Pa-
tricia, 7 Jul 1980 (Y fl), L. V Ferreira et al. 1341 (INPA,
MG, NY, U, US); Mun. Tucuruf,Rio Tocantins, 1-5 km
upriver from Represa Tucuruf, (Y fl-fr), Plowman et al.
9883 (F, GH, INPA, MG, MICH, MO, NY, U, US), 22 Mar
1980 (d), Plowman et al. 9886 (F, GH, MG, MO, NY, U,
US); Rio Maicuru,Lageira, 19 Jul 1981 (Y fr), Strudwick
et al. 3187 (INPA, MG, NY). ROND6NIA:PortoVelho, Rio
Machado,27 Nov 1977 (Y fl), Goulding 7 (INPA);Rio Ma-
deira, Calama,Apr 1980 (st), Goulding Ila (MG); Rio Ma-
chado, Jan 1981 (Y fr), Goulding 1252 (MG); Upper Rio
Machado,nr. Tabajara,24 Nov 1931 (d), Krukoff1438 (A,
BM, G, K, MICH,NY, P, S, U). RORAIMA: Ilha de Maraca,
Mun. Alto Alegre, Rio Uraricuera,Furo Santa Rosa, 8 Jun
1986 (d), Hopkinset al. 872 (BG, INPA, NY); Ilha de Mar-
aca, FuruParanddo Firmino,3 Jul 1987 (Y fl-fr), Milliken
et al. 388 (BG, E, K); Rio Uraricuera,between 63010' &
63?30'W,13 Mar 1979 (d), Pires et al. 16963a (U), 16963b
(NY); Rio Mucajai,24 Mar 1971 (' fr), Prance etal. 11207
(INPA, K, MG, NY, S, U, US).
BOLIVIA. BENI: Prov.Yacuma,Rio Yacuma,Espiritu,
9 Apr 1981 (Y fr), Beck et al. 5292 (LPB), 28 Feb 1990 (Y
fl-fr), Moraes R. et al. 1239 (BG, LPB, MO, USZ); Prov.
Marban,ParqueIsiboro-Secure,5 Jun 1992 (st), Seidel et al.
6674 (LPB). PANDO:Prov.Manuripi,Rio Manuripi,Puerto
Rico, 23 Jan 1983 (9 fr), FerndndezC. et al. 8410 (G, NY).
SANTACRUZ:Prov.Velasco,ReservaEcol6gica El Refugio,
107
Rfo Paragua,between CampamentoEl Refugio & Campa-
mento Toledo, 7 Feb 1995 (9 fr), Guillenet al. 3170 (USZ).
In the upper Amazon basin, particularlyin the
northwesternpart, the spikes of the fruitingpistillate
inflorescenceare short, usually to 10 cm long. In the
Guianas,VenezuelanGuayana,andpartsof Brazil,the
fruitingspikes can be to 25 cm long, and the pedun-
cles are usually longer as well. The species is com-
monly found in periodically inundated areas along
black-waterrivers or in depressions with sandy and
poorly drainedsoil away from white-waterrivers(cf.
Lamotte, 1992). It is also found in periodically in-
undatedwhite sand savannasin the Guianas.It is eco-
logically clearly distinct from Cecropia membrana-
cea, the other species of the genus that tolerates
extended inundation.
Cecropia latiloba and C. utcubambanaare mor-
phologically very close and almost certainlyclosely
related.
Vernacular names and use. Venezuela:simbra-
potro (Apure); tokorodek (Pen6m, Bolivar); uaru
(Warao, Delta Amacuro). Guyana: wanasoro. Suri-
name:bospapaja.Peru:kabeari(Machiguenga,Madre
de Dios); setico colorado (Ucayali). Brazil: imbauiba
branca, imbauba da varzea (Amazonas); imbauba
branca(Mato Grosso); imbaubarana(Rondonia);to-
kori (Uaica'-Mucajai,Roraima).
In Madre de Dios, bark fibers are used to make
arrowstrings (D. W. Yu, pers. comm.).
26. Cecropia litoralis Snethlage,Notizbl. Bot. Gart.
Berlin-Dahlem 8: 392. 1923. Type. Ecuador.
"CoastalPlain,"withoutlocality,Rimbach15 (ho-
lotype: B, destroyed, duplicates not found; pho-
tographsin F, G, GH, MICH, MO), herewithre-
placed by: Ecuador.El Oro: Between Pasaje &
Uzhcurrumi,2 Feb 1981 (9), Berg 1238 (neotype:
QCA; isoneotypes:BG, COL, U). Fig. 30
Tree,to 10(-25) m tall, the trunkwith prominent
stipularscars. Leafy twigs 1-3 cm thick, green with
conspicuous lenticels, densely hispidulous (with
curved to straighthairs). Laminachartaceousto sub-
coriaceous, ca. 40 X 40 cm to 75 X 75 cm, the seg-
ments 7-9, the free parts of upper segments elliptic
to oblong to (sub)obovate,the incisions down to 6/
10-8/10; apices obtuse to short-acuminate;uppersur-
face scabridulous,sparselyhispidulousand (initially)
with sparse arachnoid indumentum;lower surface
(minutely) puberulousto subtomentellous(or partly
to hirtellous), with arachnoid indumentum in the
areoles and on the smaller veins, extending to the
108 FLORA NEOTROPICA

1014~

FIG.30. Cecropia litoralis. 1. Leaf. 2. Apex of lamina and venation.3. Stipules and base of petiole with trichilium
(Berg et al. 1043). 4. Leafy twig with staminate inflorescences in various stages of development (with spathes and at
anthesis) and bases of petioles with trichilia (Balslev et al. 84740). 5. Staminateflower and stamen (Balslev et al. 97161).
6. Pistillate inflorescencewith spathe (Dodson 7334). 7. Pistillate inflorescenceat anthesis. 8. Pistillate inflorescenceafter
anthesis and base of petiole with trichilium.9. Pistillate flower. 10. Fruit(Berg et al. 1046).
SYSTEMATICTREATMENT 109

main veins; lateral veins in the free part of the mid- Vrda. El Tigre, 23 Feb 1995 (9 fl-fr), Franco et al. 5544
segment 13-19 pairs, marginally loop-connected, (BG, COL, HUA, US), (6), Franco et al. 5545 (BG, COL,
mostly branched;petiole 25-75 cm long, puberulous HUA, US), (9 fl-fr), Franco et al. 5546 (BG, COL, HUA,
to hirtellousor also with arachnoidindumentum;tri- US). PUTUMAYO:Mun. PuertoAsfs, rd. to Kanakas,16 Feb
1995 (9 fl-fr),Francoet al. 4681 (BG, COL,HUA, LP,MO,
chilia fused, the brown indumentumintermixedwith
NY, US).
short white hairs; stipules 8-17 cm long, yellowish ECUADOR. EL ORO: Nr. Piedras, 23 Nov-16 Dec
green to reddish, when dry usually chartaceousto 1978 (9 fl-fl), Albert de Escobar 984 (QCA, TEX, U); be-
membranaceous, puberulous to strig(ill)ose and tween Pasaje & Uzhcurrumi,2 Feb 1981 (6), Berg 1237
with sparse arachnoidindumentumoutside, (rather) (BG, COL, QCA); 30 km SW of SantaRosa, rd. to Pinias,6
sparsely sericeous inside. Staminate inflorescences Oct 1979 (9 fl), Dodson et al. 8869 (F, MO); Puyango, 29
solitary or in pairs,the peduncleerect to deflexed,the Dec 1997 (9 fl-fr), P. Lozano et al. 902 (BG, LOJA),(d),
spikes pendulous; peduncle 6-16 cm long, puberu- P Lozano et al. 903 (BG, LOJA);nr. Portovelo, 6-15 Oct
lous to hispidulousand/or(initially)with brownplur- 1918 (6), Rose 23411 (GH, NY, US). ESMERALDAS:Can-
icellular hairs and sparse arachnoid indumentum; t6n San Lorenzo, La Tola, 21 Nov 1993 (6), Alvarez 740
(QCNE); nr. Borb6n, 6 Aug 1967 (9), Jditivaet al. 2160
spathe 7-14 cm long, white to/or pinkish or yellow-
(NY, S); Rio Santiago, between Borb6n & Palma, 10 Aug
ish, when dry usuallychartaceousto membranaceous, 1967 (9 fl-fr), Jdtiva et al. 2223 (AAU, NY), (6) Jdtiva et
sparsely puberulousor partly to strig(ill)ose to sub- al. 2224 (AAU, MO, NY); nr. Esmeraldas,29 Sep 1965 (9
hirtellousand with very sparseto ratherdense arach- fl-fr), Little 21176 (NY, QAME, QCNE, US). GUAYAS:
noid indumentumoutside, glabrousinside; spikes 6- Cerro Azul, W of Guayaquil,9 Feb 1955 (9 fr), Asplund
15, 5-12 X 0.2-0.3 cm, with stipes 0.2-1 cm long 15377 (S); nr. Naranjito, 6-7 Jun 1945 (9 fl-fr), Camp
and hirtellous to puberulous;rachis glabrous.Stami- E.3634 (BG, NY, S); Cant6nNaranjal,Taura,ReservaEcol-
nate flowers: perianth tubular,ca. 1 mm long, gla- 6gica Manglares-Churute,22 Feb 1991 (6), Ceron et al.
brous, the apex plane; filamentsslightly swollen; an- 13470 (QCNE);rd. Guayaquil-Salinas,km 15, 2 Nov 1995
thers ca. 0.5 mm long, (short-)appendiculate,at (9 fl-fr), Clark et al. 1530 (BG, QCNE). LOJA:Rd. Ala-
mor-Arenillas, nr. bridge over Rio Puyango, 30 Dec 1997
anthesis remainingattachedto the narrowedapex of
(d6), P. Lozano et al. 912 (BG, LOJA). Guayachuma,rd.
the filament (but after anthesis detached). Pistillate Loja-Machala, ca. 1400 m, 23 Sep 1967 (9 fl), Sparre
inflorescences solitary or in pairs, erect to deflexed, 18866 (S). Los Rios: Rio PalenqueBiological Station,rd.
becoming pendulousin fruit;peduncle9-16 cm long, Quevedo-Santo Domingo de los Colorados,km 56, 23 Jul
sparselypuberulousto hirtellousbelow the spatheand 1975 (6), Dodson 5880 (QCA, MO, US), 1 Dec 1978 (9
with sparse to dense arachnoidindumentum,some- fr), Dodson 7334 (BG, QCNE); Cant6nVinces, rd. Quev-
times subglabrous;spathe 8-19 cm long, the color, edo-Palenque, km 70, Jaunecheforest, 26 Mar 1980 (9 fr),
texture,and indumentumas in the staminateinflores- Dodson et al. 9922 (BG, F, MO); 18 km S of Empalme,nr.
cence; spikes 4(-5), 7-16 X 0.4-0.6, to 28(-37) X Guayas border,5 Feb 1974 (9 fl-fr), Gentry 9673 (MO);
between Babahoyo & Montalvo, 18 Feb 1967 (6), Sparre
1-1.5 cm in fruit, with stipes 0.3-1 cm long and pu-
14534 (S). NAPO:Cant6nPompeya,La Joya de los Sachas,
berulous to glabrous;rachis hairy. Pistillate;flowers:
Isla de Pompeya, 6 Ec 1992 (9 fl-fr), Gudifnoet al. 2021
perianthca. 1.5 mm long, with arachnoidindumen- (BG, QCNE);Lagunade Yuturi,28 Feb 1990 (9 fl-fr),Jar-
tum on and below the apex, also in the style channel amillo et al. 11475 (GB, NY, QCA). PICHINCHA:SantoDo-
inside, the apex plane; stigma peltate. Fruit oblon- mingo de los Colorados,22 Aug 1986 (d), Vivaret al. 2844
goid, 1.8-2 mm long, tuberculate. (LOJA). SUCUMBiOS: Reserva Faunistica Cuyabeno, Rio
Cuyabeno, between Laguna Grande& 5 km upstream, 11
Distribution (Fig. 8.3). Coastal regions of Ecua-
Mar 1990 (6), Balslev et al. 97160 (AAU, BG, QCA,
dor and of Narinlo(Colombia);very common in wet QCNE); Rio Cuyabeno, ca. 0?10'S, 75?55'W,17 Feb 1980
to relatively dry forest, coastal swamps, and second- (6), Berg et al. 1043 (BG, COL, GB, MO, QCA, TUR, U),
ary growth;at elevations to 1500 m. Also occurring (9 fl-fr),Berg et al. 1046 (AAU, BG, COL, GB, MO, QCA,
in the upperAmazonbasin,probablydiscontinuously, TUR, U).
known from Colombia (Caqueta'and Putumayo),Ec- PERU. LORETO: Nr. Iquitos, 30 Nov 1940 (9 fl-fr),
uador (Sucumbios), and Peru (Loreto), along (black- Asplund 14810 (S); Prov. Requena,nr. JenaroHerrera,Co-
water) rivers in places that are not inundated or cha Supay,Arboretumde Braga, 18 Jul 1988 (9 fl-fr), Berg
inundatedonly during short periods, often in associ- et al. 1586 (BG, USM), 16 Dec 1987 (9 fr), Lamotte375
ation with Cecropia latiloba. (BG); Prov. Alto Amazonas, Andoas, Rio Pastaza, 10 Nov
1980 (6), R. Vdsquezet al. 776 (BG).
Representative specimens examined. COLOMBIA.
CAQUETA: Rio Caqueta, below mouth of Rio Orteguaza, Cecropia litoralis is closely related to C. engleri-
Solano, 8 km SE of Tres Esquinas,8 Mar 1945 (Y fl), Little ana, which is sympatric with C. litoralis in the north-
et al. 9684 (COL). NARINO: Mun. Tumaco, rd. to Pasto, western part of the Amazon basin. Cecropia litoralis
110 FLORANEOTROPICA

can be easily distinguishedfrom C. englerianaby the peduncle ca. 30-90 cm long, hirtellous;spathe 7-12
smallernumberof lateralveins, to ca. 15 pairs,but in cm long, the color unknown, the indumentumas in
the other features the two taxa cannot be distin- the staminate inflorescence; spikes 6-12, often ?
guished. One could arguethatthese two taxa are sub- crinkled,4-10 X 0.6-0.8 cm, to 8 X 1.1 cm in fruit,
species ratherthanspecies (see "Ecology,"discussion with stipes 0.3-0.8 cm long and puberulousto hirtel-
of the C. peltata-group). It is on phytogeographical lous; rachishairy.Pistillateflowers: perianthca. 1.5-
and ecological considerationsthat they are kept dis- 2 mm long, with arachnoidindumentumbelow the
tinct at the species level for the time being. In the apex outside, also in the style channelinside, the apex
coastal region of Ecuador and the adjacent part of + convex, muriculate;style short,muriculate;stigma
Colombia, C. litoralis is a ? weedy species in vari- comose. Fruit subobovoid to oblongoid, ca. 2 mm
ous habitatsbut not clearly associated with riverine long, smooth, pale brown.
habitats.However,in the Amazon basin it is common
Distribution (Fig. 8.3). Eastern Panama and
along Rio Cuyabeno, a black-watertributaryof Rio
northwesternColombia,mostly in secondarygrowth,
Aguarico, and is there associated with C. latiloba.
at low elevations.
The species has also been collected in Peru (Loreto)
in a C. latiloba habitat.The association with C. lati- Representative specimens examined. PANAMA.
loba indicates ecological preference different from COLON:S of Fort Sherman, 27 Jul 1955 (9 fl-fr), L M.
that of C. engleriana, which is an uplandspecies, or Johnston 1503 (A, MO); nr. MaddenDam, 17 Jul 1938 (9
fr), Woodsonet al. 1312 (A, MO, NY). DARIEN: Cruce de
if it is found in riverine habitats,then ratherin asso-
Mono, 5 Nov 1989 (st), Fisher 53 (BG); Rio Tuquesa,lower
ciation with C. membranacea.
TuquesaMining camp, 5 Jul 1975 (9 fr), Mori 6998 (BG,
MO). PANAMA: El Llano-Cartird., nr.El Llano, 1 Sep 1977
(9 fl-fr), Berg 410 (BG); 1.5 mi beyond intersection of
27. Cecropia longipes Pittier,Contr.U.S. Natl. Herb. USARSO chemical trainingcamp area, 15 Jun 1967 (9 fl-
18: 227. 1917; Croat, Fl. Barro Colorado Island fr), Correa et al. 27 (COL, MO, PMA, SCZ); Barro Colo-
347, t. 195. 1978. Type. Panama.Panama':Nr. Ta- radoIs., 30 Apr 1970 (9 fl), Croat10115 (COL,MO, SCZ),
bernilla, 6 Jul 1911 (d), Pittier 3823 (holotype: 23 Jun 1971 (6), Croat 15083 (MO, NY, SCZ); Capitana,
US). Chepo, 9 Jun 1971 (6), Lao 120 (MO, PMA, SCZ), (9 fl-
fr), Lao 121 (H, MO, PMA); nr. El Real, 22 May 1969 (9
Tree, to 18 m tall. Leafy twigs 1.5-4 cm thick, fl), Lazor et al. 3440 (MO); Curundu,20 Jul 1985 (9 fl-fr),
green, hispid(ulous)with uncinatehairs to hirtellous. Nevers et al. 5994 (BG, MO); 2 mi E of Santa Fe, 18 Jul
Laminachartaceous,ca. 20 X 20 cm to 70 X 70 cm, 1966 (9 fr), Tysonet al. 4854 (MO, SCZ).
the segments 7-10, the free parts of the upper seg- COLOMBIA. ANTIOQUIA: Uraba-Chigorod6-
Malag6n, CafnoMalag6n, 22 Mar 1986 (9 fl-fr), Renteria
ments ovate to elliptic, the incisions in the upperpart
et al. 4735 (JAUM).BOLiVAR: Mun. Turbaco,8 Jun 1982
of the lamina down to 4/10-6/10; apices roundedto
(9 fl), Cuadros V 1374 (COL, MO, US); Rfo San Jorge,4
short-acuminate;upper surface ? scabrous,puberu- Jul 1948 (9 fl), Romero-Castaneda1137 (COL, F, US).
lous to hispidulous;lower surfacepuberulousto sub- CHocO: Mun. Acandi:Vrda.Coquita,El Paramo,Quebrada
tomentose on the veins, with arachnoidindumentum Zardi, 26 May 1989 (9 fl-fr), Betancur et al. 1255 (HUA,
in the areoles or also on the reticulum;lateral veins MO); Mun. Turbo,rd. Tap6ndel Dari6n,between Rio Le6n
in the free partof the midsegment5-8 pairs, margin- & Lomas Aisladas, km 37, 28 May 1984 (9 fl-fr), Brand
ally loop-connected, often branched;petiole ca. 20- 1201 (JAUM,MO); Mun. Riosucio, Cgto. La Honda,Vrda.
60 cm long, puberulousto hirtellous;trichilia fused, La Balsa, Canal La Balsa, 15 Apr 1987 (6), Echavarriaet
the brown indumentumintermixedwith dense long al. 149 (JAUM);Mun. Riosucio, rd. Peye-Sautata, 5 Jun
Foreroet al. 1890 (COL, MO); Mun. Acandi,
whitish hairs; stipules 4-9 cm long, color unknown, 1976 (9 fl),
Cgto. Ungufa, Rio Cuti, 27 Jul 1957 (9 fl), Romero-
subhirsute to hirtellous or to subsericeous outside,
Castaneda 6453 (COL); ParqueNacional NaturalLos Ka-
sparsely hairy inside. Staminate inflorescences in tios, Sautana,3 May 1982 (6), ZuluagaR. 194 (COL).
pairs, at least the spikes pendulous (?); peduncle 5-
15 cm long, hirtellousto subhirsute;spathe 12-17 cm This species is quite distinct because of long pe-
long, color unknown,puberulousto hirtellousoutside, duncles of the pistillate inflorescences. It does not
glabrousinside; spikes ca. 30-80, + crinkled, 4-15 show clear affinities to other species from the same
x 0.1-0.2 cm, with stipes ca. 0.4-1 cm long and region.
hairy; rachis hairy. Staminateflowers: perianth tu-
bular,ca. 0.5-0.7 mm long, with arachnoidindumen-
tum below the apex, the apex almost plane; stamens 28. Cecropia marginalis Cuatrecasas,Revista Acad.
not seen. Pistillate inflorescencesin pairs,pendulous; Colomb.Ci. Exact. 6(22/23): 285. 1945. Type.Co-
SYSTEMATICTREATMENT
lombia. Putumayo: Rio Putumayo, between Rio
Giiamues& PuertoAsis, 21 Dec 1940 (Y), Cua-
trecasas 11243 (holotype:COL; isotype: F).
Fig. 31
Tree,to 25(-40) m tall. Leafy twigs 2-5 cm thick,
green, hispidulous or hirtellous to hirsute. Lamina
subcoriaceousto chartaceous,ca. 50 X 50 cm to 85
X 85 cm (to 120 X 120 cm), the segments 7-8, the
free parts of the upper segments ovate, the incisions
down to 3/10-5/10; apices rounded; upper surface
(almost) smooth, almost glabrous,with sparsebrown
pluricellularhairs and sparse long, unicellularhairs;
lower surface densely to rather sparsely hirtellous
or subhirsute on the main veins, with arachnoid
indumentumin the areoles and on the smaller veins;
lateral veins in the free part of the midsegment6-15
pairs, submarginallyloop-connected, many of them
branched;petiole relatively short,in adult specimens
ca. half the diameterof the lamina, 25-45(-60) cm
long, densely hirsute, denser toward the apex, often
also with sparseto dense arachnoidindumentum;tri-
chilia separateor occasionally fused, the brown in-
dumentumintermixedwith dense shortor long white
hairs; stipules ca. 15-45 cm long, caducous, yellow-
ish to darkred to red-brownor pale green,with sparse
unicellularhairs at the base or confinedto the margin
or largely ? densely hirsute to subvillous outside,
glabrousinside; terminalbuds usually ? curved.Sta-
minate inflorescencesin pairs,the peduncleerect, the
spikes erect to pendulous; peduncle 6-9 cm long,
sparselyto densely hirtellousto subhirsute;spathe8-
13(-16) cm long, darkred to red-brownto orangeor
to pink, or pale green, puberulousor hirtellousto sub-
hirsute outside, glabrous inside; spikes ca. (20-)30-
50, 6-11 X 0.2-0.3 cm, with stipes 0.5-1.5 cm long
and tomentose; rachis hairy. Staminateflowers: per-
ianthdeeply 2-parted,1.2-1.3 mm long, glabrous,the
apex convex; filamentsswollen; anthers0.6-0.7 mm
long, without appendages,remainingattachedat an-
thesis. Pistillate inflorescences in pairs, sometimes
solitary, initially erect, pendulous in fruit; peduncle
7-8(-17) cm long, subhispidulousto hirtellous,more
densely so at the apex; spathe6-11(-14) cm long, the
color and indumentumas in the staminateinflores-
cence; spikes 4(-7), ca. 8-12 x ca. 0.5 cm, to 22 X
1.5 cm in fruit, sessile or with stipes to 0.5 cm long
and tomentose; rachis glabrous. Pistillate flowers:
perianth 1.3-1.5 mm long, with arachnoidindumen-
tum below the apex outside, absent inside, the apex
convex to subconical, sometimes 2-lobed, granulate
or muriculateand with a few short bristles near the
aperture,the apertureslit-shaped; style ratherlong,
111
almost straight;stigma penicillate. Fruit oblongoid,
ca. 2 mm long, smooth.
Distribution (Fig. 8.2). From Colombia (Meta)
throughAmazonianEcuadorto Peru (Amazonas),in
non-inundatedor occasionally or briefly inundated
places, mostly in secondary growth, at elevations to
ca. 1200(-1500?) m. It is the most common species
in AmazonianEcuador.
Representativespecimensexamined.COLOMBIA.
CAQUETA:Mun.Florencia, rd.to Gabinete,18Oct1993(Y
fl), Franco et al. 4508 (BG,COL).META: Mun.Villavicen-
cio, rd. to Guayabetal,900-1000 m, 13 Feb 1995 (Y fl-fr),
Franco et al. 4676 (BG, COL). PUTUMAYO:Mun. Puerto
Asis, rd. to Kanakas, 16 Feb 1995 (Y fl-fr), Franco et al.
4680 (BG, COL);rd. Mocoa-Pasto, nr.Mocoa, 18 Feb 1995
(Y fl-fr), Franco et al. 4686 (BG, COL, HUA, US); rd. Mo-
coa-Puerto Asis, 3 May 1994 (? fl-fr), Franco et al. 5518
(BG, COL), (6), Franco et al. 5519 (BG, COL).
ECUADOR.MORONA-SANTIAGO: Rio Blanco, be-
tween Macas & Sucua, 26 Jan 1981 (6), Berg 1217 (BG,
COL, QCA, U); rd. Zamora-Gualaquiza,Estaci6n Exp. El
Padmi,ca. 5 km N of Los Encuentros,4 Jan 1991 (Y fl-fr),
Berg et al. 1659 (BG, LOJA,QCA); Cord.de Cutucu,5-10
km E of Logronlo,1200-1500 m, 7-9 Oct 1975 (? fl-fr),
Little et al. 669 (COL, LOJA,Q, QAME, QCNE); 11.7 km
SW of Taisha, 11 Sep 1976 (Y fl-fr), Ortega U. 148 (Q).
NAPO: Rio Payamino, ca. 10 km NW Coca, 27 Feb 1980
(Y fl), Berg et al. 1091 (AAU, BG, COL, MO, QCA, U);
ReservaBiol6gica JatunSacha, 8 km E of Misahualli,4 Sep
1987 (6), Cer6netal. 2149 (BG, QAME,QCNE);rd.Tena-
Rfo Pano, km 6-12, 5 Aug 1984 (Y fl-fr), Dodson et al.
15011 (QCNE); San Jos6 de Payamino,40 km W of Coca,
Rio Tutapishu,5 Nov 1982 (Y fl-fr), Irvine 210 (BG, F,
QCA, QCNE);Archidona,1865 (6), Isern SPB (GB, MA);
rd. Puyo-Puerto Napo, km 18, 8 Apr 1969 (Y fl-fr), Lugo
1065 (BG, GB); ArmeniaVieja, Rio Napo, 12 km SW of
Coca, 12 Jan 1973 (6), Lugo 2650 (BG, GB); Cafi6nde los
Monos, 15 km N of Coca, Rio Coca, 18-20 Apr 1985 (Y
fl-fr), Neill 6337 (BG, MO, QAME, QCNE); Afiangu,
ParqueNacionalYasuni,30 Jun-9 Jul 1982 (st), SEF 10108
(AAU, BG, QCA). PASTAZA: Between Mera & Moravia,
ca. 1000 m, 16 Dec 1955 (Y fl-fr),Asplund18867 (S); Mera,
ca. 1100 m, 21 Dec 1955 (6), Asplund 18944 (S); Rio Bo-
bonaza, Cuansha-Nalpi,between Cabeceras & Canelos, 9
Feb 1971 (Y fl-fr), Lugo 1450 (BG, GB); Rio Bobonaza,at
Rio Tinguiza,SE of Pacayacu,15 Mar 1971 (6), Lugo 1685
(BG, GB); Rio Bobonaza, Puerto Ubilla, nr. Pacayacu, 17
Mar 1971 (d), Lugo 1705 (BG, GB). SUCUMBfOS:Rio Cuy-
abeno, ca. 0?20'S, 75?55'W,20 Feb 1980 (Y fl-fr), Berg et
al. 1072 (AAU, BG, COL, GB, K, MO, NY, QCA, TUR,
U), 1073 (BG, COL, QCA); Rio Aguarico, ca. 0?20'S,
75?05'W,21 Feb 1980 (6), Berg et al. 1074 (AAU, BG,
COL, GB, K, MO, NY, QCA, TUR, U, WIS);Rio Aguarico,
nr.confluencewith Rio Pavayacu,Sep 1981 (Y fl-fr),Bravo
et al. 291 (QCA); Rio Aguarico, Dureno, 22 Feb 1980 (Y
fl-fr), Holm-Nielsen et al. 21754 (AAU, MO). ZAMORA-
CHINCHIPE: Rio Nangaritza,nr. Nangaritza, 12 Dec 1990
112 FLORA NEOTROPICA

fX;~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~Vv
5z~ ~~~~~~~~~~~~~~~~~/

FIG. 31. Cecropia marginalis. 1. Leaf, reduced. 2. Apex of lamina and venation (Berg et al. 1072). 3. Stipules
(Dodson et al. 15011). 4. Pair of staminate inflorescences with spathes and base of petiole with trichilium. 5. Pair of staminate
inflorescences, one with spathe, the other at anthesis, and base of petiole with trichilium (Berg et al. 1074). 6. Staminate
flower and stamen (Franco et al. 4508). 7. Pistillate inflorescence with spathe and base of petiole with trichilium. 8. Pistillate
inflorescence at anthesis and base of petiole with trichilium. 9. Pistillate inflorescence after anthesis and base of petiole
with trichilium. 10. Pistillate flower. 11. Fruit with remnant of style (Berg et al. 1072). 12. Trichilia (Berg et al. 1072).
SYSTEMATICTREATMENT
(d), Neill et al. 9703 (AAU, BG, QCNE); Cant6n Nanga-
titza, Pachicutza,18 Oct 1991 ( Y fl-fr), Palacios et al. 8293
(BG, QCNE, USZ).
PERU. AMAZONAS: QuebradaAintami, 1 May 1973 (?
fl-fr), Kayap 680 (GH); QuebradaHuampami,5 Jul 1974
(Y fl), Kayap 1052 (BG); Rfo Santiago,nr.La Poza, 12 Nov
1979 (st), Tunqui16 (MO); Rfo Santiago, nr. Caterpiza,14
Dec 1979 (Y fl), Tunqui350 (MO); Prov.Bagua, Dtto. Im-
aza, Rio Comaina,Kusu-kubaim,17 Aug 1994 (d), R. Vds-
quez et al. 18822 (MO).
The species is quite uniformexcept for the length
of the hairs. It has longer hairs and often also denser
indumentumon the leafy twigs, stipules, spathes,pet-
iole, lower surface of the lamina, and in the trichilia
in Colombiathanin collections made southward.The
trichilia are occasionally fused (as in Franco et al.
5519). Cecropiamarginalisshows affinitiesto C. stri-
gosa.
Vernacular names. Ecuador:suu or tsu (Shuar,
Morona-Santiago);tselan dundu (Quichua, Napo).
Peru: sui, suu, tsake kumpari(Amazonas).
29. Cecropia maxima Snethlage,Notizbl. Bot. Gart.
Berlin-Dahlem 8: 360. 1923. Type. Ecuador.Pi-
chincha:Poluahua(= Pululahua?),( Y + d ?), So-
diro 2a and 3 (syntypes: B, destroyed,duplicates
not traced; photographs in F, G, GH, MO),
herewith replaced by: Ecuador. Pichincha: Rd.
Quito-Chiriboga-Empalme, km 50, 2000-2100
m, 21 Jul 1987 (Y), Zak et al. 2198 (neotype:
QCA; isoneotypes: BG, G, GB, GH, K, L, MO,
NY, TEX, US). Fig. 32
Tree,to 30 m tall. Leafy twigs 4-9 cm thick, dark
brown with pale brown to orange lenticels, sparsely
puberulousto hirtellousand with rathersparsearach-
noid indumentumandbrownpluricellularhairs.Lam-
ina coriaceous, ca. 40 X 40 cm to 90 X 90 cm, the
segments 10-15, the free partsof the uppersegments
subobovate to (ob)lanceolate, the incisions down to
5/10-8/10; apices rounded; upper surface (almost)
smooth, sparselyhirtellouson the main veins, on the
whole surface dense or sometimes (very) sparse
arachnoidindumentum;lower surface(initially) with
brown pluricellularhairs, with arachnoidindumen-
tum (almost) confined to the areoles, but (initially)
also present on the main veins; lateral veins in the
free part of the midsegment 15-27 pairs, submargin-
ally (to marginally)loop-connected,most of them un-
branched;reticulumwith ratherthick veinlets;petiole
ca. 50-90 cm long, red to red-brown,sparselypuber-
ulous and with brown pluricellularhairs, moreover,
with (rather)dense arachnoidindumentum;trichilia
113
fused, the brown indumentumintermixedwith dense
short whitish hairs; stipules ca. 20-40 cm long, dull
dark red, sparsely to densely hirtellous puberulous
and with dark brown pluricellularhairs or also ?
sparse arachnoid indumentum outside, densely
(sub)sericeous (or glabrous) inside. Staminateinflo-
rescences often solitary, the erect or the spikes ?
spreading;peduncle 6.5-12 cm long, (sub)glabrous;
spathe 10-15 cm long, dull red to purplish,hirtellous
and with brown pluricellularhairs or also arachnoid
indumentumoutside, sparsely hairy or glabrous in-
side; spikes 4-6(-8), 8-12.5 X 0.6-1 cm, (sub)-
sessile; rachiswith stiff hairsandcrinkled(arachnoid)
hairs. Staminateflowers sessile or shortlypedicellate;
perianthtubular,ca. 2-3 mm long, puberulousbelow
the apex, the apex plane to slightly convex with short
hairs near the aperture;filaments flat; anthersca. 1
mm long, appendiculate,detachedat anthesis(?). Pis-
tillate inflorescences solitary, erect, or the peduncle
deflexed in fruit, subtendedby caducous (or subper-
sistent) bractswith dense arachnoidindumentumand
to 8 cm long; peduncle 3-15 cm long, sparsely hir-
tellous and with brown pluricellularhairs and also
arachnoidindumentum;spatheca. 10-20 cm long, the
color and indumentumas in the staminateinflores-
cence; spikes (l-)2-3, 12-18 X ca. 1.5 cm, to 48 X
2.8 cm in fruit,(sub)sessile;rachisglabrous.Pistillate
flowers: perianth ca. 3-4 mm long, with arachnoid
hairs below the apex, absent inside, the apex slightly
convex, muriculateto puberulousaroundthe aperture;
style long, ? S-shaped,minutelypuberulous;stigma
comose. Fruit oblongoid, 2-2.5 mm long, smooth,
pale brown.
Distribution (Fig. 10.4). From western Ecuador
(Cotopaxi) to Colombia (Narifno),in montaneforest;
at 1500-2400 m, or in Narifioalso at 900-1200 m (in
vegetationwith the physiognomyof montaneforest).
Specimensexamined.COLOMBIA:NARINO: Mun.
Barbacoas, rd.El Barro-Junfn,
Cgto.Altaquer, El Mirador,
1150 m, 12 Dec 1993 (Y fl-fr), Franco et al. 5175 (BG);
Mun. Barbacoas, rd. Junfn-El Diviso, 900 m (!), 21 Feb
1995 (Y fr), Franco et al. 4694 (BG, COL, HA, US); rd
Mun. Mallama, QuebradaLa Chorrera,2350 m, 27 Dec
1993 (d), RamfrezP et al. 6061 (COL).
ECUADOR.COTOPAXI:Rd. Pilal6-Quevedo, between
Pilal6 & Macuchi, 1500 m, 29 Jul 1980 (st), Holm-Nielsen
et al. 24706 (AAU, BG). IMBABURA: Cotacachi, above
Apuela, 2000 m, 24 Apr 1992 (d), Palacios et al. 10036
(MO, QCNE).PICHINCHA:BetweenSanJuan&Chiriboga,
ca. 2250 m, 3 May 1955 (d), Asplund16167 (S); Guarumal,
ca. 2100 m, 20 Jan 1956 (d), Asplund19039 (S); rd. Quito-
San Juan-Chiriboga-Empalme,ca. 7 km before Chiriboga,
ca. 2300 m, 14 Dec 1990 (Y fl-fr), Berg et al. 1645 (BG,
QCA); Calacali, Reserva Geobotanicadel Pululahua,1600
114 FLORA NEOTROPICA

Q,~~~~~~~,,
Ip I

FIG. 32. Cecropia maxima. 1. Leaf, reduced.2. Apex of lamina and venation (Ervik36971). 3. Stipules and young
leaf (Franco et al. 4694). 4. Staminate inflorescence with spathe. 5. Staminate inflorescence and base of petiole with
trichilium(Palacios et al. 10036). 6. Staminateflower.7. Stamen (Ervik36871). 8. Pistillate inflorescencewith spatheand
base of petiole with trichilium(Cero'net al. 4776). 9. Pistillate inflorescenceafter anthesis. 10. Pistillate flower (Berg et al.
1645). 11. Fruit(Franco et al. 4694).
SYSTEMATICTREATMENT 115

m, 7 Sep 1988 (? fl-fr), Cer6n et al. 4776 (BG, MO, (sub)glabrousor subsericeous on the lower part in-
QCNE); rd. Nano-Mindo, Los Cedros, 2000 m, 27 Aug side. Staminate inflorescences solitary or in pairs,
1991 (d), Ervik 36871 (BG, QCNE); rd. Quito-Tandapi- sometimes subtended by caducous bracts, to 1 cm
SantoDomingo de los Colorados,9.5 km E of Tandapi,1900
long, the peduncledeflexed andthe spikespendulous;
m, 28 Sep 1979 (? fl-fr), Gentry et al. 26662 (AAU, BG,
peduncle 10-18 cm long, green, (sub)setose; spathe
MO).
28-45 cm long, white to pinkish, hirsute to
This white-leavedspecies is conspicuous in mon- (sub)setose and with dense arachnoidindumentum
tane forest on the western slopes of the Andes in Ec- outside, glabrousinside; spikes 12-32, 25-36 X ca.
uador and adjacent Colombia. However, in Nariiio 0.6-1 cm, with stipes 0.2-1 cm long and subsetoseor
(Colombia), the species also occurs at distinctly with arachnoidindumentum;rachis with short stiff
lower elevations (900-1200 m) in forest showing hairs and arachnoidindumentum.Staminateflowers
physiognomic similarities to montane forest. At with pedicels to 1.2 mm long, these hispidulous to
these low elevations the trees lack the typically white puberulousor glabrous;perianthtubular,1.5-2 mm
upper surface of the lamina as found at higher ele- long, puberulousto hispidulous below the apex, the
vations. The trees are usually inhabited by ants. apex plane, hispidulous,mainly along the marginsof
There is a clear discrepancybetween the presentde- the aperture;filamentsflat;anthers0.6-0.7 mm long,
scription of the staminateflowers and that by Sneth- appendiculate,detachedat anthesis,reattachedto the
lage (1923), who stated that the perianthis 3-4 mm marginsof the apertureby the appendages.Pistillate
long and anther2-3 mm long. Because of these fea- inflorescencessolitary,deflexedto pendulous;pedun-
tures, Cecropiamaximawas includedin sect. Tomen- cle green, 8-18 cm long; spathe ca. 30-40 cm long,
tosae. The differences in the descriptionscannot be the color and indumentumas in the staminateinflo-
explained. rescence; spikes 2 or 3, ca. 25-40 X ca. 2 cm, to 60
X 3 cm in fruit, (sub)sessile; rachis hairy only with
stiff short hairs. Pistillate flowers: perianth 3.5-4.5
30. Cecropia megastachya Cuatrecasas, Revista mm long, with arachnoidindumentumbelow the apex
Acad. Colomb. Ci. Exact. 6(22/23): 284. 1945. outside, also in the style channel inside, the apex ?
Type. Colombia.Valle: Piedrade Moler, 1000 m, convex, punctate to muriculate;style long, straight,
19 Jul 1943 (d + Y), Cuatrecasas 14910 (sheet minutely hairy; stigma comose. Fruit oblongoid to
with pistillateinflorescence,stipules,andleaf here subovoid, ca. 3 mm long, finely tuberculate,pale-
designatedas lectotype:COL;isolectotype:COL). brown.
Tree,to 17 m tall, often unbranched.Leafy twigs Distribution (Fig. 18.2). Colombia, on the west-
5-8 cm thick, pale green with conspicuous white to ern slopes of the western Cordillera,from Antioquia
pinkish lenticels, hirtellous to hirsute to subsetose. to Valle, in forest, at ca. 400-1300 m.
Laminacoriaceous, ca. 50 X 50 cm to 140 X 140 cm
Specimens examined. COLOMBIA.ANTIOQUIA:
(to 170 x 170 cm), the segments 10-12, the free parts
Mun. Frontino,Cgto. Nutibara,rd. Nutibara-LaBlanquita,
of the upper segments oblanceolate, the incisions
Murri region, 1250 m, 4 Mar 1995 (Y fl-fr), Franco et al.
down to 8/10-9/10 or to 1-2 cm from the petiole; 5555 (BG, COL, HUA, US), (d), Franco et al. 5556 (BG,
apices acute to subacuminate;upper surface smooth COL, HUA, US); Mun. Frontino,Cgto. Nutibara,valley of
with rather sparse arachnoidindumentumor (sub)- Rio Cuevas, 1000 m, 13 Apr 1987 (Y fl-fr), D. SdnchezS.
glabrous;lower surfacesparselyminutelypuberulous et al. 1125(MEDEL).CHOc6:Mun.Trujillo,rd.Trujillo-
on the main veins largely with brown pluricellular Naranjal,km 32, Rio Sanquininf,ca. 900 m, 11 Sep 1993
hairs), with (very) sparse to dense arachnoidindu- (Y fl-fr), Franco et al. 4485 (BG), (d), Franco et al. 4486
mentum in the areoles and on the smaller veins, ex- (BG); Mun. San Jose del Palmar,ca. 1 km W of San Jos6
tending to the main veins or not; lateralveins in the del Palmar,ca. 900 m, 19 Mar 1994 (cd), Franco et al. 4564
free part of the midsegment20-30 pairs, submargin- (BG, HUA), (Y fl-fr), Franco et al. 4565 (BG, HUA).
VALLE: Rio Anchicaya, nr. CVC hydroelectricplant, 400-
ally loop-connected, unbranched;reticulumveinlets
500 m, 16 Dec 1981 (Y fl-fr), Gentry 35680 (BG, COL,
thick; petiole 70-120 cm long, with ? sparsearach-
MO, TULV).
noid indumentumand (especially at the basal and
upper part) hirsute to subsetose or setose (with irri- The habitand the presenceof irritatingbristlesare
tating hairs); trichilia fused, the brown indumentum indicationsof close relationshipto Cecropiahispidis-
intermixedwith sparsebristles;stipulesca. 20-70 cm sima. Both species are not inhabitedby ants of the
long, partly pinkish to red, largely whitish, (sub)- genus Azteca, but by representativesof the genus Pa-
setose and with dense arachnoidindumentumoutside, chycondyla.
116 FLORANEOTROPICA

31. Cecropia membranacea Trecul, Ann. Sci. Nat. brown indumentum intermixed with sparse short
Bot., Ser. 3, 8: 83. 1847; Berg, Acta Amazonica white hairs; stipules 8-20(-26) cm long, usually
8(2): 174. 1978. Ambaiba membranacea(Trecul green, sometimes dull red, almost glabrousor (very)
Kuntze, Revis. Gen. P1. 2: 624. 1891. Type. Bra- sparsely hirtellous to hirsute outside, sparsely to
zil?. Without locality, Anonymous(possibly Fer- rather densely sericeous inside. Staminate inflores-
reira) s.n. (holotype:P). Fig. 33 cences in pairs,pendulous;peduncle4-6 cm long pu-
berulousto hirtellous;spathe 8-10 cm long, yellow-
Cecropia robustaHuber,Bol. Mus. ParaenseHist. Nat.
6: 61. 1910. Type. Brazil. Para: Lower Rio Ama- ish, hirtellousoutside, glabrousinside; spikes ca. 30-
zonas, Feb 1904 (Y), Huber MG 4169 (holotype: 40, 7-11(-16) X 0.2 cm, with stipes 0.5-1 cm long
MG; isotype: G). and hirtellous;rachis hairy. Staminateflowers: peri-
CecropiabifurcataHuber,Bol. Mus. ParaenseHist. Nat. anth cup-shaped,0.8 mm long, at the upperpartwith
6: 62. 1910. Type. Brazil. Amazonas:LowerRio Pu- stiff hairs;filamentsswollen; anthersca. 0.4-0.5 mm
rus, 13 Mar 1904 (v), Huber MG 4184 (holotype: long, not appendiculate,remainingattachedto the fil-
MG; isotypes: BM, G). ament at anthesis. Pistillate inflorescences in pairs,
Cecropia laetevirens Huber, Bol. Mus. Paraense Hist. pendulous; peduncle 5-15(-20) cm long, hispid,
Nat. 6: 63. 1910. Type.Brazil.Amazonas:Rio Purus,
sometimes only just below the spathe;spathe 12-18
below Boca do Acre, 28 Mar 1904 (Y), Huber MG
cm long, the color and indumentumas in the stami-
4237 (syntype:MG; isosyntypes:BM, G, U) andRio
Purus, Ponta Alegre, 17 Apr 1904 (st), Huber MG
nate inflorescence;spikes (2-)4, (7-)10-16 X 0.4-0.6
4522 (syntype: MG). cm, to 30 X 1.4 cm in fruit,with stipes 0.5-1 cm long
Cecropia vageleri Burret, Notizbl. Bot. Gart. Berlin- and hirtellous;rachisglabrous.Pistillateflowers: per-
Dahlem 9: 49. 1924. Type. Colombia. Middle Rio ianthca. 1.5-2 mm long, with arachnoidindumentum
Magdalenavalley, 13 Aug 1923 (Y), Vageler69 (ho- below the apex, also below the apertureinside, the
lotype: B, destroyed, duplicates not found, photo- apex slightly convex, minutely and often ? sparsely
graphsin F, G). muriculate;style ratherlong, straight;stigmapenicil-
Cecropia tessmannii Mildbraed, Notizbl. Bot. Gart. late. Fruit oblongoid, ca. 2.5 mm long, smooth.
Berlin-Dahlem 9: 260. 1925. Type. Peru. Ucayali:
Rio Ucayali,Yarinacocha,Tessmann3454 (holotype: Distribution (Fig. 8.6). Upper Amazon basin
B, destroyed,photographin F). (fromBolivia to Venezuela),the Pacificcoastalregion
Cecropia setico Snethlage ex Macbride, Publ. Field of Ecuadorand Colombia, extending to easternPan-
Mus. Nat. Hist., Bot. Ser. 13(2.2): 289. 1937. Type. ama, in the lower Rio Magdalenavalley (Colombia),
Peru.Loreto:Rio Marafi6n,nr.Uanana, 1 Aug 1924 and the lower Cauca valley (Colombia), common in
(Y), Tessmann3713 (holotype: B, destroyed, pho-
periodicallyinundatedplaces along (white-water)riv-
tographsin F, MO; isotypes: NY, S).
ers, also in non-inundatedplaces, but then often near
Cecropia occidentalis Cuatrecasas,Revista Acad. Col-
rivers, at elevations to ca. 1000(-1250) m.
omb. Ci. Exact. 6(22/23): 277. 1945; 7(27): t. 5.6, t.
6.2. 1947. Type. Colombia. Valle: Rio Calima, La Representative specimens examined. PANAMA.
Trojita,22 Feb 1944 (6), Cuatrecasas 16347 (lec- DARIEN: CerroPirre,Rio Pirre,S of El Real,750-1030m,
totype, Berg & FrancoRosselli, Fl. Ecuador48: 36. 11Aug 1962 (6), Duke 5355 (MO); Crucede Mono, 5 Nov
1993: COL; isolectotype: F). 1989 (Y fr), Fisher 55 (BG); nr. El Real, Rio Tuira,7 Nov
1989 (9 fl-fr), Fisher 56 (BG).
Tree,to 25(-35) m tall. Leafy twigs 2-4 cm thick,
COLOMBIA.AMAZONAS:Mun. Leticia, Cgto. Puerto
usually pale to dark green (to reddish), hispid to Santander,Rio Caqueta, Santa Isabel, 7 Apr 1994 (9 fl),
(sub)setulose. Lamina chartaceousto subcoriaceous, Ca'rdenaset al. 4599 (COL);between Leticia & Rio Ama-
ca. 40 X 40 cm to 80 X 80 cm, the segments (7-)8- cayacu, Isla Santa Soffa, 15 Aug 1972 (D fr), Idrobo 6518
10, the free parts of upper segments obovate, the in- (AAU, COL); Rio Igara-Parana,15 km below La Chorrera,
cisions down to ca. 5/10-7/10; apices acuminate;up- 18 May 1975 (d), Idrobo8315 (AAU, COL);Rio Caqueta,
per surfacesmooth, puberulouson the veins and with nr. mouth of QuebradaQuinch6,7 May 1988 (st), M. Sdn-
very sparsearachnoidindumentumor glabrous;lower chez S. et al. 280 (BG, U); Rio Caqueta,Isla El Tigre, 19
1988 (st), M. SanchezS. et al. 1321 (BG, U); Rio Ama-
surfacesparselyto densely minutelypuberulous,also Sep
zonas, above Leticia, Isla de Monos, 25 Apr 1987 (9 fl-fr),
with longer uncinatehairs or almost glabrouson the
Gentryet al. 57097 (BG, MO); Dtto. PuertoNarinio,Parque
veins, the arachnoidindumentumconfinedto the mar- Nacional Amacayacu, 2 Aug 1989 (d), R. Vdsquezet al.
gin, occasionally present in the areoles; lateralveins 12447 (COL, MO). ANTIOQUIA: ParqueNacional Natural
in the free part of the midsegment 10-13 pairs, sub- Los Katios, Rio Atrato, Isla de Tanggara,24 Oct 1979 (9
marginallyloop-connected,severalof thembranched; fl), Barbosa 1334 (COL); Mun. Necoclf, Reserva Indigena
petiole 40-75 cm long, glabrous;trichilia fused, the CaimanNuevo, Rio Caiman,Aug 1992 (6), Castano et al.
SYSTEMATICTREATMENT 117

FIG. 33. Cecropia membranacea. 1. Leaf, reduced. 2. Apex of lamina and venation (Berg et al. 1092). 3. Twig,
longitudinalsection. 4. Stipules (Oldeman et al. 90). 5. Pair of staminateinflorescences with spathes and base of petiole
with trichilium (Balslev 2280). 6. Staminateinflorescence at anthesis (Berg et al. 1090). 7. Staminateflower and stamen
(Ferreira s.n.). 8. Pair of pistillate inflorescences with spathes and base of petiole with trichilium. 9. Pair of pistillate
inflorescencesat anthesis. 10. Pistillate inflorescence after anthesis (Berg et al. 1092). 11. Pistillate flower. 12. Fruit(Berg
et al. P.17584). (From Fl. Ecuador48: 37. 1993, modified.)
118
200 (HUA); betweenVilla Arteaga& Chigorod6,1 Oct 1961
(6), Cuatrecasaset al. 26107 (COL, P, US), (9 fl-fr), Cua-
trecasas et al. 26113 (COL, P, US); Taraza',Cgto. El Doce,
28 Mar 1994 (9 fl), Franco et al. 4601 (BG, HUA); Rfo
Atrato,rd. to Codelsa, km 9-10 from entrance,11 Mar 1970
(9 fl-fr), Idrobo 6318 (COL); Casabe, 30 Oct 1979 (9 fl-
fr), Renterfa et al. 1929 (COL, HUA, JAUM, MO). BOL-
iVAR: Rio Magdalena, 18 Dec 1969 (d), Idrobo 6300
(COL), (9 fl), Idrobo 6301 (COL). CAQUETA: Mouth of
Rfo Yari, nr. Araracuara,15 Jul 1986 (9 fl-fr), Berg et al.
1563 (BG, COL);rd. PuertoRico-San Vincentedel Caguan,
4 Oct 1975 (9 fl), Cabrera 3753 (F); Rio Orteguaza,Ve-
necia, 31 May 1940 (9 fl), Cuatrecasas 8953 (COL, F);
Florencia,El Venado,25 Jan 1969 (9 fl-fr), Cuatrecasaset
al. 27240 (COL, NY, US); Rfo Caqueta,Araracuara,Centro
Experimentallas Penias,14 Mar 1991 (9 fl), Franco et al.
3349 (COL);ca. 25 km N of Solano, on Rio Caqueta,8 km
SE of TresEsquinas,below mouthof Rio Orteguaza,14 Mar
1945 (st), Little et al. 9787 (COL, US); Morelia,2 Oct 1941
(6), SneidernA.1053 (A, MICH, NY, S, US). CAUCA: Nr.
San Francisco,23 Sep 1968 (9 fl), Espinal 2746 (MEDEL).
CESAR: Rfo Magdalena,15 Oct 1969 (6), Irusta & Fortoul
Ltd. s.n. (COL). CHOC6:Rfo San Juan, Quebradadel Ta-
paral, 30 May 1946 (st), Cuatrecasas 21511 (F, VALLE);
Rfo San Juan,between Palestina& Agua Negra, 1 Jun 1946
(9 fl), Cuatrecasas 21577 (F, VALLE);Rfo Atrato, below
Quibd6,3 Apr 1958 (st), Cuatrecasaset al. 24203 (US); Rio
Salaquf,17-29 Oct 1989 (d), Mahechaetal. 6244 (UDBC);
Rio Taparal,tributaryof Rfo San Juan, 15 Nov-6 Dec 1979
(6 fl-fr), Rooden et al. 675 (BG, NY, U); Nuquf, Alto de
Buey, 1000 m, 26 Jun 1940 (6), Sneidern A.72 (A, LL,
MICH, NY, US). META: Rio Meta, Bocas del Tua, 6 Nov
1938 (st), Cuatrecasas4490 (COL, F, S); nr. La Macarena,
11 Feb 1995 (6), Franco et al. 4669 (BG, COL, HUA, MO,
US); Reserva Nacional La Macarena,Rfo Guayabero, 13
Mar 1959 (6), JaramilloM. 2153 (COL);nr.PuertoLopez,
30 Jul 1944 (9 fl-fr), Little 8337 (COL, US); ParqueNa-
cional Natural Tinigua, SerranfaChamusa,Apr 1991 (6),
Stevenson 294 (COL). NORTE DE SANTANDER:Rio Mag-
dalena,mouth of Rfo Lebrija, 17 Dec 1969 (9 fl-fr),Idrobo
6298 (AAU, COL), (6), Idrobo 6299 (COL). PUTUMAYO:
Rfo Putumayo,PuertoOspina, 29 Nov 1940 (9 fl-fr), Cua-
trecasas 10867 (COL,F); Rfo San Miquel, betweenBermeja
& Rio Conejo, 13 Dec 1940 (9 fl-fr), Cuatrecasas 11063
(COL, F); Mun. Orito, Vrda. El Parafso, 17 Feb 1995 (d),
Franco et al. 4683 (BG, COL, HUA, LP, MO, NY, US);
Mocoa, Jul 1989 (9 fr), Mahecha et al. s.n. (UDBC 7853);
Rfo Caucaya,nr.PuertoJaramillo,18 May 1942 (6), Schul-
tes 3778 (COL, GH). SANTANDER:Rio Magdalena,nr.Bar-
ranca Bermeja, 20 Apr 1926 (st), Bosse' 7633 (LE); Mun.
Barrancabermeja, Cgto. Chucurf,23 Apr 1983 (d), Moreno
I (COL). VALLE: Mun. Buenaventura,Cgto. El Tigre,Que-
bradaTatama,26 Oct 1979 (9 fl-fr), CuadrosV 891 (MO,
TULV), (d), CuadrosV 892 (MO, TULV);Rio Calima,La
Trojita,22 Feb 1944 (9 fl-fr), Cuatrecasas16349 (VALLE);
Rfo Calima, Pail6n, 23 May 1946 (st), Cuatrecasas21242
(F, VALLE);Mun.Versalles,Vrda.El Cedro, 1000 m, 1 Sep
1983 (9 fl-fr), Devia 301 (COL, TULV);Mun. Trujillo,rd.
Trujillo-Naranjal,Rfo Sanquininf,1250 m, 11 Sep 1993 (9
FLORANEOTROPICA
fl-fr),Francoet al. 4487 (BG); Buenaventura,7-9 May 1922
(st), Killip 5312 (NY, US); Mun. Buenaventura,San Isidro,
15 Nov-6 Dec 1979 (Y fl-fr), Rooden et al. 341 (BG, NY,
U), (d), Rooden et al. 344 (BG, MO, NY, U). VAUPES:
Upper Rio Guaviare, 9 Nov 1939 (9 fl-fr), Cuatrecasas
7617 (COL, US).
VENEZUELA. AMAZONAS:Nr. San Carlosde Rfo Ne-
gro, 14 Dec 1978 (9 fl-fr), Clark 6880 (MO, QCNE); San
Carlosde Rio Negro, 21 Mar-17 Apr 1988 (9 fl), Delascio
et al. 9304 (VEN); base of CerroDuida, Jan-Feb 1969 (st),
Farifias 349 (MO, NY); 5-6 km NE of San Carlos de Rio
Negro, 12 Apr 1979 (9 fl-fr), Liesner 6573 (MO, U); San
Carlosde Rio Negro, ca. 20 km S of confluenceof Rio Negro
& Brazo Casiquiare, 1986-1988 (6), Uhl 520 (MO); So-
lano, Lower Rio Casiquiare,10 Mar 1942 (9 fr), LI. Wil-
liams 14733 (US).
ECUADOR. Los Rios: Rio Palenque Biological Sta-
tion, rd. Quevedo-SantoDomingo de los Colorados,km 56,
2 Oct 1976 (st), Dodson et al. 6235 (QCA). NAPO:Between
Tena & Archidona,12 Oct 1939 (9 fl-fr),Asplund9261 (S);
Rio Payamino,nr. Coca, 27 Feb 1980 (c), Berg et al. 1090
(AAU, BG, COL, MO, NY, QCA, TUR, U), (9 fl-fr), Berg
et al. 1092 (AAU, BG, COL, MO, QCA, TUR, U); Reserva
Biol6gica JatunSacha, 8 km E of Mishualli,29 Jan 1989 (9
fl), Ceron 6081A (BG); Rio Napo, SantaRosa, 28 Apr 1972
(d ), Lugo 2003 (BG, GB); Rio Sumino,tributaryof the Rio
Napo, ca. 5 km NE of Santa Rosa, 2 May 1972 (4), Lugo
2104 (BG, GB); Mun. Orellana, Parque Nacional Yasuni,
Rio Indillama,Comuna Pompeya, 26 Aug 1992 (9 fl-fr),
Neill et al. 10107 (BG, QCNE), (6), Neill et al. 10130 (BG,
QCNE); Cononaco, 27 May 1976 (9 fl), Oldemanet al. 90
(QCA); ParqueNacional Yasuni,Afiangu,nr. mouth of Rio
Aniangu,30 Jun-9 Jul 1982 (st), SEF 10099 (AAU, QCA,
QCNE,U). PASTAZA:Lorocachi,Rio Curaray,29 May 1980
(9 fl-fr), Jaramillo 31352 (AAU). SUCUMBiOS:Nr. mouth
of Rio San Miguel, Aug 1980 (9 fl-fr), Andrade 33149
(AAU); nr. Limoncocha, 9 Apr 1982 (6), Balslev 2280
(QCA, NY); Reserva Faunistica Cuyabeno, nr. Laguna
Grande, 1988-1990 (st), Christensen 82750 (AAU); Rfo
Aguarico, 75?22'W,0?37'S, island midway between Zan-
cudo & LagartoCocha, 30 Aug 1979 (6 + 9 fl-fr), Holm-
Nielsen et al. 20205 (AAU); SantaCecilia, rd. Lago Agrio-
Baeza, ca. 16 km W of Lago Agrio, 27 Feb 1973 (6), Lugo
3542 (BG, GB).
PERU. AMAZONAS: Rio Numpatkin, Kusu, 10 Mar
1975 (9 fl-fr), Ancuash 50 (GH); Prov. Bagua,Yamayakat,
29 Jan 1996 (9 fl-fr), Jaramillo et al. 994 (MO); Rio Ce-
nepa, QuabradaHuampami,16 May 1973 (9 fl-fr), Kayap
805 (F, GH, USM). Cuzco: Prov. Paucartambo,Pilcopata,
9 Feb 1975 (9 fl-fr), Plowman et al. 5100 (GH, US); Prov.
Quispicanchis,between Suamsari& Fortaleza,9 Dec 1966
(9 fl-fr), Vargas C. 18486 (US). HUANUCO:Nr. Tingo
Maria,Cueva de las Lachusas, 16 Jul 1957 (9 fl-fr), Ellen-
berg 2385 (U); Prov. Puerto Inca, Dtto. Yuyapichis,DAN-
TAS, 1-15 Dec 1988 (9 fl), Kroell S. 190 (G). LORETO:
Rd. JenaroHerrera-Angamos,km 10, 18 Jul 1988 (9 fl-fr),
Berg et al. 1600 (BG, UMS); nr. mouth of Rio Nanay, 29
Sep 1972 (9 fl-fr), Croat20818 (F,MO, NY); Prov.Maynas,
Rio Gueppi,above PuertoPeru, 14 May 1978 (9 fl), Gentry
SYSTEMATICTREATMENT
et al. 21817 (F, G, U, USM); Prov.Maynas,Rio Amazonas,
between Yanomono& mouth of Rfo Manati, 29 Jun 1983
(Y fl-fr), Gentry42327 (BG, MO);Allapahuayo,39 km SW
of Iquitos, 24 Feb 1988 (d), Gentryet al. 61909 (MO); Rio
Sucusari, nr. Rio Napo, 15 Jul 1981 (Y fr), Hahn 6 (MO);
Iquitos,23 Feb 1924 (Y fl-fr),Kuhlmann1486 = RB 19903
(RB); Prov.Alto Amazonas,Rio Huasaga,nr.Wishintsa,16-
26 Jun 1986 (Y fl), Lewis 11206 (MO); Prov.Maynas,Yan-
amono, 21 May 1983 (Y fl), R. Vdsquezet al. 4136 (NY).
SAN MARTiN: Prov. San Martin,rd. Tarapoto-Lamas,ca.
km 7, 4 Dec 1997 (6), Berg et al. 1771 (BG, COL, MOL);
Prov. San Martin,rd. Tarapoto-Chazuta,km 13, nr. bridge
over Rio Mayo, JuanGuerra,6 Dec 1997 (Y fl), Berg et al.
1782 (BG, COL, MOL). MADRE DE DIos: Prov. Tambo-
pata,ComunidadNativo de Infierno,HermosaChica, 24 Feb
1989 (st), Alexiades et al. 353 (BG, MG, MO); Rfo Tam-
bopata,40 km upriverfrom PuertoMaldonado,15 Jan 1991
(Y fl-fr), Alexiades et al. 1097 (BG, K); Prov. Tambopata,
15 km ENE of Puerto Maldonado, 1991 (6), Fisher 177
(BG); ParqueNacional Manu,Rio Manu,Cocha CashuSta-
tion, 3 Nov 1986 (d), Foster 12084 (BG), 7 Nov 1986 (st),
Foster et al. 12202 (INPA, MOL); ParqueNacional Manu,
Rio Manu,PakitsaStation,26 Dec 1988 (Y fl), Fosteret al.
12825 (BG, LPB), (d), Fosteretal. 12827(BG, LPB);Prov.
Manu, Cocha Juarez, 28 Sep 1989 (Y fl-fr), Foster et al.
13284 (BG, F); Prov. Tambopata,Rio Heath, Peru-Bolivia
border, 2 Mar 1990 (? fl-fr), Gentry et al. 69785 (MO);
Prov. Manu, Rio Salvaci6n, 5 Dec 1991 (Y fl), Ntiuiezet al.
14627 (BG); Prov. Tambopata,Cuzco Amaz6nico, 1 Jun
1989 (Y), Phillips et al. 172 (MO). PAsco: Prov. Oxa-
pampa,Rfo Pichis valley, nr. Paujil, 10 km downriverfrom
PuertoBermudez,28 Sep 1982 (d), Fosteret al. 8948 (MO,
NY, U, USM). SANMARTfN: Rd. Tocache-TingoMaria,N
of El Progreso, 15 Mar 1979 (Y fl), Gentry et al. 25762
(BG, MO). UCAYALI: Prov. Coronel Portillo,Pucallpa,Pu-
callpillo, 8 Apr 1953 (Y fl-fr), Ferreyra9030 (USM); Prov.
Coronel Portillo, nr. Pucallpa,Isla Progresso,(6), Ferreyra
9044 (USM).
BRAZIL. AcRE: Upper Rio Moa, Fazenda Arizona,
24-30 1984 (st), Campbellet al. 8197 (BG); Serrade Moa,
30 Apr 1971 ( fl-fr), Maas et al. P 12674 (INPA, K, MO,
NY, U, US); Rio Jurua-Mirim,nr. Porangaba,24 May 1971
(Y fl-fr), Maas et al. P.13281 (INPA, MO, K, NY, P, S, U,
US); Mun. Mancio Lima, Rio Moa, Parque Nacional da
Serra do Divisor, 13 May 1996 (Y fr), M. Silveira et al.
1327 & 1329 (NY). AMAZONAS: Rio Solimoes, S of Ma-
naus, 2 Oct 1973 ( fr), Berg et al. P 17584 (C, COL,INPA,
MG, NY, P, U, US); Rio Solimoes, 10 km E of Boca de
Januaca,2 Oct 1973 (Y fl-fr), Berg et al. P 17585 (F, INPA,
K, NY, S, U); Mun. Humaita,Rio Madeira,Igarapedo Ban-
heiro, 2 km N of Humaita,2 May 1975 (Y fl-fr),Gottsberger
35-2575 (INPA);Tef6, Rio Solimoes, 23 Jul 1972 (Y fl-fr),
Kriegeret al. 12284 (GUA, INPA);Mun. Sao Paulo de Oli-
ven,a, nr. Palmares, 11 Sep-26 Oct 1936 (9 fl), Krukoff
8571 (A, BM, F, G, K, MICH, MO, NY, P, S, U, US); Rio
Solim6es, nr. Manaquiri,27 Jul 1979 (9 fl-fr), Rodrigues
8892 (INPA);Rio Solimoes, Lago do Capitari,25 Jun 1988
(9 fr), Ziburski88/50 (INPA). PARA:Mun. Afua, Rio Ca-
juuna, 4 Dec 1992 (st), Maciel et al. 2113 (MG).
119
BOLIVIA. BENI: Prov.Moxos,Rio Chirizi,130km S
of San Ignacio, 12 Oct 1991 (st), Del Aguila et al. 168
(BOLV);Prov.Yacuma,Estaci6nBiol6gicaBeni,Rio Cu-
riraba,25 Dec 1987 (9 fl), Moraes R. 982 (BG, LPB, MO,
km 38,
USZ);Prov.Ballivian,rd.Yucumo-Rurrenabaque,
7-14 Apr 1989 (9 fl-fr), D. N. Smith et al. 12897 (LPB,
km 35,
MO);Prov.Ballivian,rd.Yucumo-Rurrenabaque,
Mar-Jul 1990 (st), D. N. Smith et al. 14349 (BG, MO).
COCHABAMBA: Prov.Carrasco,Estaci6nValle del Sajta
UMSS, 29 Oct 1991 (st), Atahuachiet al. 23 (BOLV,USZ);
nr.Chipiri,ca. 5 kmN of VillaTunari,6 Dec 1995(9 fl),
Berget al. 1729 (BG,BOLV,COL,LPB,USZ).LA PAZ:
Nr.Sapecho,26 Feb 1994(9 fl-fr),Berg1698 (BG,COL,
LPB).PANDO: Prov.NicolasSuarez,"enla zonadel Na-
reuda,juntoa la barracaSanJose,"12 Jan1983(d), Fer-
ndndezC. et al. 8203 (G);Prov.Manupiri,
nr.Independen-
cia, 23 Aug 1985(st),MoraesR. 263 (BG,LPB).SANTA
CRUZ: Prov.Ichilo,Rio Ichilo,ReservaForestalEl Chor6,
15 Mar 1990 (9 fl-fr), Saldias 1038 (USZ).
Cecropia membranaceaand C. litoralis are the
only lowland species representedboth in the Amazon
basin and in the Pacific Coastal region. The species
is abundantalong white-waterrivers,as a component
of early successional stages on well-aeratedsoil (cf.
Lamotte, 1992). However, it can also be found in
flood plain forest, although often in wet places and/
or inundatedfor shortperiods,or (less often) faraway
from rivers. In such places the species is often less
ephemeral and can become large trees. The white
arachnoidindumentumon the lower surface of the
laminais normallyconfinedto the margin;sometimes
some sparsearachnoidis presentin the areoles, but it
soon disappears. However, in Cuatrecasas et al.
26107 from Antioquia (Colombia) the arachnoidin-
dumentumin the areoles appearsto be more persist-
ent. Some collections from Ecuador(Cer6n 712 and
Hurtado et al. 2403), erroneouslyincluded in C. id-
roboi by Berg & FrancoRosselli (1993: 27), and one
from Colombia, Putumayo(Franco et al. 4679), are
distinct by the presence of minute arachnoidindu-
mentumin the areoles of the lamina, which is thicker
(coriaceousto subcoriaceous)thannormalin C. mem-
branacea. They largely match C. membranaceain
other morphological vegetative parts. Collections
with staminateinflorescences,which could make af-
finitiesclearer,arewanting.The collections examined
have not been madefrom the typical C. membranacea
habitat,and they may representa type with aberrant
phenology. They are presently included in C. mem-
branacea with some doubt.
Specimens examined. COLOMBIA. PUTUMAYO:
6 Feb1995(9 fl-fr),Franco
PuertoAsis,on rd.to Kanakas,
et al. 4679 (BG,COL).
ECUADOR. NAPO: ReservaBiol6gicaJatunSacha,8
120 FLORANEOTROPICA

km E of Misahualli, 17 Jan-6 Feb 1987 (Y fl-fr), Cer6n 712 to purplish brown with conspicuous lenticels (or
(BG, QCNE), 22 Feb 1990 (st), Ceron et al. 8865 (BG); white), densely hispidulousto puberulouswith curved
Cant6n Archidona,rd. Hollin-Loreto, km 65, Huaticocha, to straighthairs,sometimesalso with dense arachnoid
14-19 Jul 1989 (9 fl-fr), Hurtadoet al. 2403 (BG, QCNE).
indumentum.Lamina chartaceousto subcoriaceous,
The incomplete collection from Peru. Loreto, ca. 15 X 15 cm to 60 X 60 cm, the segments (7-)8-
Santa Rosa, nr. Yurimaguas, 11 Nov 1929 (9 fl), Ll. 10(-1 1), the free parts of upper segments elliptic to
Williams 4949 (F), for which the vernacular name oblong to (sub)obovate(when juvenile often ? lo-
"pungara" is noted, might also belong to this type. bate), the incisions down to 6/10-8/10; apices obtuse
In addition to this type of Cecropia membranacea to short-acuminate; upper surface ? scabrous,
with arachnoid indumentum on the lower surface of sparsely to ratherdensely hispidulous and (initially)
the lamina, two other types have to be mentioned: one with sparse arachnoid indumentum;lower surface
indicated as "pungara" in southern Amazonian Peru (minutely) puberulous with longer uncinate hairs,
(Madre de Dios) and the other, which can be indicated arachnoid indumentum in the areoles and on the
as "herrerensis," in northern Amazonian Peru (Lor- smaller veins, extending to the main veins; lateral
eto); see page 220. The latter is probably represented veins in the free part of the midsegment9-16 pairs,
by Berg et al. 1600, which was not made in a typical marginallyloop-connected,mostly branched;petiole
C. membranacea habitat and distinct by reddish stip- 10-50 cm long, densely to sparselypuberulousto hir-
ules. With the very scarce and incomplete material tellous and with ? dense (or sparse, occasionally
available, these two types can hardly or not be distin- very sparse) arachnoidindumentum;trichilia fused,
guished morphologically from typical C. membra- the brownindumentumintermixedwith shortor often
nacea. They may produce trichilia earlier than normal long white hairs; stipules 6-13 cm long, white, hir-
in the species and they are not inhabited by Azteca tellous to subhirsuteand with dense arachnoidindu-
ants. The ecology of these types (may) deviates more mentumoutside, sericeous inside. Staminateinflores-
or less, also with regard to light regimes, and conse- cences in pairs or solitary, the peduncle erect to +
quently habitat preferences (Davidson & Fisher, deflexed, the spikes erect to pendulous;peduncle 6-
1991). Cecropia membranacea is in cultivation in the 15 cm long, ? densely puberulousto hirtellousand
Botanical Garden of Universidad Nacional Agraria, with ? dense (or sparse) arachnoid indumentum;
La Molina, Lima (Peru). spathe 7-11 cm long, white, hirtellousto subhirsute
Vernacular names. Colombia: guanibarae, uda- or to puberulousand with (very) dense arachnoidin-
gomo (Mui., Amazonas); yarumo corante (Ama- dumentum outside, sericeous inside; spikes (4-)6-20,
zonas); guanibarae, yarumo (Caqueta'); yarumo ma- 3-12 X 0.2-0.3 cm, with stipes 0.1-0.8 cm long and
cho (Choco). Venezuela: yagrumo rojo (Amazonas). densely puberulous or with arachnoidindumentum;
Ecuador: bokarawa (Auca, Napo); dondo, tsitica rachis hairy. Staminate flowers: perianthtubular,ca.
dundu (Quichua, Napo). Peru: setico blanco (Huain- 1 mm long, glabrous,the apex plane to slightly con-
uco); suui, sutl, sutlk, or sutiik (Jivaro, Loreto); ka'a- vex; filaments flat; anthersca. 0.5 mm long, appen-
bona or taa-bona (Ese-ejhal, Madre de Dios). Bolivia: diculate, detached at anthesis, reattachedto the mar-
ambaibo blanco (Beni). gins of the apertureby the appendages. Pistillate
inflorescencesin pairs or solitary, erect to deflexed,
becoming pendulousin fruit;peduncle3-12(-17) cm
long, ? densely puberulousto hirtellousto hispidu-
32. Cecropia metensis Cuatrecasas, Revista Acad. lous and with ? dense (or sparse) arachnoidindu-
Colomb. Ci. Exact. 9(36/37): 338. 1956. Type. Co- mentum, sometimes subglabrous; spathe 8-19 cm
lombia. Meta: Puerto L6pez, 29-30 Jul 1946 (9 long, the color and indumentumas in the staminate
fl-fr), Jaramillo M. et al. 378 (holotype: US; iso- inflorescence;spikes 2-4, 5-9 X 0.4-0.6 cm, to 16
type: COL). X 1.3 cm in fruit, sessile (or with stipes 0.5-0.8 cm
Cecropiapeltata Linnaeusvar.candidaVelasquez,Acta long and sparselypuberulousand with arachnoidin-
Bot. Venez. 6: 54, t. 12. 1971. Syntypes.Venezuela. dumentum);rachis glabrous(but hairs at the base of
Guarico: Nr. Calabozo, Morichal Villasmil, nr. Es- the free partof the perianth).Pistillateflowers basally
taci6n Biol6gica de Los Llanos, 8 Aug 1966 (9), connate; perianth ca. 1.5 mm long, with arachnoid
Veldsquez82 (VEN; isosyntype:US), (d), Veldsquez indumentumbelow the apex outside, absent (or pres-
83 (VEN; isosyntype: US).
ent in the style channel)inside, the apex slightly con-
Tree, to 12(-20) m tall, the trunk with prominent vex; stigma peltate. Fruit oblongoid, 1.8-2 mm long,
stipular scars. Leafy twigs 1-3 cm thick, dark green tuberculate.
SYSTEMATICTREATMENT
Distribution (Fig. 18.2). The Ilanosregion of Co-
lombia and Venezuela and savannas of Guyana, in
(white sand) savannas, gallery forests and swamps
(morichales),at low elevations.
Representative specimens examined. COLOMBIA.
CASANARE: Orocue, CanloSan Miguel, Jul 1989 (9 fl-fr),
Mahecha 5588 (UDBC), (5), Mahecha 5589 (UDBC).
META: Mun. Remolinos, CentroRecreacionalCafamllanos,
CafnoMatamata, 30 Sep 1995 (5), Bottia I (COL); rd.
Puerto Gaitan-San Pedro de Arimena, Alto Neblinas, 15
Mar 1986 (5), Foreroet al. 10239 (COL);Mun.Villavicen-
cio, rd. Villavicencio-PuertoL6pez, 10 Feb 1995 (9 fl-fr),
Franco et al. 4655 (COL), (5), Franco et al. 4656 (COL);
Mun. Puerto L6pez, Alto Menagua, 10 Feb 1995 (5),
Franco et al. 4660 (COL); Mun. Puerto L6pez, Puenta La
Balsa, 10 Feb 1995 (9 fl), Franco et al. 4661 (BG, COL,
HUA, MO, US), (juv), Franco et al. 4662 (COL,HUA, MO,
US); nr. La Macarena,11 Feb 1995 (5), Franco et al. 4465
(BG, COL, HUA, MO, US); nr. La Macarena,nr. rapidsof
Rio Guayabero,11 Feb 1995 (5), Franco et al. 4670 (BG,
COL, HUA, MO, US), (9 fl-fr), Franco et al. 4671 (BG,
COL, HUA, NY, US); San Juande Arama,Rio Giiejar,5-
20 Dec 1950 (5), Idrobo et al. 604 (US); San Martfn,22
Jan 1972 (5), Idrobo et al. 6489 (AAU, COL); PuertoL6-
pez, 27 Jul 1944 (9 fr), Little et al. 8242 (COL, F, P, US);
10 km S of Puerto Gaitan, 12 Mar 1971 (9 fl), Pinto et al.
1165 (COL, U); ParqueNacional NaturalTinigua,Serrania
Chamusa,Apr 1992 (5), Stevenson361 (COL);Rio Melda,
Finca La Florida, Nov 1993 (5), Stevenson545 (COL).
VENEZUELA. AMAZONAS:Depto. Atures,Rio Catan-
iapo, 25 Jul 1981 (5), Castillo 1337 (VEN); 8 km S of
Puerto Ayacucho, 13-15 Apr 1978 (9 fl-fr), Davidse et al.
14998 (MO, U, VEN); Depto. Atabapo, 15 km SW of Santa
Barbaradel Orinoco, 26 Jun 1979 (9 fl-fr), Huber 3869
(VEN). ANZOATEGUI: Bella Vista, Rfo Orinoco, 19 Nov
1970 (st), Morillo 113 (VEN); Rio Guaraguara,nr.Santome,
17 Jun 1941 (9 fl-fr), Pittier 14842 (US), (5), Pittier 14845
(VEN), 23 Jun 1941 (5), Pittier 14878 (US); nr. La Leona,
Rio Chives, 25 Jun 1941 (5), Pittier 14907 (VEN). APURE:
Dtto. Pedro Camejo, 22 km WNW of Buena Vista, 2-3 km
N of Isla Algarrobe, 17 Feb 1978 (9 fl), Davidse et al.
14302 (MO, U, VEN); Dtto. PedroCamejo,ca. 62 km NNE
of PuertoPdez, 28 Feb 1978 (5), Davidse etal. 14637(MO,
NY, U, VEN); Dtto. PedroCamejo, ca. 53 km NE of Puerto
Paez, 21 Feb 1979 (9 fl), Davidse et al. 15555 (MO, U,
VEN); Dtto. Pedro Camejo, 16 km NW of Mata de Guan-
abana,between Rio Meta & Rio Cinaruco,27 Feb 1979 (9
fr), Davidse et al. 15831 (MO, NY, U, VEN); Dtto. Pedro
Camejo/Anchagua,rd. San Fernando-PuertoPaez, between
Rio Capanaparo& CafioLa Guardia,7 Jul 1988 (d), G6mez
et al. 216 (VEN). BARINAS: Corozo, La Vizcaina, 5 May
1980 (5), MarcanoBerti et al. 94-980 (U). BOLiVAR: Mun.
Piar, Sector Las Patillas, Feb 1987 (9 fl), Fernandez2993
(BG, VEN). GUARICO: Morichal Largo, 17 km SE of Cal-
abozo, 15 Dec 1982 (5), Brito 83 (MO, NY, VEN); nr. Par-
mana, 3-10 Apr 1950 (d), Croizat 196 (F); Dtto. Miranda,
MorichalEl Recreo, Jul 1982 ( 9 fl-fr),Delascio et al. 11720
(VEN); nr. Calabozo, MorichalVillasmil, nr.Estaci6nBiol-
121
6gica de Los Llanos, (Y fl-fr), Veldsquez84 (US); rd. El
Sombrero-Calabozo,8 Aug 1966 (9 fl-fr), Veldsquez89
(US, VEN); nr. Calabozo, nr. Lagunade los Patos, Estaci6n
Biol6gica de Los Llanos, 8 Aug 1966 (9 fl-fr), Veldsquez
90 (US), (d ), Veldsquez91 (US). SucRE: Montafiade Mo-
chima, 20-21 km SW of Cumana, 16 Sep 1973 (9 fl-fr),
Steyermarket al. 108727 (MO, NY, VEN).
GUYANA. Nr. Aishalton Hospital, 19 Nov 1982 (st),
Stofferset al. 457 (BG, NY, U); Lake Awakawau,5 km SE
of Aishalton, 20 Nov 1982 (d), Stofferset al. 516 (U).
BRAZIL. RORAIMA: 12 km S of Boa Vista, 13 Apr
1989 (d), Guillaumetet al. 5850 (INPA),(9 fl-fr), Guillau-
met et al. 5851 (INPA).
Cecropia metensis is characterized by dense white
arachnoid indumentum on stipules, spathes, and usu-
ally also petioles and peduncles, and by its occurrence
in savanna regions. The presence of arachnoid indu-
mentum distinguishes C. metensis from C. sararensis
in most cases, and if not by this arachnoid indumen-
tum, then by the dense indumentum of straight to un-
cinate hairs on the petiole. Although the areas of C.
metensis and C. sararensis touch (or slightly over-
lap?) each other, intermediates between them are rare,
suggesting the presence of rather solid genetic barri-
ers. The collections Franco et al. 4655, 4656, and
4663 lack more or less dense arachnoid indumentum
on the petioles, but both the stipules and spathes are
covered by dense arachnoid indumentum. Moreover,
the relatively long hairs are not present in the trichilia
of Franco et al. 4663. Because of these features, these
collections could be regarded as transitional to C. sar-
arensis. Material of C. sararensis from Arauca also
shows features more or less transitional to C. meten-
Sis.
33. Cecropia montana Snethlage, Notizbl. Bot. Gart.
Berlin-Dahlem 8: 368. 1923. Type. Peru. San Mar-
tin: Tarapoto, Cerro de Escaler, 1300 m, Mar 1903
(9), Ule 6845 (holotype: B, destroyed, fragment
and photograph in F; isotype: MG, photographs in
F, GH). Fig. 34
Cecropialanciloba Cuatrecasas,RevistaAcad. Colomb.
Ci. Exact. 6(22/23): 292. 1945. Type. Colombia.Pu-
tumayo:Mocoa, Rfo Mulato,26 Dec 1940 (d), Cua-
trecasas 11308 (holotype:COL; isotype: F).
Cecropia alexandrina Cuatrecasas,Revista Acad. Col-
omb. Ci. Exact. 9(36/37): 339. 1956. Type. Peru.
Ucayali: San Alejandro,8 Apr 1953 (9), Ferreyra
9037 (holotype:US; isotype: USM).
Tree, to 25 m tall. Leafy twigs 1.5-4 cm thick,
green to red-brown or brownish to whitish, hispidu-
lous or hispid to subhirtellous. Lamina chartaceous to
subcoriaceous, ca. 35 x 35 cm to 85 x 85 cm, often
122 FLORA NEOTROPICA

~ ~~~~~~~~~~~~~~~~~~~~~~~~~~~~4
~~~~~~~~~~~4\*

10 ~9 3

FIG. 34. Cecropia montana. 1. Lamina, reduced.2. Apex of lamina and venation (Berg et al. 1661). 3. Young leaf
(Palacios 8178). 4. Base of petiole with trichilium(Neill et al. 9704). 5. Stipules (Berg et al. 1656). 6. Staminateinflorescence
with spathe (Palacios 8178). 7. Staminateinflorescence at anthesis and base of petiole with trichilium (Berg et al. 1661).
8. Staminate flower and stamen (Berg et al. 1656). 9. Pistillate inflorescence with spathe. 10. Pistillate inflorescence at
anthesis, opened prostoma,and base of petiole with trichilium(Neill et al. 9704). 11. Pistillate inflorescenceafter anthesis
(Berg et al. 1662). 12. Pistillate flower. 13. Fruit(Franco et al. 4684).
SYSTEMATICTREATMENT
reflexed in young leaves, the segments (1I-)12-15,
the free parts of upper segments oblanceolate(to su-
bobovate),the incisions down to ca. 7/10-9/10; apices
obtuse to subacute; margin with long, weak, white
hairs;upper surface scabrous,densely hispidulousor
partly hirtellous to strigose; lower surface (rather)
densely puberulousto hirtellousto subtomentoseand
with (rather)dense brown pluricellularhairs on the
main veins, rather sparsely subtomentose on the
smallerveins, with arachnoidindumentumvery short
in the areoles and much longer and loosely covering
the areasbetween the lateralveins, often extendingto
the lateral veins or the main veins; lateral veins in
the free partof the midsegment(25-)30-50 pairs,of-
ten >35 pairs, submarginallyto marginally loop-
connected, most of them unbranched;petiole ca. 30-
70 cm long, puberulousto hirtellous,towardthe apex
subhirsute;trichiliafused, the brownindumentumin-
termixedwith dense shortandratherlong white hairs;
stipules 5-12 cm long, white to pinkish,subhirsuteto
subsericeous to villous and with (very) dense arach-
noid indumentum outside, densely (sub)sericeous
inside. Staminate inflorescences in pairs, erect;
peduncle 6-12 cm long, hirtellous; spathe 4-9 cm
long, white to pinkish, hirtellous to subhirsuteand
with very dense arachnoidindumentumoutside, ?
densely to sparsely (sub)sericeous inside; spikes 8-
40, (0.8-)2-8 X 0.2-0.5 cm, with stipes 0.2-0.5 mm
long and sparsely puberulous to hirtellous or gla-
brous;rachis (sparsely)hairy.Staminateflowers with
pedicels to 0.2 mm long; perianthtubular,1-1.5 mm
long, puberulousor also with sparse arachnoidindu-
mentum below the apex, the apex almost plane, gla-
brous; filaments flat; anthers 0.5-0.7 mm long, ap-
pendiculate, detached at anthesis, reattachedto the
marginsof the apertureby the appendagesat anthesis
(?). Pistillate inflorescencesin pairs,erect, often sub-
tendedby caducousbracts, 1-2 cm long; peduncle2-
6(-9) cm long, puberulousto hirtellous; spathe 4-6
cm long, the color and indumentumas in the stami-
nate inflorescence;spikes 4-9(-20), 2-5.5 X 0.4-0.8
cm, to 1OX 1-1.5 cm in fruitand ? curvedandoften
broadened, sessile (or with short stipes). Pistillate
flowers: perianthca. 1.5 mm long, with arachnoidin-
dumentum below the apex outside, also below the
style channel inside, the apex + convex, muriculate;
style short; stigma comose. Fruit ellipsoid to oblon-
goid, 1.5-2 mm long, tuberculate.
Distribution (Fig. 8.7). Eastern slopes and foot-
hills of the Andes, disjunctly in Colombia (Putu-
mayo) and adjacentEcuador (Sucumbios) and from
Ecuador (Morona-Santiago) to Peru (Madre de
123
Dios), mostly in wet premontaneforest, at 400-1200
(-1500) m.
Specimensexamined.COLOMBIA. PUTUMAYO:Nr.
Mocoa, 18 Feb 1995 (Y fl-fr),Franco et al. 4684 (BG, COL,
HUA, US), (d), Franco et al. 4685 (BG, COL, HUA, US);
Mun. Mocoa, between La Campucana& San Antonio, 1000
m, 2 May 1994 (5), Franco et al. 5512 (BG), (Y fl-f),
Franco et al. 5516 (BG).
ECUADOR. MORONA-SANTIAGO:10 km W of Rio Za-
mora & 2 km S of Rio Bomboiza, nr.Misi6n Salesiana,800
m, 13 Mar 1986 (cTfl-fr),Baker6724 (BG, QAME,QCNE);
rd. Zamora-Gualaquiza,between Yantzatza& Los Encuen-
tros,ca. 900 m, 3 Jan 1991 (5), Berg etal. 1656 (BG, LOJA,
QCA); rd. Zamora-Gualaquiza,Estaci6nExp. El Padmi,ca.
5 km N of Los Encuentros,ca. 900 m, 4 Jan 1991 (Y fl-fr),
Berg et al. 1658 (BG, LOJA, NY, QCA); rd. Gualaquiza-
San Juan Bosco, nr. Gualaquiza,16 Feb 1994 (5), Berg et
al. 1691 (BG, QCNE). SUCUMBiOS: Lumbaqui,1000 m, 13
Aug 1975 (Y fl), Littleet al. 1681 (MO, Q, QAME,QCNE).
ZAMORA-CHINCHIPE:Nr. Zamora,ca. 1000 m, 4 Jan 1991
(5), Berg et al. 1661 (BG, GB, LOJA,NY, QCA), (Y fl-fr),
Berg et al. 1662 (BG, LOJA,QCA); Pachicutza,70 km NE
of Zamora, 1000 m, 14 Sep 1975 (? fl-fr), Little et al. 347
(COL, LOJA,Q, QAME); Cant6nNangaritza,Rfo Nangar-
itza, Pachicutza,900 m, 12 Dec 1990 ( Y fl), Neill et al. 9704
(BG, MO, QCA, QCNE);Cant6nNangaritza,Yanzatza,Rio
Yacuambi,17 Oct 1991 (5), Palacios 8178 (BG, QCNE).
PERU. HUANUCO: Rd. Tingo Maria-Aucayacu,Puma-
huasi, 650 m, 25 Nov 1997 (9 fl-fr), Berg et al. 1742 (BG,
COL, MOL); rd. Tingo Maria-Aucayacu,nr. Milagro, 650
m, 25 Nov 1997 (5), Berg et al. 1743 (BG, COL, MOL);
10 km S of Nuevo San Martin,60 km S of Shumanza,550
m, 16 Jul 1982 (9 fl-fr), Gentry et al. 37566 (BG, MO);
Prov. Leoncio Prado, Dtto. Rupa Rupa, E of Tingo Maria,
Cerro Quemado,700-800 m, 13 Sep 1978 (9 fl), Schunke
V 10595 (BG, MO). JUNIN: Prov. Chanchamayo,Catarata
del Tirol, 3 Feb 1999 (9 fr), NuiiiezI. 87 (BG). MADRE DE
Dios: Manu National Park, Yomybato vill., Aug 1997
(5), Yuet al. 1173 (BG). PASCO:Valley of Rfo Palcazu,
300-600 m, 7 Dec 1984 (5), Hartshorn et al. 2694 (F,
MO); Prov. Oxapampa, Gran Pajonal, trail Chequitavo-
Shumahuani,1200 m, 24 Sep 1983 (9 fl), D. N. Smith5225
(BG, MO). SAN MARTiN: Prov. Lamas, rd. Lamas-Pa-
mashto, ca. 800 m, 4 Dec 1997 (5), Berg et al. 1768 (BG,
COL, MOL); Prov. San Martin,rd. Tarapoto-Yurimaguas,
ca. km 13, ca. 700 m, 5 Dec 1997 (5), Berg et al. 1773
(BG, COL, MOL); Prov.Lamas,rd. Tarapoto-Moyobamba,
ca. km 62, ca. 1000 m, 8 Dec 1997 (9 fl-fr), Berg et al.
1788 (BG, COL, MOL); nr. Moyobamba,600-700 m, 18
Dec 1971 (5), Ferreyra17878 (US); Rio Huallaga valley,
between PuertoPinzana& PuntaArenas(Shumanza),4 Feb
1984 (9 fl-fr), Gentry et al. 44925 (BG, MO, NY); Prov.
Lamas, rd. Tarapoto-Moyobamba,km 55, 700 m, 10 Oct
1984 (d), Maas et al. 6035 (BG, U); Prov. Moyobamba,
Dtto. Calzada, Morro Calzada, 1050-1350 m, 11 Nov 196
(9 fl-fr), Sdnchez V et al. 8611 (BG). UCAYALI: Rd. Pu-
callpa-Tingo Maria,km 135, ca. 400 m, 26 Nov 1997 (5),
Berg et al. 1749 (BG, COL, MOL).
124
Cecropia montana is probably closely related to
C. angustifolia. It has also inflorescenceswith short
peduncles and spikes and the lamina is also (usually)
reflexed in the bud. It can be distinguishedfrom C.
angustifolia by the dense arachnoidindumentumon
the stipules (and spathes), by more numerouslateral
veins in the free partof the midsegmentof the lamina,
usually 30-50 pairs, whereas C. angustifolia (in the
non-villous type) normallyhas at most 30 pairs, and
usually also by the long white hairs intermixedwith
the brown indumentumof the trichilia (a featurenot
found in all specimens of C. montana, and rarelyin
C. angustifolia, e.g., in Berg 1233 and Franco et al.
5515). Moreover, the length of the stipules can be
used to distinguishthe two species (especially in the
northernpart of the range of distributionof C. mon-
tana): to 12 cm long in C. montana and often >12
cm long in C. angustifolia.Intermediatesbetweenthe
two species have not been encountered. Cecropia
montana is ecologically different from C. angusti-
folia, occurringat lower elevations.
34. Cecropia multisecta P. Franco& C. C. Berg, Cal-
dasia 23(1): 77, t. 1. 2001. Type. Colombia. An-
tioquia:Mun. Anon',rd. Anorf-Medellin,between
Vrda.El Roble & Anon', 1600 m, 8 Mar 1995 (Y),
Franco et al. 5573 (holotype:COL;isotypes: BG, Mun. San Rafael,nr.campamentoVersalles,1350 m, 23 Mar
COL, HUA). Fig. 35 1994 (Y fl-fr),Francoet al. 4574 (BG, HUA), (d), Franco
et al. 4575 (BG, HUA); Mun. Anorf, rd. Anori-Medellin,
Tree,to 13 m tall. Leafy twigs 1-4.5 cm thick,dark between Vrda. El Roble & Anorf, ca. 1600 m, 8 Mar 1995
green to brown or white to grayish, sparsely to (Y fl-fr), Franco et al. 5569 (BG, COL, HUA), (d), Franco
densely puberulousto hirtellousto subhispid(ulous), et al. 5572 (BG, COL, HUA).
also with rathersparse to dense arachnoidindumen-
tum and sparse to dense brown pluricellularhairs.
Lamina coriaceous, ca. 25 X 25 cm to 65 X 65 cm,
the segments 10-12, (ob)lanceolate, the incisions
down to the petiole; apices obtuse to acute to shortly
acuminate;upper surface smooth, sparsely hirtellous
to strigillose (on the veins); lower surface with ?
dense brown pluricellularhairs on the veins or also
sparsely puberulouson the main veins, with arach-
noid indumentumin the areoles; lateral veins in the
free partof the midsegmentca. 20-25, submarginally
loop-connected, unbranched;petiole ca. 25-55 cm
long, with dense arachnoidindumentum;trichiliasep-
arate or fused, the brown indumentum intermixed
with rather short whitish to brownish (unicellular)
hairs; stipules ca. 15-27 cm long, pink to yellowish,
sparsely to densely hirtellous to subsericeous and
with dense brownpluricellularhairsoutside, glabrous
or towardthe apex subsericeousinside. Staminatein-
florescences in pairs, the peduncle erect, the spikes
erect to ? spreading;peduncle 5-6 cm long, hirtel-
FLORANEOTROPICA
lous to puberulous and with dense to rathersparse
arachnoidindumentum;spathe ca. 12-13 cm long,
pink to greenish, + densely puberulousto hirtellous
outside, glabrous inside; spikes ca. 9-12, ca. 10-12
X 0.5 cm, with stipes 0.2-0.3 mm long and with
brown pluricellularhairs; rachis short-hairy.Stami-
nate flowers: perianth tubular, 1.8-2.2 mm long,
sparselyand minutelypuberulousbelow the apex, the
apex slightly convex;filamentsflat;anthers0.8-1 mm
long, appendiculate.Pistillate inflorescencesin pairs,
erect to pendulous;peduncle 3-6 cm long, hirtellous
to puberulous,and with + dense arachnoidindumen-
tum; spatheca. 14-15 cm long, pink, densely puber-
ulous to subsericeousoutside, glabrousinside; spikes
2-3(-6), ca. 11-12 X 1-1.3 cm, to 23 X 2 cm in fruit,
sessile; rachis short-hairy.Pistillateflowers: perianth
1-1.5 mm long, with arachnoidindumentumbelow
the apex outside, also in the lower part of the style
channel inside, the apex convex, punctateto muricu-
late or aroundthe apertureminutelypuberulous;style
rather short, minutely puberulous; stigma comose.
Fruit ellipsoid to oblongoid, ca. 2.5-3 mm long, tu-
berculate.
Distribution (Fig. 11.3). Colombia (Antioquia),
in forest, at 1300-1600 m.
Specimens examined. COLOMBIA. ANTIOQUIA:
This species shows affinitiesto Cecropiaplicata,
from which it clearly differs by the incisions of the
lamina extendingdown to the petiole.
35. Cecropia mutisiana Mildbraed,in Cuatrecasas,
Trab. Mus. Ci. Nat. (Madrid), Bot. Ser. 26: 29.
1933. Type. Colombia. Cundinamarca:La Esper-
anza, 1500 m, 18 Apr 1932 (?), Cuatrecasas3082
(holotype: MA-n.v., photographin COL).
Cecropia ibaguensis Cuatrecasas,Revista Acad. Col-
omb. Ci. Exact. 6(22/23): 289. 1945. Type. Colom-
bia. Tolima:Between Ibague & Cajamarca,1800 m,
7 Jul 1939 (Y), PerezArbeldez& Cuatrecasas5737
(holotype: COL; isotypes: F, US).
CecropiatolimensisCuatrecasas,RevistaAcad. Colomb.
Ci. Exact. 6(22/23): 282. 1945. Type. Colombia.To-
lima: Between Fresno & Mariquita,1150 m, 7 May
1940 (d), Cuatrecasas 9397 (holotype: COL; iso-
type: F).
SYSTEMATICTREATMENT 125

1998~~~~~~~~

FIG.35. Cecropia multisecta. 1. Lamina,reduced.2. Apex of lamina and venation.3. Stipules and bases of petioles
with trichilia. 4. Pair of staminateinflorescences at anthesis and base of petiole with trichilium. 5. Staminateflower and
stamen (Franco et al. 4575). 6. Pair of pistillate inflorescenceat anthesis and base of petiole with trichilium(Franco et al.
4574). 7. Pistillate flower. 8. Fruit (Franco et al. 5569). (From Caldasia 23: 78. 2001.)
126
Tree, to 12(-20) m tall. Leafy twigs 2.5-4 cm
thick, hispidulousto puberulouswith straightto un-
cinatehairs.Laminachartaceousto subcoriaceous,ca.
35 X 35 cm to 75 X 75 cm, the segments (9-)10-
12(-13), the free parts of the upper segments oblan-
ceolate to (sub)obovateto elliptic, the incisions down
to ca. 7/10-8/10; apices obtuse to rounded;uppersur-
face scabrous to scabridulous, hispidulous or (ini-
tially) also with arachnoidindumentum;lower surface
puberulousto hirtellousto subtomentoseon the veins,
with arachnoidindumentumin the areoles or extend-
ing to the main veins; lateralveins in the free partof
the midsegment 10-21, submarginally(to margin-
ally), loop-connected(the lowerones often faintlyso),
unbranchedor branched;petiole ca. 20-60 cm long,
puberulousto subhispidulousor (initially) also with
arachnoidindumentum;trichiliafused, the upperrim
often irregularlylobate, the brown indumentumin-
termixed with short white hairs; stipules 6-23 cm
long, darkred to red-brown,hirtellousto puberulous
or also with arachnoidindumentumoutside, (partly)
white sericeous inside. Staminate inflorescences in
pairs,the peduncleerect and the spikes pendulous(or
spreading); peduncle 5-10 cm long, hispidulous;
spathe 8-18 cm long, (red-)brown,sparselyto rather
densely hirtellousto puberulousoutside, densely yel-
lowish subsericeousinside; spikes ca. 15-25, 4.5-14
X 0.2-0.3 cm, with stipes 0.4-1 cm long and sparsely
hairy; rachis hairy. Staminateflowers: perianth tu-
bular,ca. 1 mm long, sparsely and minutely puberu-
lous below the apex, the apex plane; filaments flat;
anthersca. 0.4 mm long, appendiculate,detachedat
anthesis,reattachedto the marginsof the apertureby
the appendages(?). Pistillate inflorescencesin pairs
or solitary,erect, soon deflexed to pendulous;pedun-
cle 3-7 cm long, hispidulousor also with arachnoid
indumentum;spathe 10-13 cm long, the color and
indumentumas in the staminateinflorescence;spikes
(2-)4-5, 6-12 X 0.4-0.6 cm, to 31 X 1.3 cm in fruit,
sessile; rachis hairy. Pistillate flowers: perianth ca.
1.5-2 mm long, with arachnoidindumentumbelow
the apex outside, also in the style channel inside, the
apex convex, minutely puberulous to muriculate;
style short; stigmas comose. Fruit ellipsoid, ca. 1.5
mm long, smooth, darkbrown.
Distribution (Fig. 8.3). Colombia, upperMagda-
The species is probably related to Cecropia obtu-
lena valley, in forest and secondarygrowth,normally
at 800-1800 m, but also found as low as 500 m, or in sifolia.
lected at 510 m, is that of C. peltata, but the flowers
Caldas even down to 200 m.
match those of C. mutisiana, although the stigma is
Representative specimens examined. COLOMBIA. relatively large due to long papillae.
CALDAS:Nr. La Dorada, 1-20 Feb 1946 (9 fl), Duque-
Jaramillo 2590 (COL, NY). CAUCA:Rd. Ricaurte-Guadu-
alejo, 1165-1270 m, 19 Jul 1993 (i), Agudelo et al. 3100 blanco (Caldas).
FLORA NEOTROPICA
(COL);rd. Guadualejo-Inza,1270-1810 m, 19 Jul 1993 (9
fl-fr), Agudelo et al. 3114 (COL). CUNDINAMARCA:S of
Silvania, 7 km on rd. to Tibacuy from km 37 of old rd. to
Fusagasuga, 1500 m, 15 Jun 1972 (6), Barclay et al. 3506
(COL, US); Mun. San Bemardo, Vrda. Portones, 1650 m,
28 Jul 1981 (d fl-fr),Diaz P 3263 (COL,MO);Mun.Pueblo
Nuevo, rd. to San Antonio, 15 Mar 1986 (9 fl-fr), Franco
et al. 2409 (COL);Zipac6n, El Ocaso, rd. to CartagenaLa
Mesa, 14 Mar 1988 (d), Franco et al. 2441 (BG, COL);
Mun. La Magola, 1600 m, 11 Apr 1988 (6), Franco et al.
2483 (COL, MO), (9 fl-fr), Franco et al. 2485 (COL,MO);
Mun. Guaduas,Alto La Mona, 1250 m, 18 May 1990 (9 fl-
fr), Franco et al. 2973 (COL), (6), Franco et al. 2974
(COL); Silvania, at km 62, 1520 m, 12 Mar 1993 (6),
Franco et al. 4344 (COL);Mun. Melgar, Rio Sumapaz,nr.
La Nariz del Diablo, 12 Mar 1993 (9 fl), Franco et al. 4345
(COL, MO); Mun. Tena, rd. Bogota-La Mesa, 1650 m, 6
Feb 1995 (6), Franco et al. 4644 (BG, COL; HUA, MO,
US); Mun. Guaduas,10 km from Guaduas,12 Feb 1985 (9
fl), Galeano et al. 512 (COL), (6), Galeano et al. 513
(COL); Mun. San Francisco,Vrda. El Pifion, 25 Feb 1985
(9 fl-fr), Galeano et al. 552 (COL); rd. Caquera-Puente
Quetame,km 68, PuenteQuebradaBlanca, 13 Mar 1985 (9
f-fr), Galeano et al. 645a (COL);La Esperanza,RR to Gi-
rardot, 1300 m, 23 Jul 1934 (9 fr), Garcia-Barriga4692
(COL, F, UDBC, US); rd. Fusagasuga-Silvania,1680 m, 21
Mar 1970 (9 fl-fr), Idrobo 6319 (COL), (6), Idrobo 6320
(COL); Mun. Bituima, 1400 m, 6 Mar 1972 (6 + 9 fr),
Pinz6n-Rodrfguezs.n. (COL, US); nr. Anolaima, ca. 1500
m, 25 Jul 1963 (9 fl-fr),Soejartoet al. 112 (GH, K). HUILA:
Mun. Garz6n,Vrda. Las Quebraditas,21-26 Feb 1947 (9),
Bermtidez34923 (F); between Gigante & Rioloro, 19 Mar
1940 (9), Cuatrecasas 8334 (COL, F); Guadalupe,below
Gabinete, 1700 m, 16 Oct 1993 (6), Franco et al. 4505
(BG), ( 9 fr), Franco et al. 4506 (BG); Caucho,E of Palacio,
ca. 20 km NE of Santa Ana, ca. 1800 m, 24 Feb 1944 (6),
Little 7343 (COL, P, US); 15 km NE of Algeciras, ca. 1550
m, 1 Apr 1944 (9 fr), Little 7562 (COL, F, US); 3 km SW
of Acevedo, ca. 1450 m, 15 Aug 1944 (9), Little 8478
(COL, F, US); Mun. Colombia, Cerro Bellavista, 1340 m,
15 Apr 1989 (9 fl-fr), Llanos et al. 1372 (COL); between
Gigante & Rioloro, 19 Mar 1940 (9 fl-fr), Perez Arbeldez
& Cuatrecasas8334 (COL);beyondGuadalupe,1000-1300
m, 20 Mar 1940 (9), Perez Arbeldez & Cuatrecasas8402
(F); Pitalito,Calamo, 1300 m, 30 Dec 1942 (d), Schulteset
al. 5128 (GH, NY). QUINDIO:Mun. Quimbaya,Vrda. Pa-
lermo, 23 Nov 1989 (d), Velezet al. 724 (COL). TOLIMA:
Mun. Mariquita,rd. Mariquita-Honda,17 May 1990 (9 fl),
Francoet al. 2970 (COL,MO), 2973 (COL);withoutprecise
locality, 1000 m, 6 Jan 1882 (6), Lehmann2341 (G).
The general aspect of Franco at al. 4345, col-
Vernacular names. Colombia: calentano, yarumo
SYSTEMATICTREATMENT 127

36. Cecropia obtusa Trecul,Ann. Sci. Nat. Bot., ser. minutelypuberulousor also with arachnoidindumen-
3, 8: 79. 1847; Berg, Acta Amazonica 8(2): 169. tum; rachis hairy. Pistillate flowers: perianth ca. 2
1978. Ambaiba obtusa (Trecul) Kuntze, Revis. mm long, with arachnoidindumentumbelow the apex
Gen. P1.2: 624. 1891. Type. FrenchGuiana.With- outside, also below the style channelinside, the apex
out locality, 1834 (d), Leprieur 195 (lectotype, convex, densely muriculate; style rather long,
Berg, Fl. Suriname5(1): 288. 1975: P). straight;stigma comose. Fruitoblongoid to ellipsoid,
CecropialisboanaSnethlage, Notizbl.Bot.Gart.Berlin- ca. 2.5 mm long, ? tuberculate.
Dahlem9: 171. 1924.Brazil.Maranhao: Tury-assd, Distribution (Fig. 8.4). Guianas and the lower
Snethlage283 (holotype:B, destroyed, photographs Amazon basin, in forest and secondarygrowth,at low
in MICH,MO).
elevations.
Tree,to 12(-25) m tall. Leafy twigs 1-5 cm thick,
Representative specimens examined. GUYANA. Ru-
green, hispidulouswith curvedhairs. Laminacharta- pununiDistr.,between KuyuwiniLanding& KassikaityuR.,
ceous to (sub)coriaceous,ca. 25 X 25 cm to 60 X 60 18 Oct 1992 (Y fl-fr), Jansen-Jacobset al. 2997 (BG, U);
cm, the segments 6-8(-9), the free partsof the upper MazaruniR., Kamarang,21 Aug 1977 (Y fl-fr),Maas et al.
segments usually obovate to subobovate,sometimes 2594 (NY, U); S of Temehri, 13 Sep 1979 (Y fr), Maas et
to subspathulate,the incisions down to 5/10-7/10(-8/ al. 3525 (K, NY, U); RupununiDistr., foothills NW of Ka-
10); apices obtuse to subacute(short-acuminate);up- nuku Mtns., nr. Moco-Moco, 27 Oct 1979 (c), Maas et al.
per surface ? scabrousor smooth, (very) sparselyto 3848 (MO, NY, U); base of Mt. Makarapan,MakarapanCr.,
ratherdensely hispidulous and with ? dense arach- 17 Sep 1988 (Y fl-fr), Maas et al. 7501 (K, NY, U); Marudi
noid indumentum;lower surface puberulousto pu- Mtns.,MazoaHill, 13 Nov 1982 (d), Stofferset al. 339 (BG,
K, NY, S, U).
bescent on the veins, with arachnoidindumentumin
SURINAME. Watramiri(tree no. 1534) 2 Aug 1916 (Y
the areoles, extendingto the main veins; lateralveins
fl), BW 2241 (U), (tree no. 1533), 29 Jul 1917 (Y fl), BW
in the free part of the midsegment 9-15 pairs, sub- 2967 (U),(tree no. 1633) 18 Mar 1918 (Y fr), BW3818 (U);
marginally loop-connected, unbranched or faintly Sectie 0 (tree no. 527), 25 Jul 1917 (d), BW 3031 (AAU,
branched;petiole 15-50 cm long, puberulousto pu- U), 9 Aug 1918 (i), BW 3953 (U, US); Brownsberg(tree
bescent and with ? dense (persistent) arachnoid no. 1163), 3 Apr 1916 (st), BW 1711 (U); Cottica R., nr.
indumentum;trichilia fused, the brown indumentum Moengo, 15 Aug 1933 (9 fr), Lanjouw453 (U); Lely Mtns.,
intermixedwith short and/orratherlong white hairs; 25 Sep 1975 (9 fl), Lindeman& Stofferset al. 367 (NY);
stipules 7-12(-20) cm long, dark to bright red or Sipaliwini savanna,nr.Brazilianborder,Feb 1970 (i), Old-
grayish when covered by dense arachnoidindumen- enburger et al. 1297 (U); Jodensavanne-MapaneCr. area,
tum,puberulousto hirtellousto strigillose andusually Feb 1961 (st), Schulz (LBB) 8628 (U).
also with sparseto dense arachnoidindumentumout- FRENCH GUIANA. Charvein,20 Nov 1913 (6), Be-
side, (sub)sericeous inside. Staminateinflorescences noist 279 (P); Saul, Monts La Fumee, 1982 (9 fl-fr), Boom
et al. 2086 (NY), 19 Oct 1982 (d), Boom et al. 2162 (NY);
in pairs, the peduncleerect and the spikes pendulous;
nr. Cayenne, 27 Jun 1921 (9 fl), Broadway582 (GH, NY,
peduncle 4-8.5 cm long, hispidulousor partlypubes-
US); LowerOyapockbasin, SavaneRoche du QuatorzeJuil-
cent; spathe8-17 cm long, red-brownto yellowish or let, 19 Apr 1991 (st), Cremers12330 (CAY); Region Paul
whitish, puberulousto hirtellous and with sparse to Isnard,Camp Voltaire,28 Oct 1991 (9 fl), Cremerset al.
dense arachnoid indumentumoutside, glabrous in- 12490 (BG); Regina region, Mt. Tortue, 11 km WNW of
side; spikes 15-26, 4-13 X 0.2-0.3 cm, with stipes ApprouagueR., 18 Jun 1988 (6), Feuillet 10313 (BG); Cay-
0.5-1.5 cm long and densely minutely puberulous; enne, Montabo,Mar 1978 (9 fr), Grenand1542 (U); Piste
rachis hairy. Staminateflowers: perianth 1-1.5 mm de St. Elie, 7 Jan 1986 (9 fl-fr), Lauri 187 (BG); nr. Saul,
long, minutely puberulouswith straighthairs or also 11 Aug 1987 (9 fl), Mori et al. 18706 (NY), 15 Sep 1989
with arachnoidindumentumbelow the margin, the (6), Mori et al. 20907 (BG, NY); Criquede la Folie, 23 Jul
apex plane to slightly convex or truncate,muriculate, 1965 (9 fl-fr), Oldeman 1486 (P); piste de St. Elie, 19 Jul
the apertureoften surroundedby a rim; filaments 1979 (J), Prevost 706 (P, U); Cayenne, CentreORSTOM,
17 Aug 1979 (9 fl), Prevost 732 (P,U); OyapockR., Zidock,
slightly swollen; anthers0.5-0.7 mm long, appendi-
11 Sep 1980 (9 fl), Prevost et al. 1009 (P, U); Acarouany,
culate, detachedat anthesis,reattachedto the margins
1857 (d), Sagot 515 (BM, G, P, S, U, US); Saint Laurent
of the apertureby the appendages(?). Pistillate info- de Maroni, 12
Aug 1948 (d), Service des Eaux et Forets
rescences in pairs, erect; peduncle 6.5-8(-14) cm 4070 (P).
long, hispidulousor partlypubescent;spathe8-12 cm BRAZIL. AMAPA:Mun. Mazagao,Rio Jari,Morrosdo
long, the color and indumentumas in the staminate Felipe, 10 Oct 1987 (9 fr), H. T Beck et al. 32 (BG, INPA,
inflorescence;spikes 4, 3-10 X ca. 8 cm, to 16 X 1.5 NY); Rio Oiapoque,Boa Esperanqa,opposite mouth of Ca-
cm in fruit, sessile or with stipes to 0.8 cm long and mopi R., 18 Aug 1960 (d), Egler 47658 (COL,GH, K, MG,
128
NY, S, US); Rio Oiapoque,between Oiapoque& Santo An-
tonio, 26 Jul 1960 (9 fl-fr), Irwin et al. 47159 (BG, K, MG,
NY, US); Rio Araguari,ca. 1?40'N,51?56'W,28 Aug 1961
(9 fl), Pires et al. 50533 (MG, MO, NY, U, US); P6rto
Platon, 13 Oct 1976 (d), Ribeiro 1502 (INPA, MG, MO,
NY, RB, U); Rio Oiapoque, CachoeiraCamaraua,3 km S
of mouthof CamopiR., 29 Sep 1960 (9 fl-fr), Westra48523
(BG, MG, NY). MARANHAO:Mun. Carutapera,Gurupiuna,
3 Nov 1986 (st), Balee et al. 2783 (BG, NY). PARA: Mun.
Altamira,IgarapeIpixuna,4 Nov 1985 (st), Balee et al. 1830
(BG); Serrados Carajas,SerraNorte,ca. 20 km N of AMZA
ExplorationCamp, 16 Oct 1977 (9 fl-fr), Berg et al. 585
(A, AAU, COL, MG, NY, RB, U); Belem, groundsof IAN,
7 Nov 1962 (9 fl-fr), Cuatrecasas26654 (P, US); Rio Jar-
amacarii,1 Jun 1957 (9 fl), Egler 437 (MG); Belem, Santa
Izabel, Oct 1906 (9 fl-fr), Goeldi (MG) 7727 (BM, G, MG,
S, U); Tucuruf,20 Oct 1983 (9 fl-fr), E. Lima et al. 64
(INPA, NY); rd. Bel6m-Braslfia, km 92, 24 Sep 1959 (9
fr), M. Kuhlmannet al. 284 (MG, SP, US); Mun. Altamei-
rim, Monte Dourado, 10 Oct 1986 (d), Pires et al. 1425
(BG, MG); rd. BR.22, nr. Cachoeira,27 Oct 1965 (9 fl-fr),
Prance et al. 1700 (GH, K, MICH,MO, NY, S); rd. BR.163,
Cuiaba-Santar6m,km 1223, nr. Igarape Kazuo, 18 Nov
1977 (d), Prance et al. 25548 (K, MG, NY, U, US), (9 fl-
fr), Prance et al. 25549 (K, MG, MO, NY, U, US); rd. Cu-
iaba-Santarem(BR.163), km 1300, nr. IgarapeJose Preto,
23 Nov 1977 (st), Prance et al. 25708 (MG, NY, U); Rio
Jarn,Monte Dourado, 11 Oct 1968 (9 fl-fr),N. T Silva 1178
(NY, U, US), 30 Oct 1968 (9 fl-fr), N. T Silva 1340 (NY,
U): Mun. Almeirim, 18 Sep 1984 (d), N. T Silva 5393
(INPA, MG); rd. Altamira-Maraba,km 19, 30 Aug 1974
(st), N. SmithB.38 (INPA);Serrade Carajas,AMZA camp,
8 Jun 1982 (9 fl-fr), Sperling et al. 5966 (MG, NY).
The species appears to be closely related to the
allopatric Cecropia ficifolia. The two species have
many featuresin commmon. Cecropia obtusa differs
from C. ficifolia, e.g., in the largernumberof spikes
in the staminateinflorescence,the shorterpeduncleof
the pistillate inflorescence,and the shorterstipules. If
the lamina is deeply incised (5/10), then the free part
of the midsegmentis ? abruptlynarrowedand thus
+ distinctly spathulate,whereas usually obovate to
subobovateand rarely subspathulatein C. obtusa. In
C. obtusa, the petiole is coveredwith persistentarach-
noid indumentum.When such indumentumoccurs in
C. ficifolia (as in Ecuadorand Colombia),it is (often)
in combinationwith hirsuteindumentum.WhereasC.
ficifolia is a weedy species, at least in partof its range
of distribution, C. obtusa is associated with forest
habitats.
The following collection is difficult to interpret:
Serra do Navio, Rio Amapari, 12 Nov 1954 (9 fr),
Cowan 38280 (NY, US). The label indicates thatit is
made from a tree 5 m tall, unlikely for this species to
be already in fruit; the inflorescences are separate
from the partialleaf, which has narrowsegmentswith
FLORANEOTROPICA
up to ca. 25 pairs of lateral veins in the free part of
the midsegment, and the arachnoidindumentumis
dense in the petiole but sparseon the laminaand ab-
sent in the areoles beneath.It is difficultto relatethe
leaves to the juvenile state of C. obtusa.
The name Cecropiaobtusa was based on four col-
lections: the present lectotype, Bonpland 1295 from
Cuba(= C. schreberianasubsp.antillarum),Gardner
3981 (= C. saxatilis), and Ruiz & Pavon s.n. (= C.
polystachya).
Vernacular names. Guyana:congo pump, trum-
pet tree. FrenchGuiana:ama'i (Wayapi);bois canon,
kulegle (Wayana). Brazil: ama-'y-tuwir (Ka'apor,
Maranhao);imbaubabranca(Maranhao);ama'i-hete
(Tupi, Para);imbauibabranca(Para).
37. Cecropia obtusifolia Bertoloni,Fl. Guatimal.39.
1840; Woodson & Schery, Ann. Missouri Bot.
Gard.47: 175, t. 61. 1960; Burger,Fieldiana,Bot.
40: 125, t. 23. 1977. Type. Guatemala.Without
locality, J. Veldsquez s.n. (BOLO-n.v., photo-
graphsin BM, K, MICH, MO). Fig. 36
Cecropiacommutata Schottex Miquel,in MartiusFl.
Bras.4(1): 148. 1853.Type.Materialcultivatedin
HortusSchonbrunn, frommaterialcollectedin Mex-
ico by Schottin 1852(holotype:M, photographsin
MICH,MO).
CecropiamexicanaHemsley,Biol.Centr.Amer.Bot.3:
151. 1883. Ambaiba mexicana (Hemsley) Kuntze,
Revis.Gen.P1.2: 623. 1891.Syntypes:
Mexico.Ve-
racruz:"Valleede Cordova,"4 Feb 1866(9 & 5),
Bourgeau 1869 (C [5], BM, GH [5], K, P, US [9]).
CecropiapanamensisHemsley,Biol.Centr.Amer.Bot.
3: 151. 1883.Type.Panama.Nr.PanamaCity,Mar
1847(5), Seemann499 (holotype:K).
Ambaiba costaricensis Kuntze, Revis. Gen. P1. 2: 623.
1891.Type.CostaRica.Not designated,
nortraced.
AmbaibahemsleyanaKuntze,Revis.Gen.P1.2: 623.
nortraced.
1891.Typenotdesignated,
CecropiaradlkoferianaV. Richter,Biblioth.Bot. 43: 17.
1897.Type.Colombia.Choc6:RioSanJuan,Triana
865 (P,mixedcollection,thepistillateinflorescence
designated as thelectotype,Berg& FrancoRosselli,
Fl. Ecuador48: 41. 1993;isolectotype:K).
CecropialevyanaBureau&V.Richter, Biblioth.Bot.43:
20, t. 5, 1897.Type.Nicaragua. Jun1870
Chontales,
(5), Levy473 (holotype: P,fragmentin F; isotypes:
G, P).
CecropiamexicanaHemsley var.macrostachyaDonnell
Smith,Bot. Gaz. (Crawfordsville)27: 442. 1899.
Syntypes:CostaRica.Lagunade BuenosAires,Feb
1892 (9 fl-fr),Pittier6666 (US);Cartago:Atirro,
Mar 1894 (9), Donnell Smith 4934 (GH, K, US),
Rio Poas, 1200 m, Mar 1896 (9), DonnellSmith
SYSTEMATIC TREATMENT 129

FIG. 3s 1.
(< :1~~~~~~~~~~~~~~~~

,7

4 ~ ~ ~ ~ 1

.4~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~. .4

4~~~~~~~~~~~~~~~~~~~~~.

FIG. 36. Cecropia obtusifolia. 1. Lamina, reduced. 2. Apex of lamina and venation. 3. Leafy twig with stipules,
young leaf, and base of petiole with trichilium (Berg 1242). 4. Staminate inflorescence and base of petiole with trichilium
(Berg 1540). 5. Staminate flower and stamen (Berg 1541). 6. Pistillate inflorescence with spathe and base of petiole with
trichilium. 7. Pistillate inflorescence at anthesis and base of petiole with trichilium (Berg 1242). 8. Pistillate flower. 9. Fruit
(Berg et al. 432). (From Fl. Ecuador 48: 43. 1993, modified.)
130
6771 (K, US); La Concepci6n, Llanurasde Santa
Clara, Feb 1896 (Y), Donnell Smith 6774 (GH, K,
US).
Cecropia maxonii Pittier, Contr. U.S. Natl. Herb. 18:
228. 1917. Type. Panama.Chiriquf:Nr. El Boquete,
1000-1300 m, 2-8 Mar 1911 (d), Maxon 5132 (ho-
lotype: US).
CecropiaamphichloraStandley& L. 0. Williams,Ceiba
3: 111. 1952. Type. Honduras.Cortes: La Cumbre,
Sierrade Omoa, 30 Nov 1950, Molina R. 3455 (US,
mixed collection, the leaf here designated as lecto-
type; the d inflorescenceis C. peltata).
Cecropia dabeibana Cuatrecasas,Revista Acad. Col-
omb. Ci. Exact. 6(21): 67. 1944. Type. Colombia.
Antioquia:Between Dabeiba & Llanitos,Rio Sucio,
25 Mar-I Feb 1942 (Y), Metcalf & Cuatrecasas
30168 (holotype: COL; isotypes: F, US).
Cecropia burriadaCuatrecasas,Revista Acad. Colomb.
Ci. Exact. 6(22/23): 274. 1945. Cecropiaobtusifolia
Bertoloni subsp. burriada (Cuatrecasas)C. C. Berg
& P. Franco,Fl. Ecuador48: 41. 1993. Type.Colom-
bia. Valle:Rio Calima,La Trojita,26 Feb 1944 ( ),
Cuatrecasas16505 (holotype:COL;isotypes:F, US,
VALLE).
Cecropia alvarezii Cuatrecasas,Revista Acad. Colomb.
Ci. Exact. 6(22/23): 276. 1945. Type. Colombia.
Valle: Rio Cajambre,Barco, 21 Apr 1944 (v), Cua-
trecasas 16960 (holotype: COL; isotypes: US,
VALLE).
Tree, to 12(-35) m tall. Leafy twigs 1.5-5 cm
thick, green or partly purplish, hirtellous to subhis-
pidulous with straightand/oruncinatehairs. Lamina
chartaceousto subcoriaceous,ca. (15 X 15 cm to) 30
X 30 cm to 90 X 90 cm, the segments (8-)10-13 or
7-1 0, the free partsof the uppersegmentssubobovate
to oblanceolate (to elliptic or obovate), the incisions
down to ca. 5/10-8/10; apices obtuse to roundedor
acuminate;upper surface ? scabrous, hispidulous;
lower surface puberulousto hirtellous to subtomen-
tose on the veins, with arachnoidindumentumin the
areoles, also on the reticulumor (almost)lacking;lat-
eral veins in the free part of the midsegment (10-)
15-25(-50), marginally to submarginally loop-
connected, often branched;petiole (lO-)20-80 cm
long, puberulousto hirtellous and/or with arachnoid
indumentum, or subglabrous; trichilia fused, the
brownindumentumintermixedwith shortwhite hairs;
stipules 5-15 cm long, (maroon-)redor yellowish,
(sparsely) puberulousto hirtellous to hirsute to sub-
sericeous or also with arachnoidindumentumoutside,
sericeous to subvillous or to subglabrousinside. Sta-
minate inflorescences in pairs (or solitary), the pe-
duncle erect to pendulous,the spikes erect to spread-
ing to pendulous;peduncle 4-18 cm long, (minutely)
puberulousto subglabrous;spathe5-20 cm long, red
FLORANEOTROPICA
to red-brownto pink or to purple or yellowish with
reddish stripes and spots (or whitish), (sparsely)hir-
tellous to subhirsute,often also with sparse(or dense)
arachnoid indumentum, or subglabrous (and then
sometimes with a bluish waxy layer) outside, gla-
brous or sericeous inside; spikes 8-18(-24), 4-25 X
0.2-0.4(-0.8) cm, sessile or with stipes to 2.5(-10)
cm long and puberulous;rachis sparselyhairy or gla-
brous.Staminateflowers: perianthtubular,1-1.5 mm
long, usually with short hairs below the apex, some-
times glabrous,the apex plane, the apertureoften sur-
roundedby a rim; filamentsflat; anthers0.5-0.7 mm
long, appendiculate,detachedat anthesis, reattached
to the marginsof the apertureby the appendages.Pis-
tillate inflorescencesin pairs (or solitary), erect or at
least the spikes pendulous;peduncle 3-24 cm long,
(minutely) puberulousto subglabrous;spathe 12-22
cm long, the color and indumentumas in the stami-
nate inflorescence;spikes 2-4(-6), (4-)8-25 X 0.4-
0.7 cm, to 35(-55) X 1.5 cm in fruit, sessile or with
stipes to 1.5(4) cm long and puberulous; rachis
hairy. Pistillate flowers: perianth 1.5-2 mm long,
with arachnoidindumentumbelow the apex, also in
the lower part of the style channel inside, the apex
convex, muriculate to minutely puberulous or
smooth; style rather short; stigma penicillate. Fruit
ellipsoid, 2-2.5 mm long, smooth, darkbrown.
Distribution (Fig. 8.5). From Mexico to the Pa-
cific coastal region, southwardto Ecuador,in forest
and secondary growth, at elevations to 1300(-1650)
m.
Representative specimens examined. MEXICO.
CHIAPAS: 5 mi N of Tapiula,10 Jul 1966(d),
Baird1029
(US);Mun.VillaCorzo,10-12 km SW of ColoniaAgr6n-
omos Mexicanos, CerroTres Picos, 1000 m, 8 Feb 1972 (Y
fr), Breedlove 23943 (MO); Mun. Huixtla, 6-8 km NE of
Huixtla, 27 Dec 1972 (6), Breedlove et al. 30939 (MO);
Mun. Ococingo, Centro Arqueol6gico Bonampak, 19 Feb
1982 (6), Castillo 265 (MO); Escuintla,Esperanza,13 Feb
1945 (6), Matuda 16411 (EAP, MICH, NY); Mun. Angel
Albino Corzo, Rio Cuztepeques,26 Mar 1968 (? fl-fr),Shi-
lom Ton 3894 (LL, MICH); Mun. Tuxtla Chico, Curva de
Cahoacan,rd. to Talsiman,3 Jun 1992 (6), Vazquez176
(MEXU); Mun. Tapachula,El Porvenir,21 Nov 1984 (6),
et al. 729 (MEXU);Mun.Cacahoatan,
Ventura Cuatimoc,
14 Nov 1985 (6), Venturaet al. 2705 (MO). GUERRERO:
Nr. Puerto Marquez, 10 Aug 1966 (6), Baird 1041 (US);
PuertoMarquez,20 May 1979 (st), L6pez F 788 (MEXU);
Mun. Atoyac, 27 km NE of Atoyac de Alvarez, rd. to Par-
aiso, 28 Mar 1983 (Y fl-fr), Soto N. et al. 5064 (MEXU,
MO). HIDALGO: 4 km SE of San Bartolo, Tutitepec,3 Jun
1972 (Y fl-fr), Gimate L. 642 (AAU, COL, EAP, MICH,
PMA, US); Dtto. Jacala,Cantil de Espejo, nr. km 340-341
on hwy. below Chapulhuacan,30 Jan 1948 (Y fl-fr),Moore
SYSTEMATICTREATMENT
et al. 3642 (A, MICH, TEX). JALISCO: 7 km W of Villa de
Purificaci6n, 10 Dec 1988 (st), Bullock 2022 (MEXU); 0.1
km SW of rd. Barra de Navidad-PuertoVallarta,on rd. to
Manzanilla,24 Dec 1983 (Y fl), Lott et al. 2213 (MEXU);
rd. to Mazatlan,ca. 19 mi NW of Tepic, 15 Nov 1959 (d),
McVaughet al. 717 (MICH); 15-18 rd.-mi SW of Autlan,9
Apr 1951 (6), MceVaugh 11957 (MICH);SantaCruzde Val-
larta, 10 Dec 1926 (c), Mexia 1275 (A, BM, G, MICH,MO,
NY, US). MEXICO: 3 km S of San Miguel Amatepec,20 Jan
1973 (c3'),GonzalezM. et al. 5180 (MEXU);Temascaltepec,
Anonas, 3 Jul 1933 (Y fl-fr), Hinton 3568 (G, GH, K, NY).
NAYARIT: Ca. 12 km E of San Blas, 27 Apr 1962 (d + Y
fl-fr), Barr et al. 62-305 (MEXU);rd. PuertoVallarta-Tepic,
km 103, 22 Apr 1980 (i), Rooden 751 (U), (9 fl-fr),
Rooden 752 (U); 9.3 km W of Jalcocatan,rd. to Miramar,4
Feb 1989 (G), Tellez V et al. 11733 (MO). OAXACA: Rd.
PuertoEscondido-Oaxaca,8 mi N of San Gabriel,ca. 1000
m, 9 May 1965 (9 fl-fr), Breedlove 9912 (MICH); Dtto.
Tuxtepec, Presa Temazcal, 8 Dec 1985 (G), Cortes et al.
111 (MEXU); Dtto. Tuxtepec, rd. Esmeralda-Rio Manea,
11.5-13.5 mi S of Esmeralda,19 Jan 1987 (6), Croat63278
(BG, MO); Mun. Soyaltepec, 4 km S of HidroelectricaTe-
mascal, 20 May 1986 (9 fl-fr), Gereau et al. 2223 (BG,
MEXU, MO); Mun. Temascal, Cortina de la Presa M.
Aleman, 21 Mar 1964 (6), Gonzalez Q. 575 (MEXU); nr.
Rio Soyaloapan, 21-22 Sep 1971 (9 fl-fr), Graham 1395
(MICH);Mun. Juxtlahuaca,1 km NE of Tilapa, 1200 m, 25
Jan 1985 (9 fl), Lopez G. et al. 262 (MEXU); 5 km N of
Pochutla,2 Mar 1985 (9 fl-fr),L6pezG. 351 (MEXU);Dtto.
Choapam,Yaveo, trail of El Chorro,27 Mar 1938 (9 fl-fr),
Mexia 9225 (G, GH, K, MO, NY, U, US); San Felipe Usila,
9 Mar 1929 (6), Sabino s.n. (NY); Chichihua,17 km N of
Lazaro Cardenas, 14 Mar 1983 (6), Tenorio et al. 3450
(MEXU). PUEBLA: Mun. Oaonahua,rd. Teteles-Mazatepec,
Atotocoyan, 24 Mar 1976 (6), Mdrquez717 (NY); Tuza-
mapa de Galeana, 1 Apr 1980 (6), Espadas et al. 96
(MEXU); nr. La Ceiba, 19 Apr 1959 (9 fl-fr), Go'mez-
Pompa167 (MEXU);rd.Mexico-Tuxpan,km 247.3,30 Apr
1962 (st), Sarukdnet al. 2090 (MEXU). SAN Luis POTOSi:
3 mi N of Tamazunchale,13 Aug 1966 (d), Baird 1042
(US); 8 mi NE of Xilitla, 1 Apr 1961 (9 fl), King 4426
(MICH, NY, TEX, US); Mun. Xilitla, El Joba,6 May 1959
(9 fr), Rzedowski10595 (US); Ubero, May 1937 (9 fl-fr),
LI. Williams9431 (P), Jun 1937 (9 fl-fr), LI. Williams9466
(K, P). SINALOA: Mazatlan, Jun 1923 (6), Ortega 5086
(US); El Habal, 1923 (9 fl-fr), Ortega 5194 (US); nr. Col-
omas, 20 Jul 1897 (st), Rose 1776 (US). TABASCO:Without
locality, 1840 (6), Lindens.n. (G, P). VERACRUZ:51 mi S
of Tantoycua,13 Jun 1966 (9 fl-fr), Baird 1004 (US); San
AndresTuxtla,Sihuapan,9 Jan 1973 (6), Calzada927 (BM,
K, MO); between Conejos & Huatusco, 18 Nov 1956 (9 fl-
fr), Cox 561 (MEXU); Estaci6n de Biologia TropicalLos
Tuxtlas, 22 May 1981 (9 fl), Gentry et al. 32257 (LL,
MEXU, MO); rd. Martinezde la Torre-Misantla,km 3, 13
Jun 1963 (9 fl-fr), G6mez-Pompa913 (A, MEXU); 2 km
NE of Putla, 14 Mai 1981 (6), Lorenceet al. 3436 (MEXU);
Mun. Catemaco, 7 km NE of Sontecomapan,1 Nov 1981
(9 fl), Nee 22573 (LL);Dtto. Tuxtepec,EjidoBenito Juarez,
30 Aug 1967 (9 fl), Pennington et al. 9073 (NY); Mun.
131
Catemaco,rd. Catemaco-Montepfo,km 3,15 Mar 1978 (J),
Perino 3275 (NY); Cerrode Cuauhtlapan,between Cuauht-
lapan & Tuxpango, 30 Apr 1967 (i), Rosas R. 317 (A, K,
P, U); 3.1 km W of Santiago Tuxtla, 10 May 1965 (9 fr),
M. Sousa 2361 (MEXU); Mun. Hidalgotitlan,Rio Las Cue-
vas, campamentoLa Laguna,6 Mar 1984 (9 fl-fr),K. Taylor
394 (NY, RB, U); Mun. Hidalgotitlan,Cedillo rumboa Paso
Noral, 17 Jun 1974 (9 fl-fr), Valdivia800 (K, MO); Mun.
Minantitlin,2 km N of Uxpanapa,14 Mar 1982 (c), Wendt
et al. 3684 (MEXU); Fortunia,CoatzacoalcosR., Mar 1937
(9 fl), LI. Williams8665 (MICH,P).
GUATEMALA.ALTA VERAPAZ: Cubilqiiitz,Feb 1901
(9 fl-fr), Tuerckheimin herb. Donnell Smith 7666 (GH, K,
NY, US), Feb 1901, Tuerckheimin herb.Donnell Smith7890
(US). ESCUINTLA: Ca. 10 km NE of Escuintla, 1100 m, 2
Aug 1966 (i), Baird 1000 (US); Escuintla, Mar 1890 (9
fl-fr), Donnell Smith2024 (G, GH, K, NY, P, US); 5 mi N
of Escuintla,31 Jan 1970 (i), Harmon1828 (MO). IZABAL:
Nr. Izabal, 23 Apr 1966 (6), G. C. Jones et al. 3034 (NY);
S of Rio Dulce, 8 Aug 1975 (6), LeDoux et al. 2122 (NY).
PETEN: SantaTeresa,SubfnR., 14 Apr 1933 (9 fl-fr),Lun-
dell 2891 (MICH, NY, TEX); San Luis, 11 Jul 1959 (9 fl-
fr), Lundell 16355 (LL, NY); Laguna Petexbatun, 1 Apr
1964 (9 fl-fr), Lundell 18191 (LL). RETALHULEU: Retal-
huleu, Finca Concepci6nPital, 15 Jul 1966 (d), Baird 1031
(US); San Andres, Apr 1892 (st), Donnell Smith 1499 (K,
US); Telahuleu, 12 Jan 1907 (9 fl), Kellerman6039 (US).
SANTA RoSA: Malpais, ca. 1300 m, Sep 1893 (6), Heyde
& Lux in Donnell Smith6238 (G, GH, K, NY, US). SUCH-
ITEPEQUEZ: Between Escuintla & Mazatenango, 11 Jan
1978 (9 fl), Harriman14663 (MO).
BELIZE. StannCr. Distr., Cocksombbasin, JaguarRe-
serve, 21 May 1990 (9 fl), Balick et al. 2630 (BG, NY);
ToledoDistr.,SouthernMayaMtns.,BladenNatureReserva,
8 May 1996 (d), Davidse et al. 35776 (BG, MO); El Cayo
Distr., S of Belmopan, 14-21 Jun 1973 (9 fl-fr), Croat
24563 (GH, MO), 14-21 Jun 1973 (6), Croat24887 (MO);
Toledo Distr., nr. San Antonio, 1 Apr 1969 (9 fl), Lazor et
al. 1710a, (9 fr), Lazor et al. 1710d (MO, NY); El Cayo
Distr., Valentin, Jun-Jul 1936 (9 fl-fr), Lundell 6373
(MICH), (9 fl-fr), Lundell6374 (MICH);Middlesex, 2 Oct
1929 (9 fl-fr), Schipp341 (A, BM, G, K, MICH,MO, NY,
S), 1928 (6), Stevenson 14 (US); Maya Mtns., Humming-
bird Rd., 13 Aug 1992 (9 fl-fr), R. D. Worthington21363
(L).
EL SALVADOR. AHUACHAPAN: El Impossible Re-
serve, Campana, 1400 m, 24 Jan 1998 (6), Monro et al.
2058 (BG); San Benito, 5 Feb 1990 (6), Sermeno80 (MO),
23 Dec 1992 (6), Sandovalet al. 942 (MO). LA LIBERTAD:
Cafetal de Miguel EdaudoAraujo, 1000 m, 7 Jul 1988 (9),
Monro2289 (BG). SAN SALVADOR: In or nr. San Salvador,
9 Nov 1985 (6), Berendson 311 (MO), Feb 1922 (9 fl),
Calderon 104 (GH, NY, US), 1922 (6), Calderon 1264
(GH, NY, US); San Miguel Tepexontes,1923 (6), Calderon
1840 (GH, US); San Salvador,20 Dec 1921-4 Jan1922 (6),
Standley19614 (GH, NY, US), (9 fl), Standley22665 (GH,
NY), (9 fr), Standley23181 (US).
HONDURAS. ATLANTIDA: Pico Bonito, Quebrada
Grande,ca. 10 km SW of La Ceiba, 8 May 1993 (9 fl-fr),
132 FLORANEOTROPICA

Evans 1551 (EAP,MO);nr.Tela, 14 Dec 1927-15 Mar 1928, 1966 (6), Molina R. et al. 18220 (EAP,NY, US); FincaLos
(6), Standley 56637 (A); La Ceiba, 6 Aug 1938 (6), Cusingos, 8 mi SE of San Isidro del General,22 Dec 1948
Yunckeret al. 8844 (GH, NY). COMAYAGUA:El Achote, (6 fl), Norby 133 (EAP); nr. El General, Jan 1939 (9 fl),
above plains of Siguatepegue, 1350 m, 26 Jul 1936 (st), Skutch3986 (A, K, MO, NY, S, US); nr. Santo Domingo,
Yunckeret al. 6127 (NY). CORTES: 1 km SW of Omoa, 13 1100 m, 23 Dec 1974 (6 fl-fr), J. Taylor17540 (NY).
Dec 1970 (9 fl-fr), Harnon et al. 5198 (MO). OLANCHO: PANAMA. BOCASDEL TORO:1 mi W of Almirante,
Between Las Marias& La Colonia, 4 Feb 1982 (d), Black- 15 Oct 1965 (6), Blum 1383 (MO); Rio Changuinola,0.5
more et al. 1655 (BG, BM, MEXU, MO). TEGUCIGALPA: mi upstreamfrom mouth, 19 Apr 1969 (st), Lazoretal. 2653
Mont. de la Flor, 1937 (st), Hagen et al. 1264 (NY). (MO); 10-15 mi S of mouth of Rio Changuinola, 18 Dec
NICARAGUA. JINOTEGA: Rd. Jinotega-Wiwilf,NE of 1966 (6), Lewis et al. 868 (MO); Isla Bastimentos, 1-2 km
Jinotega,3 Mar 1980 (9 fl), Araquistainet al. 1718 (MO). N of TroubleHole, 14 Feb 1989 (6), Peterson et al. 6858
RiO SANJUAN:Rio Santa Cruz, 30 Sep 1982 (9 fr), Ara- (MO); nr. ChiriquiLagoon, WaterValley, 14 Sep 1940 (6),
quistain 3244 (MO); San Juandel Norte, Jan 1896 (9 fl-fr Wedel771 (GH, MO, US), 9 Nov 1940 (d + 9 fr), Wedel
+ 6), C. L. Smith 57 (EAP, GH, MO, NY, US). ZELAYA: 1559 (GH, MO). CHIRIQUi: Cerro Horqueta,ca. 2300 m,
Nueva Guinea, 11 Aug 1982 (9 fr), Araquistain3033 (BG, 24 Jul 1966 (6), Blum et al. 2607 (MO); Burica Peninsula,
MO); Slilma Sia, 6 Mar 1971 (9 fl-fr), Little 25073 (MO, above Brazo Seco, nr. Costa Rican border,3 Mar 1973 (6),
NY, US); S of Rio Wawa,60 km NW of PuertoCabezas, 11 Croat22562 (MO); Boquete, ca. 1300 m, 1 Jul 1938 (9 fr),
Mar 1971 (6), Little 25077 (MO, US); Mun. Siuna, Santa Davidson 862 (A). COL6N:Nr. Rio Piedras, rd. to Puerto
Rosa, 6 Sep 1982 (9 fl-fr), Ortiz 96 (BG, MO); rd. to Co- Bello, 16 Jul 1966 (6), Blum et al. 2496 (MO), (9 fl-fr),
loniaYolaina,Colonia La Esperanza,11-12 Feb 1978 ( 9 fl- Blum et al. 2531 (MO); 3 mi NW of Gamboa, 10 Jul 1966
fr), Stevens6359 (BG, MO); rd. to Colonia Manantiales,0.8 (9 fl-fr), Haines 467 (LL, MO); Gatun, Dec 1857 (6),
km S of ridge of Serraniasde Yolaina, 13-14 Feb 1978 (6), Hayes 671 (NY); Las CrucesTrail,8 Feb 1935 (6), Hunter
Stevens 6454 (BG, MO); rd. Rio Blanco-Rio Copalar,ca. et al. 462 (GH, MO); nr. Fort Sherman, 5 Jul 1967 (6),
29 km E of Rio Branco, 14 Feb 1979 (9 fl-fr),Stevens12193 Stimson5251 (COL, NY, PMA, US, SCZ); 7 mi N of Gam-
(BG, MEXU, MO). boa, 11 Mar 1966 (9 fl-fr), Tyson3526 (MO, PMA); Salud,
COSTA RICA. ALAJUELA: 6 km E of CiudadQuesada, 27 Jul 1971 (9 fl-fr), Lao et al. 169 (MO); 4 km NW of
22 May 1968 (9 fl-fr), Burger et al. 5201 (GH, S, US). Salamanca,13 km NE of Buenos Aires, 30 Dec 1973 (d),
CARTAGO: Turrialba,8 Mar 1966 (d), Blum 2216 (MO), Nee 9084 (MO, PMA). DARIEN: Nr. Pinogana,4 Mar 1947
(9 fl-fr),Blum2217 (MO); Atirro,Mar 1894 (9 fl-fr),Don- (9 fl-fr), Allen 4311 (EAP, MO); ParqueNacional Darien,
nell Smith4934 (GH, US); VolcdnArenal, 17 Sep 1972 (9 Crucede Mono, 5 Nov 1989 (st), Fisher 54 (BG); 16-19 km
fl-fr), Lent 2948 (U); 9 km SE of San Jose, 1500 m, 19 Dec SE of Jaque,20 Jan 1981 (9 fl), Garwood 1130 (BG, MO,
1976 (6), Lent 4016 (AAU, MO), (9), Lent 4017 (NY). PMA); 23 km SE of Jaque,24 Jan 1981 (6), Garwood1144
GUANACASTE: NE of Sabalito, rd. to La Uni6n, 1-18 Jan (BG, MO, PMA), (9 fl-fr), Garwood 1147 (MO); nr. Cana,
1967 (9 fl), Burger et al. 4549 (BM, EAP, G, MO); Pan- 23 Jun 1959 (d), Sternet al. 490 (GH, MO, US); E of Santa
American Hwy., Rio Congo, 6 Jan 1964 (6), Molina R. Fe, 18 Jul 1966 (6), Tysonet al. 4852 (MO, SCZ), (9 fl-
13571 (EAP, NY). HEREDIA:Finca La Selva, Rio Puerto fr), Tysonet al. 4853 (MO, SCZ);Rio Cocalito, 10 Feb 1982
Viejo, 16 Feb 1981 (9 fl), Folsom 8979 (MO); Finca La (9 fl-fr), Whitefoordet al. 125 (BM, MO, PMA, U), (6),
Selva, PuertoViejo de Sarapiqui,23 Jul 1979 (9 fl), Sperry Whitefoordet al. 133 (BM, MO, PMA). PANAMA: Cerro
1011 (MO); nr. Rio Sarapiquf,25 Jul 1990 (6), Wiemann Jefe, 30 Aug 1977 (9 fl-fr), Berg et al. 391 (BG); beyond
311 (NY); Rio Virilla, 10 km S of Heredia, 1000 m, 25 Mar Goofy Lake, rd. to CerroJefe, 4 Jan 1968 (6), Correaet al.
1949 (9 fr), L. 0. Williams16034 (EAP). LIMON:Pta. Ca- 575 (MO, PMA); rd. to Cerro Campana, 14 Mar 1968 (9
huita, 1 Feb 1980 (9 fl-fr), Berg 949 (U); between Siquirres fl-fr), Correaet al. 850 (MO, PMA); BarroColoradoIs., 15
& Rio Pacuare,20-22 Dec 1919 (9 fl-fr),Burgeret al. 6934 Apr 1968 (6), Croat4796 (MO, NY, US), 5 Aug 1970 (9
(GB, LL, U); 29 km W of Tortiguero,31 Aug 1979 (9 fl- fl-fr), Croat 11716 (MO, SCZ), 30 Jan 1970 (9 fl), Foster
fr), Davidson et al. 8990A (MO); ParqueTortuguero,Esta- 1484 (PMA); Cerro Azul, Feb 1968 (9 fr), G6mez-Pompa
ci6n Cuatro Esquinas, 3 Dec 1987 (9 fl-fr), Robles 1446 et al. 3413 (MEXU, MO, PMA); 5-10 km NE of Altos de
(BG, MO); Cerro Coronel, E of Laguna Danto, 15-20 Sep Pacora, 8 Mar 1975 (9 fl-fr), Mori et al. 4989 (BG, MO);
1986 (9 fl-fr), Stevens 24592 (MO, NY). PUNTARENAS: Cerro Campana,above Su Lin Motel, 25 Mar 1969 (6),
Golfo Dulce area,nr.Jalacafarm,8 Jun 1949 (9 fl-fr),Allen Porter 4194 (MO); rd. to CerroAzul, 5 Mar 1972 (9 fl-fr),
5302 (EAP,US); above PalmarNorte, trailto Buenos Aires, Tyson6170 (PMA); above El Valle, rd. to La Mesa, 11 Feb
17 Feb 1951 (6 fl-fr), Allen 5937 (EAP); EsquinasStation, 1974 (6), Tyson 6932 (PMA). SAN BLAS: Nusagandi, 16
29 Aug 1952 (9 fl-fr), Allen 6545 (EAP); ca. 17 km N of Aug 1989 (9 fl), Fisher 21 (BG), 26 Jul 1986 (6), Mc-
San Vito, 30 Mar 1978 (9 fl-fr), Davidson 7165 (MO, NY); Donagh et al. 405 (BM, MICH, MO); El Llano-Carti rd.,
Monteverde,Lower Tigre Reserve, 1150-1250 m, 29 Jan continental divide, 25 Aug 1984 (9 fl-fr), Nevers 3754
1991 (9 fl), Haber et al. 10486 (BG); CaniasGordas, 1100 (PMA);El Llano-Cartird., km 19, Nusagandi,26 Aug 1984
m, Nov 1899 (9 fl-fr), Pittier 11077 (G, US); 4 mi W of (9 fl), Nevers et al. 3769 (BG, MO, PMA). VERAGUAS:Nr.
Rinc6n de Osa, 8 Aug 1967 (9 fr), Raven21644 (NY). SAN SantaFe, Rfo SantaMarfa,26 Mar 1947 (9 fl-fr),Allen 4422
JOsE: El Generalvalley, nr. San Isidro El General, 28 Feb (MO).
1966 (9 fr), Molina R. et al. 18102 (EAP, NY, US), 2 Mar COLOMBIA. ANTIOQUIA:Mun. Turbo,rd. Tap6ndel
SYSTEMATICTREATMENT
Darien, between Rfo Le6n & Lomas Aisladas, km 37, 25
Aug 1984 (d), Brand 1083 (JAUM);Mun. Anori, Vrda. El
Carmen, 18-20.5 km NW of Anorn,19 Nov 1989 (2 fl-fr),
Callejas et al. 8793 (BG, HUA); between Villa Arteaga &
Chigorod6, Lomitas, 1 Oct 1961 (2 fl-fr), Cuatrecasaset
al. 26114 (COL, P, US); Mun. Mutata,Rio Mutata,2 May
1987 (2 fl-fr), Fonnegraet al. 2070 (BG, HUA); Mun. San
Francisco, Cgto. Aquitania, Rfo Venado, 1150 m, 4 Apr
1992 (2 fr), Fonnegra et al. 4221 (BG, MO); Mun. San
Rafael, CampamentoVersalles, 1350 m, 23 Mar 1994 (2
fr), Franco et al. 4573 (BG, COL, HUA); Valdivia, 1250 m,
27 Mar 1994 (2 fl-fr), Franco et al. 4596 (BG HUA), (d),
Franco et al. 4597 (BG, HUA); Taraza,Cgto. El Doce, Finca
Las Mercedes,28 Mar 1994 (2 fr), Franco et al. 4599 (BG,
HUA); Mun. Frontino, Cgto. Nutibara, rd. Nutibara-La
Blanquita,Murriregion, 4 Mar 1995 (2 fl-fr), Franco et al.
5558 (BG, COL, HUA, LP, MO, US); Mun. Anorf,Vrda.El
Roble, 1100 m, 8 Mar 1995 (2 fl-fr), Franco et al. 5567
(BG, COL, HUA, LP, MO, US), 1600 m, 8 Mar 1995 (d),
Franco et al. 5571 (BG, COL, HUA, LP, MO, US); Mun.
Urrao, ParqueNacional Natural Las Orquideas,Quebrada
La Agudelo, 1300-1380 m, 4 Apr 1992 (d), Muntozet al.
534 (JAUM);Mun. San Luis, QuebradaLa Cristalina, 21
May 1987 (d), Ramirezet al. 909 (JAUM),29 Jul 1987 (d),
Ramfrezet al. 1212 (JAUM);rd. Uraba-Chigorod6-Mala-
g6n, Cafio Malag6n, 22 Mar 1986 (2 fl-fr), Renterfaet al.
4713 (JAUM,MO); Mun. Frontino,upperRfo Cuevas,Cgto.
Nutibara,1630 m, 15 Jan 1987 (2 fl), D. SdnchezS. et al.
990 (BG); Mun. San Carlos, Cgto. El Jordan,Trocha Las
Planes, 10 Apr 1991 (2 fr), Veldsquezet al. 299 (HUA).
CAUCA: Rio Micay, El Chachajo,27 Feb 1943 (2 fl), Cua-
trecasas 14266 (F); Mun. El Tambo,Pepital, 1700 m, 12 Jan
1970 (2 fl), Idrobo 6316 (COL); Mun. Guapi, ParqueNat-
uralGorgona,rd. Playa Blanca-Gorgonilla,6 Sep 1987 (6),
Lozano C. et al. 5652 (COL). CHOCO: Bete, Apr 1977 (2
fl-fr), Acero et al. 89 (UDBC); Rio Atrato, nr. Quibd6, 23
Jan 1949 (2 fl), AraqueM. et al. 159 (COL, MEDEL,US);
ca. 20 km SE of Quibd6, 8 Jul 1986 (d), Berg 1540 (BG);
rd. Quibd6-Bolivar, ca. km 30, 9 Jul 1986 (2 fl-fr), Berg
1549 (BG); Mun. Acandi, Vrda. Coquital, El Paramo, 26
May 1989 (2 fl-fr), Betancuret al. 1256 (BG, HUA); Mun.
Riosucio, Cerros del Cuchillo, rd. Cuchillo Negro-Cumbre
Noroeste, 20 Jan 1988 (2 fl-fr), Cdrdenas 1115 (JAUM,
MO); Rio San Juan,Quebradadel Tarapal,30 May 1946 (2
fr), Cuatrecasas 21510 (F); Cataugad6,nr. Quibd6, rd. to
Munquirri,30 Mar 1958 (2 fl-fr), Cuatrecasaset al. 24050
(COL, US); Rio Atrato,Yuto, 2 Apr 1958 (d), Cuatrecasas
et al. 24164 (COL, US); Rio Truando,between La Nueva &
La Esperanza,6 Feb 1967 (2 fl-fr), Duke 9883 (US); Rio
Salada, Camp Teresita, 14 Jun 1967 (6), Duke 12204 (NY,
US); Rio Munguid6,nr.Altagracia,5 May 1975 (2 fr), For-
ero et al. 1554 (COL, MO); Rio Tamana,tributaryof Rio
San Juan,below Santa Rosa, 10 Apr 1979 (6 fl-fr), Forero
et al. 4969 (BG, COL); Mun. Bahia Solano, nr. Bahia So-
lano, 4 Apr 1990 (2 fl), Franco et al. 3011 (COL); Mun.
Yuto, rd. Yuto-Quibd6, 12 Apr 1990 (2 fl), Franco 3055
(COL); Quibd6, 14 Apr 1990 (2 fl), Franco et al. 3062
(COL); Rfo Baudo, opposite Quebrada Caimanerita, 11
Feb-29 Mar 1967 (2 fl-fr), Fuchs et al. 21921 (COL, G, K,
133
NY, P, S, US); Rfo Baud6, between Puerto Pizarro & La
Sierpe, 28 Feb 1967 (9 fl-fr), Fuchs et al. 22112 (COL, K,
NY, US); rd. Yuto-Certegui,17 Aug 1976 (9 fl-fr), Gentry
et al. 11791 (BG, COL); Rfo Yuto, between Llor6 & La
Vuelta, 18 Jan 1979 (6), Gentry et al. 24357 (BG); Mun.
San Jose del Palmar,nr. San Jose del Palmar, 11-15 Nov
1985 (9 fl-fr),Lozano C. et al. 4900 (COL);Mun.Riosucio,
Cgto. Truand6,confluence of Rfo Chintad6& Rio Salad6,
24 Oct 1956 (9 fl), Romero-Castanteda6140 (COL,
MEDEL);Nuqui, Alto de Buey, 1000 m, 26 Jun 1940 (9 fl-
fr), SneidernA.71 (A, COL, K, LL, MEDEL, MICH, NY,
S, US), (9 fl-fr), SneidernA.73 (A, COL, K, LL, MICH,
NY, S), 28 Jun 1940 (i), SneidernA.74 (A, MICH, NY, S,
US); Rio Jurad6,4 Oct 1940 (d), SneidernA.245 (MICH);
ParqueNacionalNaturalLos Katios,Cristales, 19 May 1982
(i), Zuluaga R. 62 (COL). CORDOBA: Tierralta,Rio Sinu,
26 Jul 1988 (9 fr), CuadrosV 4173 (MO). NARINO:Olaya
Herrera,Vrda.Naidizales,mouthof Rio Satinga,5 Mar 1991
(9 fl-fr), Carmona 132 (COL, MEDEL);Mun. Barbacoas,
Cgto. Junin,El Diviso, 12 Dec 1993 (9 fl-fr), Franco et al.
5178 (BG); Mun. Barbacoas,Llorente, 12 Dec 1993 (9 fl-
fr), Franco et al. 5181 (BG). RISARALDA: Mun. Pueblo
Rico, Rio Taiba, 12 Mar 1986 (5), Bernal et al. 952 (COL);
Mun. Mistrat6,rd. Mistrat6-Puertode Oro, Cgto. Jeguades,
26 May 1991 (9 fr), Franco et al. 3442 (COL, MO); Mun.
Mistrat6,Cgto. Rio Mistrat6,11 Sep 1991 (9 fl-fr), Franco
et al. 3508 (COL,MO); Mun.Mistrat6,Cgto. Puertode Oro,
28 Sep 1991 (6), Franco et al. 3605 (COL); Mun. Pueblo
Rico, Vrda. Piunda, 26 Apr 1991 (d), Gonzdlez 2419
(COL).SANTANDER:Mun. PuertoWilches,betweenLa G6-
mez & km 80 of Atlantico RR, 23 Apr 1960 (5), Romero-
Castaneda8409 (COL). VALLE: Mun. Buenaventura,Cgto.
Bocas del Tiger, QuebradaMondoya, 27 Oct 1979 (5),
CuadrosV 937 (BG, COL, MO); Rio Anchicaya,Quebrada
del Retiro, 19 Dec 1942 (5), Cuatrecasas13689 (COL, F);
Rio Naya, PuertoMerizalde, 20-23 Feb 1943 (d), Cuatre-
casas 13969 (F, US, VALLE),(9 fl-fr), Cuatrecasas13979
(F); Rio Yurumangui,El Papayo, 5 Feb 1944 (9), Cuatre-
casas 15988 (F, US), (5), Cuatrecasas16000 (F, US); Rio
Cajambre,Quebradade Ord6fiez, 1 May 1944 (9 fl-fr),
Cuatrecasas17268 (F,VALLE);Quebradade Guapecito,16
May 1944 (5), Cuatrecasas 17681 (COL, F); Rio Calima,
Quebradade L6pez, 23 Sep 1961 (9 fl-fr), Cuatrecasaset
al. 26042 (COL, P, US); Mun. Restrepo, campamentoRio
Azul, Al Chanco, 18 Mar 1986 (9 fr), Devia 1267 (BG,
MO); Mun. Trujillo,rd. Trujillo-Naranjal,km 43, Rio San-
quinini, 1050 m, 11 Oct 1993 (5), Franco et al. 4489 (BG);
Mun. Buenaventura,rd. to Bajo Calima, Bahia Malaga, 16
Mar 1994 (5), Franco et al. 4537 (BG), (9 fl-fr), Franco
et al. 4538 (BG), (9 fl-fr), Franco et al. 4539 (BG), (5),
Franco et al. 4541 (BG), (5), Franco et al. 4544 (BG), (9
fl-fr), Franco et al. 4546 (BG); Bajo Calima, ca. 15 km N
of Buenaventura,15 Feb 1983 (9 fr), Gentry et al. 40290
(BG, COL, JAUM, MO); Bajo Calima, Cart6n Colombia
forest, 1 Jun 1982 (9 fl), Murphyet al. 531 (NY, TULV,
US); Mun. Buenaventura,San Isidro, 15 Nov-6 Dec 1979
(5 fl-fr), Rooden et al. 257 (BG, MO, NY, U), (5 fl),
Rooden et al. 516 (BG, COL, U).
ECUADOR. AZUAY: Azuay/Guayas/Cafiar-border, E of
134
Jesus Maria, 12 Apr 1980 (5), Gentry et al. 28521 (MO).
CARCHI: Nr. TobarDonoso, 19-28 Jun 1992 (9 fl-fr), Tipaz
et al. 1460, 1479 (BG, QCNE). EL ORO: Nr. Piedras, 11
Jun 1943 (9 fl-fr), Little 6610 (F, Q, US). ESMERALDAS:
Reserva Etnica Awa, Alto Tambo, Rio Mira, 22 Mar 1993
(9 f), Aulestiaet al. 1385 (BG, QCNE);Rfo Cayapa,Zapalo
Grande, 19-25 Oct 1983 (9 fl), Barfod et al. 48433
(QCNE);Reserva IndigenaAwa, Mataje,6 Oct 1993 (9 fl-
fr), H. Beck et al. 2237 (QCNE);rd. Quininde-Esmeraldas,
km 5, 13 Sep 1977 (5), Berg et al. 430 (BG, COL, MO,
QCA, U); rd.VueltaLarga-CoronelC. Concha, 14 Sep 1977
(9 fl-fr), Berg et al. 432 (BG, COL, QCA, U); nr. Lita, 9
Feb 1981 (5), Berg 1254 (AAU, BG, COL, MO, QCA, U);
rd. Quito-San Lorenzo, km 319, Ventanas,16 Jul 1964 (9
fl-fr), Jativa et al. 802 (NY, S, US); Rio Onzole, Estero
Chontaduro,14 Jul 1966 (9 fl-fr), Jativa et al. 1099 (MO,
NY, S, US); Rfo Santiago,nr. Sucre, 10 Aug 1967 (9 fl-fr),
Jdtiva et al. 2217 (S); 2 km S of San Lorenzo, 23 Apr 1943
(9 fl-fr), Little 6361 (F, Q, US); Rio San Miguel, 40 km S
of Borb6n, Rio Cayapas, 13 Sep 1965 (9 fl), Little et al.
21052 (Q, QAME, QCNE); Anchayacu,Eloy Alfaro, May-
ronga, 28 Oct 1993 (9 fl-fr), Penningtonet al. 14530 (BG);
ca. 20 km S of Esmeraldas,10 Apr 1967 (d), Sparre 15514
(S); ReservaEcol6gica Cotacachi-Cayapas,Luis VargasTor-
res, Rfo Santiago, 28-31 Oct 1993 (9 fl-fr), Tiradoet al.
695 (BG, QCNE). GUAYAS:Rio Congo, between Empalme
& Pichincha, 27 Feb 1955 (9 fl-fr), Asplund 15548 (S).
IMBABURA: Nr. Guadalupe,RR Ibarra-SanLorenzo, 7 Feb
1981 (9 fl-fr), Berg 1242 (BG, COL, MO, QCA, U). Los
Rios: Rio PalenqueBiological Station,rd. Quevedo-Santo
Domingo de los Colorados,km 56, 19 Jun 1974 (d), Dodson
et al. 5601 (QCA, US), 2 Oct 1976 (9 fl-fr), Dodson et al.
6237 (MO, US). MANABi:Parque Nacional Machalilla,
Cerroel Pechiche, ca. 15 km NW of Machalilla,22-24 Apr
1992 (st), Josse et al. 785 (BG). PICHINCHA: Rd. Quito-
PuertoQuito, km 113, 10 km N of the rd., ReservaForestal
ENDESA, 8 Jul 1984 (9 fl), Jaramillo 6714 (AAU, GB,
MO, NY, QCA); trail Santo Domingo de los Colorados-
Quininde, km 43-68, 7 Apr 1943 (5), Little 6185 (F, Q,
US); Cant6nPedroVicenteMaldonado,ReservaRio Pitzara,
Nov 1997 (9 fl), Penningtonet al. 16239 (BG).
Cecropia obtusifolia is very variable in many
characters. The trees tend to have tall trunks with rel-
atively small, ? globose crowns. There is a tendency
to monocauly as branches may not be formed. Ac-
cording to Alvarez-Buylla & Martinez-Ramos (1992),
trees may grow to 20-35 m tall and become elements
of the forest canopy in Veracruz (Mexico). Indications
that this could happen in other parts of the species
range are lacking, and on the basis of observations, it
is very unlikely that it would happen in the southern
part of the range. The lamina varies considerably in
shape due to differences in the extension of the inci-
sions. One can recognize two types of leaves with
regard to the number of leaf segments: the most com-
mon type usually with (8-)10-13 lamina segments
FLORA NEOTROPICA
and in the free part of the midsegment >20 pairs of
lateral veins. The other is a less common one, most
typicallyfoundin the Pacificcoastalregionof Colom-
bia and Ecuador(as in Cuatrecasas13689, Franco et
al. 3011 and 5178, Pennington et al. 14530, and
Rooden et al. 516), with 7-10 segments and in the
free part of the midsegment ca. 15 pairs of lateral
veins. The latter type might have links with C. sub-
integra.Arachnoidindumentumis always presenton
the lower surfacethe laminaof materialfromMexico.
Specimens withoutor with very scarce arachnoidin-
dumentumhave been collected in Guatemala.The fre-
quency of materialwithoutor with very scarcearach-
noid indumentumon the leaves increases through
CentralAmerica to the Pacific coastal region of Co-
lombia. Specimens with arachnoidindumentumon
the leaves appearto be absentin Ecuador.The differ-
ences in the presence of arachnoidindumentumhas
led to the distinctionof two subspecies, subsp. obtu-
sifolia, comprising the material with arachnoidin-
dumentum, and subsp. burriada, comprising that
without arachnoidindumentum.We decided not to
maintainthis distinction,as the geographicalsepara-
tion of the two proved not to be clear enough. The
two can be indicatedas the "obtusifolia-type"andthe
"burriada-type"respectively. The number of lateral
veins varies considerably.Leaves with >25 pairs of
lateralveins in the free partof the midsegmentshow
a ratherregularvenationpattern,and the lateralveins
are usually unbranched.Materialwith numerouslat-
eral veins in the midsegmentappearsto be most com-
mon in northernChoc6 andPanamaandis foundmost
frequently in the "burriada-type."Throughout the
range of distribution,one can find a type with short
spikes, often sessile or with short stipes and another
type with long spikes, andoften also long stipes. This
differencein the lengthof the spikes and stipes occurs
most pronouncedlyin the "burriada-type." Material
with erect inflorescences with short (and relatively
thick) spikes can be indicatedas the "alvarezii-type."
These conspicuously different inflorescencescan be
found in the same population.Cecropiaobtusifoliais
very closely relatedto C. subintegraand(less closely)
also to C. heterochromaand to C. mutisiana.It shows
similaritiesto the Amazonianspecies C.putumayonis
and C. utcubambanain the shape of the trees and the
frequentlack of arachnoidindumentumon the lower
surface of the lamina, but features of the staminate
flowers indicatethat they are not closely related.The
trees are usually inhabited with large ant colonies.
The fruitingspikes are reportedto be favoritefood of
toucans. The species has been introducedin Hawaii,
where it has become naturalized(Wetterer,1997).
SYSTEMATICTREATMENT
Vernacular names and uses. Mexico: trompeta
(Jalisco, Sinaloa, Guerrero);chancarro, guarumbo,
huagadeug(Zapotecans,Oaxaca);changarro,huaga-
deug (Oaxaca); chancarro(Veracruz);hormigo, hor-
miguillo (Puebla).Belize: guarumo,trumpet.Colom-
bia: burriada (Cauca); burriala (Embera, Choco).
Ecuador:arumococedera (Esmeraldas);tsenguillachi
(Chaplaachi,Esmeraldas).
The plant is used medicinally,mostly to treatdi-
abetes. Bark fibers are used to make hammocks.
38. Cecropia pachystachya Trecul, Ann. Sci. Nat.
Bot., ser. 3, 8: 80. 1847; Berg & Carauta,Albertoa
1(1): 10. 1986. Ambaiba pachystachya (Trecul)
Kuntze, Revis. Gen. P1. 2: 624. 1891. Syntypes.
Brazil. Minas Gerais: Without locality, Claussen
158 (P), andCeara:Crato,Nov 1838 (9), Gardner
1845, here designated as lectotype (P; isolecto-
types: G, K, P).
CecropiaadenopusMartiusex Miquel,in Martius,Fl.
Bras. 4(1): 147, t. 50, 1. 1853. Ambaiba adenopus
(Miquel) Kuntze,Revis. Gen. P1.2: 624. 1891. Syn-
types. Brazil. Minas Gerais:Rio Francisco,nr. Sal-
gado, Martius 2834 (lectotype, here designated:M)
and Brazil. Amazonas.Rio Solimoes, nr.Manaquiri,
Martiuss.n. (not traced).The two collections belong
to differentspecies without doubt.
Cecropia carbonaria Miquel, in Martius,Fl. Bras. 4(1):
144, t. 48, 1. 1853. Ambaiba carbonaria (Miquel)
Kuntze, Revis. Gen. P1. 2: 624. 1891. Type. Brazil.
Goias, Meiaponte,Pohl s.n. (W, destroyed;isotype:
BR).
Cecropiacinerea Miquel, in Martius,Fl. Bras.4(1): 142.
1853. Ambaiba cinerea (Miquel) Kuntze, Revis.
Gen. P1. 2: 624. 1891. Type. Brazil. "Southern,"
without locality, Pohl s.n. (W, destroyed), photo-
graphin MO).
Cecropia lyratiloba Miquel, in Martius,Fl. Bras. 4(1):
144. 1853. Ambaibalyratiloba(Miquel) Kuntze,Re-
vis. Gen. P1.2: 624. 1891, as "lyratiflora."Cecropia
adenopus Miquel var. lyratiloba (Miquel) Hassler,
AnnuaireConserv.Jard.Bot. Geneve 21: 130. 1919.
Brazil. Minas Gerais:Rio Sao Marcos,Pohl s.n. (W,
destroyed,fragmentin F).
CecropiacyrtostachyaMiquel,in Martius,Fl. Bras.4(1):
146, t. 49, 1. 1853. Ambaiba cyrtostachya(Miquel)
Kuntze, Revis. Gen. P1. 2: 624. 1891. Type. Brazil.
Goias: Arraias, Mar 1840 (Y), Gardner3982 p.p.
(lectotype: K; p. 138).
CecropiadigitataTenoreex Miquel, in Martius,Fl. Bras.
4(1): 149. 1853. Type. Materialcultivatedin Hortus
Schonbrunn,collected in Brazil (holotype: M-n.v.).
Cecropiaglauca Rojas,Cat. Hist. Nat. Corrientes80.
1897. Type not indicatednor traced.
Cecropia adenopus Miquel var. macrophylla Hassler,
135
AnnuaireConserv.Jard.Bot. Geneve 21: 130. 1919.
Syntypes: Paraguay.Upper Rio Apa region, Nov
1901 (c), Hassler 7924 (BM, G), (Y), Hassler
7924a (G).
Cecropia adenopus Miquel var. vulgaris Hassler, An-
nuaire Conserv. Jard. Bot. Geneve 21: 130. 1919.
Syntypes. Paraguay.Cordillerade Altos, 1885-1895
(d), Hassler 617a (G, K, P), (?), Hassler 617b (G,
K, P); LakeYpacarayregion, May 1913 (d), Hassler
12442 (A, BM, C, E, G, GH, K, MO, NY), (Y),
Hassler 12442a (A, BM, C, E, G, GH, K, MO, NY,
US).
CecropiaadenopusMiquel var. Iata Snethlage,Notizbl.
Bot. Gart. Berlin-Dahlem 8: 359. 1923. Syntypes.
Brazil. Without locality, Sellow 2157/2146 (B, de-
stroyed);Rio de Janeiro:Mun. Rio de Janeiro,Cor-
covado, 1891-1892 (i), Glaziou 18500 (B, de-
stroyed;LE).
CecropiaadenopusMiquel var.oblonga Snethlage,No-
tizbl. Bot. Gart. Berlin-Dahlem8: 359. 1923. Syn-
types. Brazil. Rio de Janeiro.Mun. Rio de Janeiro,
Floresta de Tijuca, 1891-1892 (6), Glaziou 18501
(B, destroyed;C, G, LE, P), (Y), Glaziou 18502 (B,
destroyed;C, G, K, LE, NY, P), Glaziou 18506 (B,
destroyed;NY); Mun. Rio de Janeiro:Gavea, 1891-
1892 (d + Y), Glaziou 18504 (B, destroyed;BM,
C, G, K, LE, P), Glaziou 18507 (B, destroyed).
Cecropia catarinensis Cuatrecasas,Brittonia 11: 171.
1959. Type. Brazil. Santa Catarina:Laguna,22 Dec
1951 (6), Reitz & Klein 177 (holotype:US; isotype:
US).
Cecropia lyratiloba Miquel var. nana Andrade & Car-
auta, Bradea3(22): 164. 1981. Type. Brazil. Rio de
Janeiro:Recreio dos Bandeirantes,Reserva Biol6-
gica de Jacarapagua,7 Sep 1979 (d), Andrade 15
(holotype: GUA; isotypes: F, K, R, RB, U).
Tree, to 12 m tall. Leafy twigs 1-4 cm thick, green
(or whitish), hispidulous to puberulous, sometimes
also with arachnoidindumentum.Lamina subcoria-
ceous to chartaceous,ca. 20 X 20 cm to 60 X 60 cm,
the segments 9-13, the free parts of the upper seg-
ments oblanceolateto subobovate,often ? lobate to
sinuate, the incisions down to (5/10-)7/10-9/10,
sometimes down to 2 cm from the petiole; apices
short-acuminateto subobtuse;uppersurfacescabrous,
hispidulousor partlyhirtellous;lower surfacepuber-
ulous to subhirtellousto subtomentose (with hairs
of different length), with arachnoidindumentumin
the areoles, on the smaller veins or also on the main
veins, sometimes only on the main veins and the
margin;lateral veins in the free part of the midseg-
ment 10-20 pairs, submarginallyto marginallyloop-
connected, branched;petiole 10-55 cm long, puber-
ulous to hirtellous,mostly also with sparse to dense
arachnoidindumentum;trichiliafused, the brownin-
dumentum intermixed with white or sometimes
136 FLORA NEOTROPICA

brownish dense rather short (unicellular)hairs, dis- Mucugezinho,nr. Morrodo Pai Inacio, ca. 1000 m, 20 Dec
tinctly longer than the brown trichomes, sometimes 1984 (6), Stannard et al. (CFCR) 7271 (GUA, SPF).
ratherwhite long hairs; stipules (8-)10-22 cm long, CEARA:Fortaleza,nr.Rio Coco, 22 Jan 1968 (9 fl-fr), Car-
auta 544 (US), (5), Carauta 545 (GUA, U, US); Campos
white to pale green, subsericeous to pilose and with
da Cruz, 28 Jan 1968 (5), Carauta 551 (GUA, U, US);
? dense arachnoidindumentumoutside, glabrousor
between Ipiabina& Sao Benedicto, 21 Jul 1995 (5), A. Fer-
hairy inside. Staminateinflorescencesin pairs,the pe- nandes et al. s.n. (GUA); Serrade Araripe,Oct 1838 (9 fr),
duncle erect or deflexed, the spikes pendulous; pe- Gardner 1845 (BM, G). DISTRITOFEDERAL:Reserva Ecol-
duncle 5-12(-17) cm long, hirtellousto subhirsuteto 6gia do IBGE, 15 May 1978 (9 fl-fr), Heringer et al. 491
subtomentose or minutely puberulous, mostly also (IBGE,NY, US); Rio Sao Bartolomeu,rd. Brasilia-Unaf,11
with arachnoidindumentum;spathe 3-18 cm long, Dec 1979 (d), Heringer et al. 2902 (MG, MO, NY, US);
white to pale green, subsericeous to pilose and with Paranoa,ca. 5 km E of Brasilia, 1000 m, 11 Jul 1966 (9 fr),
? dense arachnoidindumentumoutside, glabrousin- Irwin et al. 18150 (COL, G, GH, MO, NY, RB, S, US).
side; spikes 5-20, (1-)2.5-21 x 0.5-1 cm, with stipes ESPiRITO SANTO:Vila Velha,Rodoviado Sol, 30 Aug 1980
(9), Andrade 114 (NY); Domingos Martins,31 Aug 1980
0.1-0.6 cm long and sparselyto densely hairy or gla-
(9 fl), Andrade 122 (G); Mun. Santa Teresa,Estac,o Biol-
brous hairy;rachishairy.Staminateflowers: perianth
6gica de Santa Lucia, 7 Nov 1985 (5), Boudet Fernandes
tubular,1.5-2 mm long, glabrous,the apex plane;fil- 1620 (GUA, MO, NY, US); Mun. Cariacica,Rio da Vit6ria,
aments flat; anthers0.5-0.7 mm long, appendiculate, 5 Aug 1983 (5), Hatschbach46710 (BG, US); Mun. Lin-
detachedat anthesis,reattachedto the marginsof the hares, Reserva Florestal da Cia. Vale do Rio Doce, 27 Sep
apertureby the appendages.Pistillate inflorescences 1978 (9 fl-fr),Martinelli5011 (RB); between BrejoGrande
mostly in pairs, erect, pendulous in fruit; peduncle & Aracruz,27 Oct 1992 (9 fl-fr), 0. J. Pereira et al. 4014
(2-)4-15 cm long, indumentumsimilarto that of the (GUA). GoIAs: Serra dos Pirineus, ca. 10 km NE of Co-
staminate inflorescence; spathe 3-10 cm long, the rumba de Goias, 15 May 1973 (9 fl-fr), W R. Anderson
color and indumentumas in the staminateinflores- 10370 (MO, NY, U, US); Mun. Goiania,Rio Meia Ponte, 5
Feb 1966 (9 fl-fr), Carauta301 (GUA, K, NY); Mun. Lu-
cence; spikes 3-6(-8), often 4, 4-8 X ca. 0.4-0.5 cm,
ziana, Rio Bartolomeu, 15 Aug 1980 (5), Hatschbach
to 15(-18) X ca. 1 cm in fruit, sessile or with stipes
43138 (AAU, C, NY, U); Rio Sao Marcos,Cristalina,29 Jan
to 0.2 cm long andpuberulous;rachishairy.Pistillate 1980 (9 fl-fr),Heringeret al. 17630 (K, MG, MO, NY, US);
flowers: perianthca. 1.5 mm long, with arachnoidin- Rio da Prata,ca. 6 km S of Posse, 6 Apr 1966 (5), Irwinet
dumentumbelow and on the apex outside, also in the al. 14415 (F, MG, MO, NY, US); 2 km SE of Piranhas,23
style channel inside, the apex almost plane; style Jun 1966 (9 fl), Irwinet al. 17688 (GH, MO, NY, RB, US);
short;stigmapeltate.Fruitoblongoid, 2-2.2 cm long, nr. Sao Gabriel,ca. 50 km N of Planaltina,rd. to Veadeiros,
tuberculate,darkbrown. 1000 m, 21 Jul 1966 (9 fl-fr), Irwin et al. 18307 (F, GH,
MO, NY, RB, US); Rio Javaes,27 Jul 1978 (9 fl-fr), Pires
Distribution (Fig. 18.1). Brazil, fromthe southern et al. 16258 (MG, NY, U); SerraCaiap6, 50 km S of Caia-
fringes of the Amazon basin,throughcentralandeast- ponia, 23 Oct 1964 (9 fl-fr), Prance et al. 59603 (GH, K,
ern parts of the country, extending to Paraguayand NY, S, U, US). MARANHAO: Mun. Lago Verde, rd. Alto
northernArgentina, in forest, cerrado, and restinga Alegre-Lago Verde, km 9, 26 Mar 1985 (st), Anderson et
areas, common in secondarygrowth, at elevations to al. 2184 (MG); Is. of Sao Luiz, Feb-Mar 1939 (? fl-fr),
1200 m. Froes 11634 (A, G, K, MICH, MO, NY, S, U); Nova Es-
peranca,Alto Turiaqu,5 Dec 1978 (5), Jangoux et al. 256
specimensexamined.BRAZIL.AMA-
Representative (MG, U), (9 fl-fr), Jangouxet al. 257 (MG, U); Rio Alper-
ZONAS: Mun.Humaita,ca. 20 km S of Humaita,Rio Ma- catas, ParqueEstudialde Mirador,28 Sep 1988 (9 fr), Nob-
deira, 13 Mar 1975 (6), Gottsbergeret al. 156-13375 (U). erto et al. 215 (MG). MATOGROSSO:6 km S of Xavantina,
BAHIA: Marati,6 May 1966 (Y fl-fr),Belem et al. 2074 (U); 19 Sep 1967 (9 fr), Argentet al. 6426 (NY, P, U); 47 km S
rd. Canavieiras-Una,km 30, 16 Jul 1981 (6), Berg et al. of Xavantina,rd. to Aragarcas,9 Nov 1968 (5), Harley et
1149 (BG, U); Mun. Santa Cruz de Cabralia, 16 km W of al. 10968 (E, NY, P, RB, U); Mun. Rio Verde,Serrada Pi-
P6rtoSeguro, 10 Mar 1983 (d), Brito etal. 228 (MG);Mun. menteira,28 Aug 1973 (5), Hatschbach32454 (BG, C); nr.
Jacobina,Serrado Tombador,ca. 8 km SW of Jacobina,27 Alto Gar9as,22 Jul 1974 (9 fr), Hatschbach34702 (MO,
Oct 1995 (6), Carvalho6139 (NY); Mun. I1heus,rd.Ilheus- U, US); Serra do Roncador,ca. 86 km N of Xavantina,3
Itabuna,km 22, 12 May 1981 (Y fl-fr), Hage et al. 662 Jun 1966 (9 fr), Irwinet al. 16554 (GH, MO, NY, RB, US);
(GUA); rd. to Abaira,ca. 8 km N of Rio de Contas (town), 40 km S of BR.070, rd. to Pocone, Rio Bento G6mes, 20
18 Jan 1972 (6), Harley et al. 15223 (AAU, NY, SPF, U); Jan 1989 (9 fr), Krapovickaset al. 43065 (C, K); Cuiaba,
Mun. Rio de Contas,Pico das Almas, 1500 m, 24 Dec 1988 22 Jun 1902 (5), Malme 11.1811(S); Sarar6,RADBRASIL,
(9 fl-fr), Harley et al. 27357 (SPF); rd. Ilheus-Una, ca. km 22 Aug 1978 (5), Pires et al. 16644 (MG, U); Garapuair-
50, 15 Feb 1992 (9 fl-fr), Hind et al. 40 (BG, K); Mun. strip, 1 Oct 1964 (9 fr), Prance et al. 59208 (K, NY, S, U).
Uruquca,rd. Uru,uca-Serra Grande,km 28-30, 4 Nov 1978 MATO GROSSO DO SUL: Rio Aquidauana, rd. to Rochedo,
(9 fl), Mori et al. 11060 (NY, U); between Len,ois & Rio 24 Jan 1979 (9 fl-fr), Carauta et al. 3077 (F); Mun. Co-
SYSTEMATICTREATMENT
rumba,Abobral, 11 Jun 1994 (6), Hatschbachet al. 60929
(NY); Mun. Ivinhema, Riberao Lib6rio, 1 Apr 1986 (6),
Pastore et al. 84 (RB); Nhecolandiaregion, ca. 100 km SE
of Corumbai,30 Oct 1985 (Y fl-fr), Ratteret al. 5081 (NY).
MINAS GERAIS: Mun. Januaria,nr. Hospital Sao Vincente,
3 Feb 1970 (9 fl-fr), Carauta1062 (GUA, NY, RB, U, US);
Mun. Januaria,nr. Ponte Boa Vista, 14 Feb 1972 (st), Car-
auta 1474 (F, GUA, RB), (d), Carauta 1475 (F, LE, RB,
U); Curvelo, 3 Mar 1972 (9 fl-fr), Carauta 1502 (F, GUA,
RB), (6), Carauta 1503 (F, LE, RB); Mun. Araguari,Rio
Aguarari,BR.050, 10 Jul 1979 (9 fl-fr), Hatschbach42244
(AAU, MO, NY, U); Mun. Marlieria, ParqueEstadual do
Rio Doce, 18 Sep 1975 (9 fl-fr),Heringeret al. 15037 (BG,
MO, US); Paracuta,29 Jan 1980 (9 fl-fr), Heringer et al.
17634 (K, MG, NY); between Grao & Mogol, Ribeiraodas
Mortes, 10 Dec 1989 (6), Pirani et al. (CFCR) 12431
(GUA, SPF); Caldas, 13 Dec 1864 (9 fl-fr), Regnell L415
(C, G, K, LE, P, S, US). PARA:Upper Rio Tapaj6s, Rio
Cururu,Missao Cururu,6 Feb 1974 (9 fl-fr), W R. Ander-
son 10556 (BG, US); Rio Xingu, Ilha Belo-Horizonte, 12
Oct 1986(d fl-fr),Dias et al. 219 (MG);UpperRio Tapaj6s,
Rio Cururui, Lago Boiossu, 24 Jul 1959 (9 fl-fr),Egler 1018
(MG). PARAiBA: Mun. Rio Tinto, 23 Feb 1989 (6), Agra
682 (K). PARANA: Mun. TerraRica, Rio Paranapanema,25
Jul 1992 (6), Barros 2585 (SP); Morungava,21 Jan 1915
(9 fl), Dusen 16456 (F, GH, MO, NY, P, S); Mun. Senges,
Fazenda Morungava,Rio Funi'l,10 Oct 1958 (6), Hatsch-
bach 5103 (US); Mun. Paranagua,Rio Guaraguacu,26 Oct
1960, (d), Hatschbach 7333 (US); Mun. Foz do Iguacu,
Parque Nacional, 20 Feb 1962 (9 fr), Hatschbach 9920
(US); Mun. Antonina, Rio Xaxim, 15 Feb 1979 (9 fl-fr),
Hatschbach 41973 (C, GB, INPA, MG, MICH, U); Mun.
Paranagua,Ponta do Poco, 15 May 1982 (9 fl-fr), Hatsch-
bach 44931 (MO, U, US); ParqueNacional do Iguacu, Rio
Floriano, 1200 m, 3 Dec 1966 (6), Lindemanet al. 3537
(NY, U). PERNAMBUCO:Mun. Caruaru,Brejo dos Cavalos,
21 Jan 1972 (6), Carauta1467 (U), (9 fl-fr), Carauta1468
(U); Rio Formoso, Horto Florestalde Saltinho (d + 9 fl),
Falcdo et al. 1155 (RB); Recife, Mata de Dois Irmaos, 28
Sep 1988 (d), Guedes 1581 (UEC); Itapessuma,9 Oct 1887
(9 fl-fr), Ridley et al. s.n. (BM). PIAUh: Bananeiros,Serra
Negra, Parque Nacional, 16 Sep 1977 (6), Barroso 282
(RB); Floriana,Fazenda Agua B6a, 12 Dec 1973 (d), Ra-
malho 316 (RB); Urucuf,4 Jul 1925 (st), Snethlage647 (F).
RIO DE JANEIRO: Macae, rd. to Lagoa Comprida,7 Apr
1982 (d), Araujo et al. 4918 (NY); Mun. Angra dos Reis,
Ilha Grande,4 Jan 1990 (9 fl-fr), Araujo 9055 (GUA); nr.
Cabo Frio, 16 Nov 1980 (9 fl-fr), Berg et al. 1135 (BG,
NY, RB, U); Mun. Rio de Janeiro,Restinga de Itapeba,23
Oct 1963 (d), Carauta 201 (COL, U, US); Mun. Rio de
Janeiro,ReservaBiol6gica de Jacarepagua,9 Sep 1964 (st),
Carauta225 (K, MG, NY); Itatiaia,nr. Hotel Donati, 7 Jan
1966 (9 fl), Carauta296 (GUA, RB, US); Mun. Nova Ig-
ua,u, Vila de Mesquita,base of Macico de Gericin6, 19 Nov
1967 (9 fl-fr), Carauta et al. 491 (NY, RB, U, US); Mun.
Silva Jardim,nr. Lagoa Jutunaiba,2 Apr 1976 (6), Carauta
1899 (RB, U); Resende, P6rto Real, Rio Paraibado Sul, 22
Oct 1981 (d), Carautaet al. 3893 (GUA, K, NY); Barrado
Piraf,between Dorandia& Sao Jose do Turvo, 23 Feb 1987
137
(5), Carautaet al. 5418 (GUA); Mun. Rio de Janeiro,Res-
tinga de Jacarepagua,3 Nov 1962 (Y fl-fr), Cuatrecasaset
al. 26644 (P, US), (5), Cuatrecasas et al. 26646 (P, US);
BarraMansa,FazendaParaizo,18 Jan 1961 (Y fl-fr),Duarte
5485 (RB, U); Morro do Pan da Fome, 19 Feb 1981 (? fl-
fr), Lira 409 (C, G, LE); Recreio dos Bandeirantes,3 Jan
1962 (d + Y fl-fr),Strang343 (K, MG, NY). Rio GRANDE
DO SUL:Sao Leopoldo, Jan 1942 (5), Leite 2300 (A, GH);
P6rto Alegre, nr. Gl6ria, 16 Dec 1901 (5), Malme II.869
(S); Sao Leopoldo, Campus da Unisinos, 27 Mar 1991 (Y
fl-fr), Quadroset al. 1677 (GUA), (5), Quadroset al. 1678
(GUA); PortoAlegre, Morroda Caloria, 16 Oct 1945 (Y fl),
Rambo29298 (S); nr. Osorio, 25 Jan 1958 (Y fl-fr),Rambo
63578 (S). SANTA CATARINA:Tijucas, 20 Jan 1983 (Y fl),
Carautaet al. 4463 (GUA); Sao Leopoldo, 7 Apr 1943 (
fl-fr),Reitz 521 (GUA); Sombrio, 19 Nov 1944 (Y fl), Reitz
853 (S); nr.Tijucas,8 Feb 1980 (d), Reitz 7903 (GUA);Vila
Velha, Sombrio, 1 Nov 1959 (d), Reitz et al. 9339 (GUA);
Mun. Ararangua,Sombrio, 27 Feb 1952 (Y fl), L. B. Smith
et al. 5897 (US); Tubarao,Jan 1889 (Y fl-fr), Ule 1106 (P,
US). SiO PAULO:Campinas, 10 Jul 1968 (5), Carautaet
al. 620 (SPF, U, US), ( fl), Carautaet al. 621 (G, U, US),
622 (GUA, U, US); Mun. Iguape,5 km WSW of Iguape, 18
Feb 1965 (Y fl-fr),Eiten et al. 6196 (K, MO, NY, US); Mun.
Canan6ia,rd. Canan6ia-Pariquera-aqd, km 19, 6 Sep 1994
(Y fl-fr), V E Ferreiraet al. 67 (SP); Ilha do Cardosa,Res-
tinga de Itacuruqc,8 Oct 1980 (Y fl), Forero et al. 8682
(COL, SP); nr. Jundiaf,5 Aug 1951 (Y fl-fr), W. Hoehne
(SPF) 13234 (F, SP, SPF, US), (5), W Hoehne (SPF) 13237
(SPF); Sao Jose dos Campos, Jacaref,30 Aug 1949 (9 fl),
M. Kuhlmannet al. 2006 (K, NY, SPF, US); BarraBonita,
Rio Jacar&Guaqu,18 Jul 1991 (5), Romaniucet al. 1199
(SP); Rinopolis, 19 Jul 1991 (9 fl-fr),Romaniucet al. 1207
(SP); Mun. TeodoroSampaio, Reservado Morrodo Diabo,
2 Dec 1986 (9 fl-fr), Tamashiroet al. 18821 (UEC), 4 Dec
1986 (5), Tamashiroet al. 18844 (RB, UEC). SERGIPE:
Mun. SantaLuzia de Itanhi,nr.SantaLuzia de Itahi,27 Nov
1993 ( 9 fr),Amorinet al. 1475 (NY); betweenAreiaBranca
& Itabaiana,Serrade Itabaiana,25 Jan 1991 (9 fl-fr),Barros
2469 (SP).
PARAGUAY.Dtto. Paraguari,Cerro Palacios, 2 Mar
1988 (9 fl), Boswaldo 1363 (BG, MO, NY, TEX); Dtto.
Itapua,between CapitanMeza & PuertoTriunfo,Sep 1981
(d), FerndndezC. 3709 (G, MO, NY); Dtto. Alto Parana,
N. of Hemandarias,nr. Pto. Curupayty,Sep 1980 (5), Fer-
ndndez C. 4138 (COL, G, MO, NY), (9 fl-fr), Ferndndez
C. 4139 (G, NY); Dtto. Canendiyd,between Las Palomas&
Salto del Guaira,Sep 1981 (9 fr), FerndndezC. 4171 (COL,
G, MO, NY), (5), FernandezC. 4172 (G, MO, NY); Dtto.
Paraguari,Ybycui National park, 19-28 Mar 1980 (9 fr),
Hartshorn2490 (MO, NY); nr.Villa Rica, 10 Feb 1929 (9
fr), J0rgensen 3836 (A, C, F, MO, NY, S, US); Depto. Mi-
siones, Santiago, Estancia La Soledad, 20 Apr 1961 (d),
Pedersen 5923 (L); Dtto. Cordillera,Tobaty,22 Mar 1975
(9 fr), Schinini 10866 (G); Dtto. Amambay,35 km SE of
Bella Vista, 24 Aug 1980 (9 fl-fr),Schininiet al. 20584 (G);
Dtto. Guaira, Colonia Independencia,23 Dec 1986 (9),
Schinini et al. 25365 (G, K, MO); Dtto. Caaguazu, Caa-
guazu, 26 Jan 1951 (9 fl-fr), Sapper et al. 2178 (P); Dtto.
138
Central,Memby, 18 Nov 1981 (d), Vavreket al. 451 (MO,
US); Dtto. San Pedro, Primavera,15 Jul 1957 (d + 9 fr),
Woolston835 (C, K, NY, S).
ARGENTINA. CHAco: Las Palmas, 9 Sep 1917 (d),
J0rgensen 2158 (GH,MO, US). CORRIENTES: Dtto.Itu-
zaing6, Esteros del Ibera,LagunaIsip6, 15 Nov 1976 (d),
Arbo et al. 1463 (C, F, MO); Dtto. GeneralPaz, Ita Ibate, 1
Jan 1946 (d), Ibarrola 4060 (BM, K, S); Dtto. Ituzaing6,
Puerto San Antonio, Isla Apipe Grande,8 Dec 1973 (9 fl-
fr + d), Krapovickaset al. 23862 (G, RB); Dtto. Capital,
Corrientes, 15 Dec 1986 (9 fl-fr), Krapovickas40699 (A,
F, G, GB, INPA, LPB, MICH, MO, NY, SPF, TEX, USZ),
2 Jan 1987 (d), Krapovickas40700 (A, C, G, GB, INPA,
K, LPB, MICH, MO, NY, SPF, TEX, USZ); EstanciaSanta
Teresa, 13 Jan 1951 (d + 9 fl), Pedersen 945 (C, G, GH,
K, MO, NY, P, S, US); Dtto. Concepci6n, Carambola,24
Nov 1982 (9 fl-fr), Pedersen 13463 (A, C, L, MO, NY).
MISIONES: Nr. PuertoLe6n, 12-21 Jul 1914 (9 fr), Curran
714 (US); Posadas, La Granja, 15 Nov 1907 (d + 9 fl),
Ekman2013 (F, GH, NY, S); Dtto. Gral. Belgrano,Rd. 101,
km 5, 13 Dec 1983 (9), Hunzikeret al. 11026 (TEX); San-
tiago, Estancia La Soledad, 20 Apr 1961 (d), Pedersen
59231 (C, MO); SantaAna, 4 Jan 1913 (9 fl-fr),Rodr(guez
240 (BM, F); Dtto. Cainguas,rt. 14, km 252, 4 Jan 1950 (9
fl-fr), Schwindt2983 (LL, US).
Cecropia pachystachya is very common and wide-
spreadsouth of the Amazon basin. It is very variable
in the marginof the segments, being lobed or not, in
the density of the arachnoidindumentum,and in the
size of the trees. Leaves with dense arachnoidindu-
mentumare common in the southernand easternpart
of the species range, e.g., in Santa Catarinaand in
Paraguay.The material with dense arachnoidindu-
mentumon petioles, stipules,andspathesis quitesim-
ilar to that of C. metensis. Small trees are found in
the restingasalong the coast. Carauta1061 from Mi-
nas Gerais is distinct in the incisions of the lamina
down to near the petiole; in that respect it is similar
to C. concolor, but the indumentumof the trichilia
and the lobation of the lamina segments indicatethat
this collection belongs to C. pachystachya.Some col-
lections from Para (Dias et al. 219 and Egler 1018)
also show similaritiesto C. concolor in the incisions
of the lamina.
In K the collections Gardner3981 and 3982 are
representedby sheets with materialcorrectlyreferred
to Cecropia pachystachya, but also by sheets with ma-
terial referred to C. obtusa (and syntype for that
name) or sheets with leaves of C. saxatilis, on a Gar-
dener 3982 sheet combined with a pistillate inflores-
cence (with sessile spikes) of C. pachystachyaand on
a Gardner3981 sheet combined with a pistillate in-
florescence(with stipitatespikes), possibly belonging
to C. glaziovii or C. palmata. Althoughit is not quite
clear from the descriptionandplate, Miquelhas prob-
FLORA NEOTROPICA
ably based C. cyrtostachyaon a leaf of C. saxatilis
and a pistillate inflorescenceof C. pachystachya,the
latteris presentlydesignatedas the lectotypeelement.
The collections Glaziou 18500-18507 are partly
mixed, comprisingboth materialbelonging to C. gla-
ziovii and to C. pachystachya.
The species is probably introducedin the Cook
Islands (W R. Philipson 10145, US) and Guam (J. B.
Thompson390, US). The collections examined have
well-developedtrichilia.Whethermaterialintroduced
in Indonesia and Malaysia belong to this species or
to C. peltata is not clear.
The species is or has been in cultivationin Cuba,
HarvardTropical Garden, Soledad, Cienfuegos (20
Mar 1927, J. G. Jack 4355, A); Philippines, Lamao
(P. J. Wester48, A), and Los Banos (28 May 1914
[ Y], E. Quisumbing7933, MICH);Burundi,Bujum-
bura, University campus (6, M. Reekmans 9622,
MO); Zaire, Eala (6, CorbusierBoland 2061, MO;
J. Leemans83 P; 6 Oct 1945 [Y], J. Leonard76, P);
see also the discussion of C. peltata. Indicationsthat
this species has become naturalizedin Zaireare lack-
ing.
Vernacular names. Brazil: imbauba vermelha
(Bahia); torem (Ceara); embauibabranca (Pernam-
buco); pau de formiga, torem (Piauf); ambahu,
ambaiba-tinga,ambai, ambati, arvore da preguica,
ibaifba,imbauibabranca,imbaubao,pau de lixa (Rio
de Janeiro;also for C. glaziovii and C. hololeuca, see
Duarte, 1959). Paraguay:amba'i, amba'y or ambay.
39. Cecropia palmata Willdenow,Sp. PI. 4(2): 652.
1806; Berg, Acta Amazonica8(2): 162. 1978.Am-
baiba palmata (Willdenow) Kuntze, Revis. Gen.
PI. 2: 624. 1891. Type. Brazil. Para:Bel6m, Sieber
in herb. Hoffinannsegg s.n. (holotype: B, herb.
Willdenow,destroyed,photographsin G, US), re-
placed by Brazil. Para:Mun. Belem, Mocambo,8
Oct 1995 (6), Berg 1720 (neotype: MG; isoneo-
types: B, BG, COL, INPA, K, NY, RB, U, UB,
US).
V. Richter,Biblioth.Bot. 43: 19.
Cecropiabureauiana
1897. Type. French Guiana.Without locality, 1845
(6), Melinons.n. (holotype:P).
Tree,to 20 m tall. Leafy twigs 2-5 cm thick,green,
hispidulouswith uncinatehairs,also partlyhirtellous.
Lamina (sub)coriaceous,ca. 20 X 20 cm to 60 X 60
cm (to 100 x 100 cm), the segments 7-11, the free
parts of the upper segments (sub)obovateto elliptic
to oblong or to ovate, the incisions down to 5/10-8/
10; apices obtuse; upper surface scabrousto scabri-
SYSTEMATICTREATMENT 139

dulous, hispidulous;lower surface minutely puberu- 1978 (Y fl), Prevost 292 (P,U); Cayenne,CentreORSTOM,
lous (with straightto uncinatehairs)on the veins, with 18 Aug 1979 (5), Prevost 731 (P, U, US); Piste de St. Elie,
arachnoidindumentumin the areoles and on the mar- 28 Oct 1984 (st), Riera 650 (BG, P).
BRAZIL. AMAZONAS: Manaus,campusINPA, 31 Aug
gin; lateral veins in the free part of the midsegment
1973 (Y fl), Berg 252 (F, K, MO, RB, U, US); rd. Manaus-
ca. 15-20 pairs, marginally loop-connected, often
Caracarai,km 20, 21 Sep 1973 (st), Berg 284 (U); Manaus,
branched;petiole 20-60(-90) cm long, minutely pu- campusINPA, 8 Nov 1979 (Y fr), D. Coelho (INPA)90393
berulous;trichilia fused, the brown indumentumin- (INPA);Rio Branco, Caracarai,8 Jan 1924 (Y fl-fr), Kuhl-
termixed with short white to brownish (unicellular) mann 1085 (RB 20199); Manaus,Pensador,Aug 1910 (d),
hairs;stipules 7-15 cm long, pale to dark(brownish) Ule 8837 (G, K, MG, US). CEARA: Nr.Lagoade Messejana,
red, often subpersistent, sparsely hirtellous, some- 20 Jan 1968 (5), Carauta 538 (GUA, U); Crato, Chapada
times also with sparsearachnoidindumentumoutside, do Araripe,20 Jan 1968 (st), Carauta553 (GUA, U), (5),
densely to sparsely hairy or glabrous inside. Stami- Carauta 554 (G, GUA, U); Serra de Maranguabe,14 Sep
1908 (5), Ducke (MG) 1690 (MG); Mun. Pacatuba,Itai-
nate inflorescencesin pairs, pendulous;peduncle 6-
tinga, 7 Aug 1957 (5), Ducke 2615 (MG); Guaraciaba,19
18 cm long, minutely puberulousor also with sparse
Jul 1979 (5), JordyFilho 53 (RB). MARANHAO:Mun.Lago
arachnoidindumentum;spathe 10-17 cm long, white Verde,rd. Alto Alegre-Lago Verde,km 9, 22 Mar 1985 (5),
or sometimes reddish, with dense arachnoid indu- A. B. Anderson et al. 2073 (MG, NY); Mun. Moncfio, Rio
ment, also sparsely hirtellous outside, mostly gla- Turiacu,nr.Urutawy,12 Feb 1985 (5), Balee etal. 809 (K);
brous or sometimes sparsely hairy inside; spikes 4- Mun. Buriti Bravo, Rio Itapecuru,Serro dos Papagaios,23
6(-11), (3-)8-15 X 0.8-1.8 cm, with stipes 1-2.5 cm Feb 1983 (Y fl), Rosa et al. 659 (BG); rd. BR.136, km 15,
long andminutelypuberulous;rachishairy.Staminate 10 km from Araguana,8 Dec 1978 (Y fl-fr), Rosa et al.
flowers: perianth tubular, 2.5-4.5 mm long, with 2901 (MG, NY, U); Mun. Buriti Bravo, Rio Itapecuru,23
Feb 1983 (Y f-fr), J. U. Santos et al. 659 (MG); Sao Luiz,
dense arachnoid indumentum below the apex, the
Andiroba,8 Aug 1980 (st), M. G. Silva 5724 (MG). MATO
apex convex andglabrous;filaments ? swollen, often
GROSSO: Nr. Base Camp, ca. 270 km N of Xavantina,
basally connate;anthers1-2 mm long, appendiculate, 12049'S, 51046'W, 26 Oct 1968 (5), Harley 10838 (E, K,
detached at anthesis, remainingattachedto the fila- NY, P, RB, U); Mun. Vila Bela da Sma. Trinidade,rd. Vila
ment by stretchedspiral thickenings.Pistillate info- Bela-Rio Verde,FazendaPousadado Guapore,18 Aug 1997
rescences usually in pairs, pendulous;peduncle 20- ( fr), Hatschbachet al. 66999 (BG). PARA:Mun. Barcar-
40 cm long, hispidulous, or also with arachnoidin- ena, Itupanema,9 Dec 1985 (Y fl-fr), Amorozo240 (MG);
dumentum;spathe 12-16 cm long, the color and in- Belem, 16 Sep 1942 (5), Archer 7645 (NY, US); Serrados
dumentumas in the staminateinflorescence;spikes 4, Carajas,Serra Norte, ca. 12 km E of AMZA exploration
camp, 16 Oct 1977 (Y fl-fr), Berg et al. 586 (K, MG, MO,
6-14 X 0.6-0.8 cm, to 20 X 1.5 cm in fruit, sessile;
NY, U); Belem, grounds of IAN, 7 Nov 1962 (Y fl), Cua-
rachis hairy. Pistillate flowers: perianth ca. 2 mm
trecasas 26653 (P, US), ( 9 fl-fr), Cuatrecasas26655, 26658
long, with arachnoidindumentumbelow the apex out- (P, US); Alto Curuadna,16 Oct 1962 (9 fl), Duarte 7189
side, also in the lower partof the style channelinside, (INPA, RB); E of Faro, Igarapedo Cauhy,28 Jan 1910 (9
the apex convex andglabrous;style long, ? S-shaped fl-fr), Ducke (MG) 10564 (MG); Maraj6,Pacoval, Sep 1896
to straight,minutelypuberulous;stigmapenicillateto (5), Huber (MG) 494 (MG); Capitao Poco, rd. Matuttii-
comose. Fruit oblongoid, ca. 3 mm long, almost Piri, 13 Jan 1974 (9 fl-fr), Oliveira 6139 (MG); Tucuruf,
smooth. Lago do Cagancho, 13 Oct 1983 (5), Revilla et al. 8642
(INPA, NY); Maraj6,Soure, Teso, 29 Nov 1906 (9 fl-fr),
Distribution (Fig. 18.2). Fromthe lower andmid- R. S. Rodrigues(MG) 7841 (G, MG), 1 Dec 1906 (d), R. S.
dle Amazon basin throughnorthernFrenchGuianato Rodrigues (MG) 7842 (G, MG). PARAiBA: Areia, Pica dos
northeasternSurinameand to northeasternBrazil and Postes, Mata de Pau Ferro,4 Dec 1980 (5), Fevereiroet al.
northeasternBolivia, in non-inundatedforest,in some M.169 (K); Mamanguape,14 Jan 1970 (5), Carauta 965
regions in savanna forest, common in secondary (GUA, U); JoaoPessoa, Miramar,18 Jan 1970 (5), Carauta
969A (C, G, GUA, U, US), (9 fl-fr), Carauta969B (GUA,
growth,at low elevations.
NY, US). PERNAMBUCO:Recife, Dois Irmaos,23 Jan 1972
Representative specimens examined. SURINAME. (9 fl-fr), Carauta et al. 1471 (GUA, U); Mun. Caruaru,
Mapane Cr., 20 May 1970 (? fl), Teunissen(LBB) 12750 Brejo dos Cavalos, 21 Jan 1972 (5), Strang s.n. (GUA).
(F, U). ROND6NIA: 1 km S of Riberao,rd. Abuna-Guajara-Mirfm,
FRENCH GUIANA. Nr. Sinnamary,29 Nov 1977 (Y 27 Jul 1968 (9 fl-fr), Prance et al. 6574 (INPA, K, MG,
fl-fr), Berg 784 (K, NY, U, US); Rorota Lake, 6 Oct 1978 MO, NY, S, U, US). RORAIMA: Foothills of Serra Tepe-
(Y fl), Grenand1690 (P, U); Oyapock R., above Saut Mou- quem, IgarapePaparu,13 Feb 1967 (9 fl-fr), Prance et al.
touci, 15 May 1970 ( fl-fr), OldemanT 737 (NY, U); Yar- 4361 (INPA, K, MG, MO, NY, U, US); Ilha de Maracd,
oupi R., nr. Saut Tainoua, 18 Apr 1970 (Y fl-fr), Oldeman SEMA Ecological Station, 27 Feb 1987 (9 fl-fr), Ratteret
3118 (U); nr.Cayenne,between Cabassou& Rnmire,11 Sep al. 5503 (BG, E, INPA, K, NY). SERGIPE: Mun. Areia
140
Branca, Estaqao Ecol6gica da Serra de Itabaiana,23 Jan
1992 (fl), Carautaet al. 6485 (GUA); Itabaiana,ca. 41 km
W of Aracaju, 5 km W of Ar6ia Branca, BR.235, 22 Jan
1992 (d ), Thomaset al. 8903 (NY).
BOLIVIA.SANTA CRUZ: Prov. Velasco,ParqueNa-
cional Noel Kempff M., E of campamentoLos Fierros, 18
May 1995 (d), Abbott 16816 (BG); Prov.Velasco, campa-
mento El Refugio, 17 May 1994 (d fl), Guilln et al. 1314
(USZ); Prov. Velasco, Parque Nacional Noel Kempff M.,
between Moira & Chore, 14 Jul 1996 (Y dl-fr), Jardimet
al. 3067 (BG); Prov.Velasco, ParqueNacionalNoel Kempff
M., campamentoLos Fierros,5 May 1994 (d), Quevedoet
al. 2572 (USZ); Prov. Velasco, Parque Nacional Noel
KempffM., between Los Fierros& Moira, 2 Jun 1993 (d),
Saldias et al. 2914 (USZ), (Y fl), Saldias et al. 2915 (USZ).
Cecropia palmata is closely relatedto C. glaziovii.
These species appearto be allopatric.Some evident
differences between these species are listed underC.
glaziovii. Consideringthe similaritiesin the staminate
flowers,these two species mightbe relatedto C. saxa-
tilis.
An occasional monoecious tree has been encoun-
tered in this species. Cavalcante 1392 is unusual in
that the staminate inflorescences have short (7-10
cm), slender(ca. 0.3 cm diam.) spikes with short(0.4-
0.5 cm) stipes; the perianths lack the characteristic
arachnoid indumentum and the stamens are small
(and the anthersprobablynot detachedin the ususal
way). These inflorescencesresemble those of Cecro-
pia concolor.
The species is and has been in cultivationin the
gardenof Museo Goeldi, Belem, representedby Ca-
valcante 1391, 1392, 1394, 1395, and 1396 (MG),
made in 1961.
Vernacular names. Brazil: torem (Ceara);ama'-
y-puku (Ka'apor, Maranhao); imbauba vermelha
(Para').
40. Cecropia pastasana Diels, Notizbl. Bot. Gart.
Berlin-Dahlem15: 368. 1941. Type.Ecuador.Pas-
taza: Mera, ca. 1000 m, 16 Nov 1938 (d),
Schultze-Rhonhof 2996 (holotype: B, destroyed,
duplicatesnot traced;here replaced by: Ecuador.
Morona-Santiago:Between Plan del Milagro &
Limon (Gral.Plaza], ca. 1500 m, 30 Jan 1981 [d ],
Berg 1236 [neotype: QCA; isoneotypes: AAU,
BG, COL, GB, MO, NY, U]). Fig. 37
Cecropiasucrensis Cuatrecasas,Revista Acad. Colomb.
Ci. Exact. 6(22/23): 297. 1945. Type. Colombia.Ca-
queta:Rio Hacha, Sucre, 1000-1040 m, 7 Apr 1940
( Y fl-fr), Cuatrecasas9204 (holotype:COL;isotype:
F).
FLORANEOTROPICA
Tree, to 25 m tall. Leafy twigs (I-)2-5(-8) cm
thick, (dark)red-brownor greenish, sometimes with
a thin bluish waxy layer, sparselypuberulousto stri-
gillose to subglabrous,usually with ? dense brown
pluricellularhairs;pith scarceandwhitish,sometimes
copious and brown. Lamina chartaceousto subcor-
iaceous, ca. (20 X 20 cm to) 35 X 35 cm to 85 X 85
cm, the segments 7-9(-10), the free parts of upper
segments ovate or elliptic to obovate, the incisions in
the upper part of the lamina down to 3/10-6/10 or
down to 8/10, in the lower part less deeply incised,
entire or lobate; apices obtuse to roundedto acute or
to subacuminate;upper surface scabridulousto al-
most smooth, puberulousto hispidulousto hirtellous
to strigillose and initially with ? dense to sparse to
arachnoid indumentum;lower surface hirtellous to
subtomentoseto tomentellous or to minutely puber-
ulous on the veins, with arachnoidindumentumin the
areoles, rathersparse and short, or along the margin
only; lateralveins 7-12 or 12-16 pairs,submarginally
loop-connected,some or most of them branched;pet-
iole (I0-)20-75 cm long, with dense to sparsebrown
pluricellularhairs or also minutely puberulous,oc-
casionally with a thin bluish waxy layer; trichiliaon
the abaxiallybulging (subscrotiform)base of the pet-
iole, fused or separate,the brown indumentuminter-
mixed with short to ratherlong whitish to brownish
(unicellular)hairs;stipules ca. 10-25, caducous,dull
darkred, red-brown,pinkish,or yellowish green,with
dense brown pluricellular hairs or also minutely
appressed-puberulousoutside, densely to sparsely
hirtellousto subsericeous inside. Staminateinflores-
cences in pairs, often subtendedby caducous bracts,
to 8 cm long, the peduncle erect and the spikes pen-
dulous; peduncle 4.5-9 cm long, densely to sparsely
subhispidto hirtellousto subhirsuteor (sub)glabrous;
spathe 15-32 cm long, dull red (to pinkish) or yel-
lowish green (with longitudinal red stripes), with
dense brown pluricellularhairs or also minutely pu-
berulous outside, (sparsely) hairy inside; spikes 10-
25, (6-)12-28 X 0.3-0.5 cm, sessile or with stipes to
0.3 cm long and sparselytomentose;rachishairy.Sta-
minate flowers: perianth tubular,0.8-1.5 mm long,
glabrous,the apex slightly convex to plane; filaments
flat; anthers0.5-0.6 mm long, appendiculate,at an-
thesis detached,remainingattachedto the filamentby
2 filiformconnectionsbetween the connectiveandthe
upper margin of the filament (?). Pistillate inflores-
cences in pairs or solitary,often subtendedby cadu-
cous bracts, to 8 cm long, the peduncle erect to de-
flexed and the spikes erect to ? pendulous;peduncle
4-10 cm long, densely to sparselyhirtellousto hirsute
and with dense brown pluricellularhairs or (sub)-
SYSTEMATICTREATMENT 141

I ( -~~~~~~~~~~~~~~~~2

9 6

FIG. 37. Cecropiapastasana. 1. Lamina and stipules. 2. Apex of lamina and venation.3. Pair of staminateinflores-
cences with spathes and base of petiole with trichilium. 4. Staminate inflorescence at anthesis and base of petiole with
trichilium(Berg 1236). 5. Staminateflower. 6. Stamen and filamentafter detachmentof anther(Franco et al. 4691). 7. Pair
of pistillate inflorescenceswith spathes.8. Pistillate inflorescenceat anthesisand base of petiole with trichilium.9. Pistillate
flower. 10. Fruit (Berg et al. 1761).
142 FLORA NEOTROPICA

glabrous;spathe 10-17 cm long, the color and indu- dense brown pluricellularhairs!) on the leafy twigs,
mentumas in the staminateinflorescence;spikes (2-) petioles, and stipules, the more or less clearly whitish
4-6(-7), 10-16 X ca. 0.4-0.8 cm, to 28 X 1.3 cm in uppersurfaceof the lamina,andonly occasionalpres-
fruit, sessile to subsessile (or with stipes to 0.8 cm ence of a waxy layer on leafy twigs and petioles. In
long), glabrous.Pistillateflowers: perianth1.5-2 mm the northernpart of the range of this species, almost
long, with arachnoidindumentumbelow the apex out- coinciding with the range of C. andina, this species
side, also in the style channelinside, the apex slightly occurs at elevations to ca. 1500 m, below the eleva-
convex, smooth or sparsely granulate style long, tional range of C. andina (1500-2300 m). In the
straight,minutely puberulous;stigma comose. Fruit southernpart of the distributionrange, C. pastasana
ellipsoid to oblongoid, ca. 1.5 mm long, smooth. can be found at elevationsto 2300 m. In the northern
part of the species range, the arachnoidindumentum
Distribution (Fig. 18.8). From Colombia (Ca-
on the uppersurfaceis ratherthin and the leaves look
queta)to Peru (Pasco), in premontaneto montanefor-
whitish only if seen from some distance, whereasin
est, at ca. 900-2300 m.
the southernpart of the species range (Huainucoand
Specimensexamined.COLOMBIA.CAQUETA:Mun. Pasco) they are more often clearly whitish above. In
Florencia, rd. to Gabinete, ca. 1300 m, 18 Oct 1993 (d), the northernpartof the species range,the incisions of
Franco et al. 4512 (BG,COL).PUTUMAYO: Rd. Mocoa- the lamina are mostly down to 5/10 or 6/10 and the
Pasto, La Teboida, 1300 m, 19 Feb 1995 (5), Franco et al. number of pairs of lateral veins in the free part of
4691 (BG, COL, HUA, US), (Y fl-fr), Franco et al. 4692 the midsegment is 7-12, but in the southernpart of
(BG, COL, HUA, US); Mocoa, Vrda.La Campucana,Finca
the range the incisions are often deeper, down to 8/
La Mariposa, 1350 m, 26 Apr 1994 (5), Franco et al. 5343
10, and the numberof pairsof lateralveins in the free
(BG), (Y fl-fr), Franco et al. 5344 (BG).
ECUADOR.MORONA-SANTIAGO: Rd. Gualaquiza- part of the midsegmentis 12-16.
San Juan Bosco, km 17, 1300 m, 16 Feb 1994 (5), Berg Cecropia pastasana is the white- or whitish-
1693 (BG, QCNE). PASTAZA: Mera, 25 Mar 1940 (Y fl), leaved species occurringon the easternslopes of the
Lugo 118 (S). TUNGURAHUA:Rd. Bafios-Puyo, between Andes, in the northernpartof its range(southernCo-
Rfo Verde & Rio Negro, ca. 1400 m, 10 Jan 1981 (9 fl), lombia and Ecuador),at elevations below that of the
Berg 1201 (AAU, BG, COL, QCA, U). ZAMORA- white-leaved C. telenitida, and in the southernpart
CHINCHIPE:Rd. Zamora-Gualaquiza,between Namirez & (Peru) below that of the white-leaved C. albicans.
Yantzatza,ca. 900 m, 3 Jan 1991 (5), Berg et al. 1655 (BG, Ants are usually present, but they are not always of
GB, LOJA,NY, QCA); rd. Loja-Zamora,between Sabanilla the genus Azteca.
& Zamora,ca. 1300 m, 4 Jan 1991 (9 fl), Berg et al. 1664
(BG, LOJA,QCA).
PERU.AMAZONAS: Prov. Bongara, 4-8 km E of Po-
macocha, rd. to Rioja, 2100-2200 m, 9 Feb 1984 (9 fl-fr), 41. Cecropia peltata Linnaeus, Syst. Nat., ed. 10.
Gentry et al. 45194 (BG, K, MO, NY). HUANUCO:Rd. 1286. 1759; Woodson & Schery, Ann. Missouri
Huanuco-Tingo Maria, below Chinchao, ca. 1800 m, 23 Bot. Gard. 47: 173, t. 60. 1960; Velasquez,Acta
Nov 1997 (9 fl-fr),Berg et al. 1733 (BG, COL, MOL), (5), Bot. Venez. 6: 51, t. 11. 1971; Burger,Fieldiana,
Berg et al. 1736 (BG, COL, MOL), 24 Nov 1997 (5), Berg Bot. 40: 126, t. 23. 1977; Berg in Gorts-vanRijn,
et al. 1738 (BG, COL, MOL); rd. Huanuco-TingoMaria,N Fl. Guianas,Ser. A, Fasc. 11: 101. 1992. Ambaiba
of Carpishpass, 54 km NE of Huanuco,ca. 2300 m, 6 Dec
peltata (Linnaeus)Kuntze,Revis. Gen. PI. 2: 623.
1981 (d), Plowman et al. 11177 (BG, NY, UMS). JUNiN:
1891. Type.Jamaica.Withoutlocality,Anonymous
Prov. Chanchamayo,rd. to Mina Pichita, 1900 m, 2 Feb
1999 (5), Nuiiez 65 (BG). PASCO: Rd. Oxapampa-Llaupi- s.n. (holotype:LINN. 1159.2). Fig. 38
Cerrode Pasco, ca. km 15, ca. 1750 m, 29 Nov 1997 (9 fl- Cecropia humboldtiana Klotzsch, Linnaea 20: 530.
fr), Berg et al. 1761 (BG, COL, MOL); rd. Oxapampa-Pu- 1847. Type. Colombia. Withoutlocality, (6), Hum-
ente Paucartambo,nr. Oxapampa,ca, 1600 m, 30 Nov 1997 boldt s.n. (holotype:B, destroyed,photographin F).
(5), Berg et al. 1763 (BG, COL,MOL).SANMARTiN: Prov. Cecropia schiedeana Klotzsch, Linnaea 20: 531. 1847.
Rioja, rd. Rioja-Pomacocha,ca. km 50, 900 m, 8 Dec 1997 Type.Mexico. Veracruz:Papantla,Schiede1114 (ho-
(9 fl-fr),Berget al. 1794 (BG, COL, MOL). lotype: B, destroyed,photographsin F, MICH,US).
Cecropia surinamensis Miquel, in Martius, Fl. Bras.
Considering the similarities in many characters
4(1): 143, t. 46, 1. 1853;Berg, Acta Amaz6nica8(2):
and similarities in the variationpatterns,this species 165. 1978. Amaiba surinamensis(Miquel) Kuntze,
appearsto be very closely relatedto Cecropiaandina. Revis. Gen. P1.2: 624. 1891. Type. Suriname.With-
It can be distinguishedby the consistent presence of out locality, (5), Fockes.n. (holotype:U).
trichilia on a subscrotiformbase of the petiole, often Cecropia digitata Tenoreex Miquel var. grisea Miquel
also by the presence of indumentum(includingoften in Martius,Fl. Bras. 4(1): 149. 1853. Type. Material
SYSTEMATICTREATMENT 143

UliA l

FIG. 38. Cecropia peltata. 1. Lamina, reduced. 2. Apex of lamina and venation. 3. Stipules and young leaf. 4.
Staminate inflorescences with spathes and bases of petioles with trichilia (Berg 978). 5. Staminate flower (Franco et al.
5561) and stamen (Franco et al. 4646). 6. Pistillate inflorescences, one with spathe and the other at anthesis, and bases of
petioles with trichilia (Stevens 3341). 7. Pistillate flower. 8. Fruit (Franco et al. 5566).
144 FLORANEOTROPICA

cultivatedin HortusSchonbrunn,from materialcol- cences erect to deflexed; peduncle 2-12.5 cm long,

lected in Mexico (holotype: M, photograph in sparsely to densely puberulousto hirtellous to sub-
MICH). hispid(ulous);spathe 2.5-7 cm long, pinkish (to pur-
CecropiapropinquaMiquel, in Martius,Fl. Bras. 4(1):
plish), greenish or whitish, puberulousto hirtellous
149. 1853. Type. Material cultivated in Hortus
and often also with arachnoidindumentumoutside,
Schonbrunn,from material collected in Mexico by
Schott in 1852 (holotype: M-n.v., photographsin F,
(sub)sericeous inside; spikes ca. (10-)15-25(-60),
MO). 0.5-6 X 0.2-0.25 cm, with stipes to 0.8 cm long and
Cecropiascabrifolia V. Richter,Biblioth. Bot. 43: 15, t. hairy; rachis hairy. Staminate flowers: perianth tu-
2. 1897. Type. Nicaragua. Grenada,Jun 1869 (5), bular, 1-1.5 mm long, with short and stiff hairs or
L,vy 52 (holotype: P; isotypes: C, G, P, fragmentex sometimes short arachnoid indumentumbelow the
C in F). apex, the apex plane, smooth; filamentsflat; anthers
CecropiaarachnoideaPittier,Contr.U.S. Natl. Herb.18: ca. 0.5-0.6 mm long, appendiculate,detachedat an-
226. 1917. Type. Panama.Panama:Nr. Matachin,23 thesis, reattachedto the marginof the apertureby the
Jun 1911 (Y), Pittier 4056 (holotype: US; isotype:
appendages at anthesis. Pistillate inflorescences in
US).
pairs, erect to pendulous; peduncle 3-10(-16) cm
Cecropia asperrima Pittier,Contr.U.S. Herb. Natl. 18:
227. 1917. Syntypes. Nicaragua. Without locality,
long, sparsely to densely puberulousto hirtellous to
1853-1856 (d + Y), C. Wrights.n. (US, 5 + Y; hirsuteor to subhispid,often also with arachnoidin-
GH, 5). dumentum;spathe 3.5-6 cm long, the color and in-
Cecropiadielsiana Snethlage,Notizbl. Bot. Gart.Berlin- dumentumas in the staminateinflorescence;spikes
Dahlem 8: 363. 1923; Berg, Acta Amaz6nica 8(2): 3-4(-5), 1-4 X 0.3-0.4 cm, to 9 X 1.1 cm in fruit,
168. 1978. Syntypes. Brazil. Roraima:Sao Marcos, sessile or with stipes to 0.5 cm long and hairy;rachis
Jan 1909 (5), Ule 7891 (B, destroyed;G, K) & (Y), hairy. Pistillate flowers basally connate;perianth 1-
Ule 7891 (B, destroyed;G, K, MG). 1.5 mm long, with arachnoidindumentumbelow and
Cecropia hondurensisStandley, Publ. Field Mus. Nat. on the apex outside, absent(or in and below the style
Hist., Bot. Ser. 4: 302. 1929. Type. Honduras.Atlan-
channel)inside, the apex flat to slightly convex; style
tida: Nr. Tela, 14 Dec 1927-15 Mar 1928 (?), Stan-
short; stigma peltate. Fruit ovoid to ellipsoid, ca. 2
dley 54528 (holotype:F; isotype: US).
Cecropia goodspeedii Cuatrecasas,Revista Acad. Col- mm long, tuberculate,darkbrown.
omb. Ci. Exact. 6(21): 67. 1944; Not. Fl. Colomb.6 Distribution (Fig. 18.3). From southemnMexico
(Trab.Com. Bot. etc. Colomb):41. 1944. Type. Co-
to northern South America (Colombia, Venezuela,
lombia. Antioquia:Rio Cauca, PuertoValdivia, 17-
Guyana,Suriname,and northernfringe of the Brazil-
20 Feb 1942 (Y), Cuatrecasas et al. 30079 (holo-
type: COL; isotypes: F, US).
ian part of the Amazon basin), also in Jamaicaand
Trinidad,in wet to deciduousforest areas,commonly
Tree, to 15(-25) m tall. Leafy twigs 1.5-4 cm in secondary growth, roadsides, pastures,etc., at el-
thick, green, hispidulous with curved to uncinate evations to 1000 m, sometimes to 1500 m, or (in Co-
hairs. Lamina chartaceousto subcoriaceous;ca. (10 lombia: Santander)to 2000 m.
X 10 cm to) 20 X 20 cm to 60 X 60 cm, the segments
Representative specimens examined. MEXICO.
(7-)8-10(-1 1), the free parts uppersegments oblong
CAMPECHE: 2 mi S of Ruinas Edzna, 27 Aug 1966 (d),
or elliptic, the incisions down to ca. 5/10-7/10; apices
Baird 1017 (US); Ruinas Edzna, 27 Aug 1966 (d), Baird
short-acuminateto rounded;upper surface scabrous, 1018(US);30 kmE of Escarcega, rd.Escarcega-Chetumal,
hispidulous; lower surface minutely puberulous,in- 12 Jul 1983 (dc), Cabrera5099 (MO). CHIAPAS: Mun.Fron-
termixed with rathersparse longer hairs, with arach- tera Comalpa, Hwy 190, 3 mi S of Rio San Gregorio, 29
noid indumentumin the areoles, also on the reticulum Apr 1965 (d), Breedlove9822 (CAS, LL, MICH,US); Mun.
or extending to the main veins; lateral veins in the Ocozocoautla,45 km N of Ocococoautla, 17 Oct 1971 (d),
free part of the midsegment (8-)10-15(-17) pairs, Breedlove et al. 20725 (MEXU); Mun. Catzadja,E of Ba-
marginally (or submarginally)loop-connected, usu- jadas Grandes, 11 Oct 1972 (d), Breedlove28781 (CAS);
ally branched;petiole (10-)20-50 cm long, puberu- nr. Judrez,NW of Pichucalco, 31 Aug 1965 (d), Chavelas
et al. 2463 (MEXU); Mun. Palenque, betweeen Salto de
lous or also with arachnoid indumentum;trichilia
Agua & Ruinas Palenque, 30 Jul 1967 (9 fr), Clarke s.n.
fused, the brown indumentumintermixedwith short (CAS, US); Mun. Ocosingo, Boca de Chajul,27 Apr 1993
(or sometimes with ratherlong) white hairs; stipules (9 fl-fr), Dminguez V 698 (MEXU); Pueblo Nuevo, Solis-
3-10(-12) cm long, pinkish or reddish, sometimes tahuacdn,1260 m, 11 Aug 1967 (d), G6mez-Pompa2568
subpersistent, strigose to hirtellous and often also (MEXU);Mun. Ocosingo, nr. San Javier,23 May 1991 (6),
with arachnoid indumentum outside, sparsely to Gonzalez-Espinosaet al. 1486 (MEXU);5.6 km E of Chiapa
densely (sub)sericeous inside. Staminate infiores- de Corzo, 15 Jul 1966 (6), Laughlin 1282 (CAS); Mun.
SYSTEMATICTREATMENT
TuxtlaGutierrez,El Zapotal,SE of TutxlaGutierrez,19 Aug
1988 (Y fr), Palacios E. 66 (CAS); nr. Ocozocoautla, 12
Nov 1988 (Y fl-fr),Reyes G. et al. 1266 (BM); nr.Simojovel
de Allende, 1000 m, 10 Oct 1967 (Y fr), Shilom Ton3075
(CAS). OAXACA: Hwy. 185, 4 km N of MatfasRomero, 24
Jul 1958 (Y fl), King 715 (LL, MICH, TEX, US); Ubero,
Apr 1937 (d), Ll. Williams9264 (MICH, US). QUINTANA
Roo: 26 mi S of Felipe Carrillo Puerto, 5 Jul 1966 (6),
Baird 1024 (US); 1 E of Cancun, 12 Sep 1982 (6), Cabrera
et al. 3483 (MO); 30 km E of Escarcega, 12 Jul 1983 (6),
Cabrera5099 (MEXU);Mun. Oth6nP. Blanco, 9 May 1982
(6), Chan et al. 1432 (MEXU); Coba, 10 Aug 1975 (6),
Lopez F 537 (MEXU);nr.Felipe CarrilloPuerto,7 Jul 1989
( fl), Serralta P. 34 (MEXU); Cancun, nr. airport,9 May
1980 (6), Tellez et al. 2189 (MEXU, MO); 17 km N of
Felipe Carrillo Puerto, 11 Jun 1980 (6), Tellez et al. 2510
(MEXU); nr. Andr6sQuintanaRoo, 4 Sep 1980 (d), Tellez
et al. 3352 (MEXU); 2 km S of Tres Garanitas,5 Sep 1980
(Y fl), Tellezet al. 3425 (MEXU). TABASCO: Nr. airportof
Villahermosa, 14 Oct 1988 (Y fl-fr), Ascencio R. 140
(MEXU); 10-40 km W of Huimanguillo,20-29 Mar 1963
(6), Barlow 30 (BM, MICH);Carreteradel Golfo, km 24,
17 Aug 1977 (Y fl-fr), Cdrdenas 186 (MEXU); Balancan,
La Palma, 1-6 Jan 1939 (6), Matuda 3240 (GH, MEXU,
MICH); Lomas de San Sebastian, 27 Feb 1889 (6), Rovi-
rosa 383 (K, NY). VERACRUZ: Isla Villa Azueta, 24 Dec
1968 (? fr), Hernandezet al. 488 (LL, MEXU, NY); Co-
atzacoalcos, Isthmus of Tehuantepec,Jan 1895 (6), C. L.
Smith 1040 (EAP, GH, MO, NY, US). YUCATAN: Chichan-
kanab,(d), Gaumeret al. 1459 (BM, C, GH, MO, US); nr.
Piste, May-Aug 1938 (6), Lundell et al. 7548 (MICH);
Xcoochle, ruins of Uxmal, 16 Sep 1865 (d), Schott 639
(BM, MO, US); ChichenItza, 29 Jun 1932 (6), Steere 1700
(MICH).
GUATEMALA. ALTA VERAPAZ: Nr. Sepacuite, Jun
1905 (Y fl-fr), Cook et al. 336 (US); Chamiquin,Apr 1889
(6), Donnell Smith1675 (US). IZABAL: Rio Sauce, 0-2 km
N of Lake Izabal, 21 May 1966 (d fl-fr), G. C. Jones et al.
3465 (EAP, LL, MICH, NY, US); Morales, 10 Mar 1907
(6), Kellerman6416 (MO, US); Sinai, ca. 30 km SW of
Santo Tomas de Castillo, 15 Mar 1988 (6), Marshall et al.
394 (BG, MO, NY); Rio Motagua,nr.Quirigua,6 Jun 1905
(6), Pittier 385 (US); Quebradas,19-22 May 1919 (9 fl),
Pittier 8626 (NY, US); nr. Quirigua, 15-31 May 1922 (d),
Standley24143 (GH, NY, US). PETEN: Lake Peten Itza, be-
tween Remate& San Jose, 25 May 1960 (9 fl-fr), Contreras
1007 (LL, NY, S); Tikal, 5 Aug 1960 (9 fl-fr), Contreras
1390 (LL, NY, S); Macanche,21 May 1966 (9 fl), Contreras
5832 (LL, US), 22 May 1966 (6), Contreras5845 (EAP,
GH, LL, TEX); SantaTeresa,Rio Subin, 13 Apr 1933 (d),
Lundell2902 (GH, MICH,NY, S); La Libertad,25 Apr 1933
(9 fl), Lundell 3006 (MICH, NY), (6), Lundell 3016
(MICH, NY, S, US); Lake Zoty, 18 May 1933 (9 fl-fr),
Lundell 3295 (MICH, NY, S), (d), Lundell 3296 (MICH,
NY); Tikal National Park,Tikal, 8 May 1959 (6), Lundell
15972 (LL, NY); SantaElena, rd. to Sacpuy,9 Sep 1971 (9
fr), Tun Ortfz 1899 (EAP, G, MO, S). ZACAPA: Quirigua,
18 Nov 1970 (9 fl-fr), Harmonet al. 4932 (GH, MO); 8 mi
145
E of Lobo, 8 Jul 1975 (9 fl-fr), Watkinset al. 726 (LL, MO,
NY).
BELIZE. Cayo Distr., 36 mi W of Belize, 3 Jul 1966
(9 fl-fr), Baird 1023 (US); Belize Distr.,NorthernHwy., mi
42.5, 27 May-6 June 1973 (6), Croat 23990 (MO); Belize
Distr.,NorthernHwy., mi 42.5, N of MaskallR., 7 Jun 1973
(9 fl), Dwyer 11017 (GH, MO); Corozal Distr., 1931-1932
(d), Gentle 455 (MICH,NY, S); Maskall, 15 May 1934 (9
fl), 8 Mar 1935 (9 fl-fr), Gentle 1566 (A, MICH,MO, NY),
29 May 1935 (6), Gentle 1664 (GH, K, MICH, MO, NY);
Corozal Distr., Alfonsville, 8 Jul 1933 (6), Lundell 4813
(MICH, NY, S), (9 fl-fr), Lundell 4814 (MICH, NY); Or-
ange WalkDistr.,between OrangeWalk& BenqueViejo, 20
Aug 1980 (9 fl-fr), Suttonet al. 152 (BM, MO); Altun Ha,
24 Aug 1970 (9 fl-fr), Wiley572 (MO, NY).
HONDURAS. ATLANTIDA: San Jose de Texiguat, 30
km NE of Tela, 20 Jun 1989 (9 fr), Galo 3 (BM); nr. La
Ceiba, 6 Aug 1938 (9 fl-fr), Yunckeret al. 8844 (BM, G,
MO, S, US). COL6N:Rfo Guaimoreto,4.5 mi NE of Tru-
jillo, 20 Feb 1981 (9 fr), Saunders 1035 (BM, NY, TEX).
COPAN:SantaRosa de Copan,10 May 1987 (6), Blackmore
et al. 3797 (BM, MO). GRACIAS A DIos: LagunaIbans,28
May 1985 (juv), Knees 2812 (BM). ISLAS DE LA BAHiA:
Isla de Roatan,30 Mar 1984 (9 fl), Nelson et al. 8737 (MO,
NY). MORAZAN: Rio Jicarito,drainageof Rio Yeguare,22
Jun 1948 (6), Glassman 1694 (EAP,NY); El Zamorano,6
Jul 1948 (6), Molina R. 427 (EAP), 15 Nov 1945 (d), Val-
erio R. 3593 (EAP, GH, US). OLANCHO: Between Campa-
mento & border Olancho/Morazan,rd. Tegucigalpa-Juti-
calpa, ca. km 100-101, 27 Oct 1996 (6), Berg 1810 (EAP);
6 km SE of Catacamas,16 May 1987 (9 fl-fr), Ortega U.
361 (EAP, MO). SANTA BARBARA: Mt. Bella Vista, above
San Pedro Sula, 4 Oct 1969 (9 fl-fr), Barkleyet al. 39574
(GH); San Pedro Sula, Jul 1887 (9 fr + 6), Thieme194 in
herb.Donnell Smith5494 (G, GH, K, US). YORO:Subirana,
ca. 1000 m, Sep 1937 (st), Hagen et al. 1063 (NY). SWAN
ISLANDS: GreatSwan, 15-24 Aug 1971 (st), Proctor32563
(IJ).
NICARAGUA. BOACO:4 km NW of Camoapa, 1 Feb
1979 (d), Grijalvaet al. 16 (MO, U). CHINANDEGA: Chin-
andega, 13 Jan 1903 (9 fl-fr), Baker 2007 (A, G, GH, K,
MO, NY, P, US); Ameya, 19-21 Jun 1923 (6), Maxonetal.
7193 (US). CHONTALES: Rd. Juigalpa-PuertoDiaz, 11 Jul
1980 (6), Guzmdnet al. 378 (MO, U); CerroLa Bateca, 8
km S of Cuapa, 21 Sep 1983 (d), Nee 28268 (MO, NY);
3.6 km E of La Gateada,rd. to Nueva Guinea, 17 Jul 1977
(d), Stevens2856 (BG, BM, MO); ca. 2.8 km N of Cuapa,
3 Sep 1977 (6), Stevens3641 (BG, BM, MO); rd. Juigalpa-
La Libertad,ca. km 17.4, 23-25 Sep 1977 (9 fl-fr), Stevens
4145 (BG, MEXU, MO); 4.9 km E of Puerto Diaz, rd. to
Juigalpa,5 Jun 1980 (d), Stevens et al. 17423 (BG, MO).
ESTELi: Isiquf, 14 Jun 1984 (9 fl-fr), Laguna 400 (BG,
MO). GRANADA: Volcan Mombacho, 2 Jul 1975 (9 fl-fr),
Atwoodet al. 70 (MO, NY), 28 Oct 1976 (9 fr), Gomezs.n.
(MO);between SantaRosa & SantaMargarita,19 Mar 1982
(6), Moreno 15989 (BG, MO). LE6N: VolcanCasita,900-
1400 m, 14 Jun 1984 (9 fr), Moreno 24261 (BG, MO).
MADRIZ: SantaTeresa,CerroMeino, 3.5 km SE of Somoto,
23 Sep 1980 (6 fl-fr),Moreno2796 (BG, MO). MANAGUA:
146
El Charco,Rfo El Carmen,2 km W of Salamina,7 Sep 1981
(i), Moreno 10791 (MO); Hwy. 12, km 24, WSW of sum-
mit of Sierra de Managua,7 Jul 1977 (9 fl), Stevens 2703
(BG, BM, MO); ca. 2.3 km from Hwy. 12, rd. to Sierrade
Managua, 29 Jul 1977 (9 fl-fr), Stevens 2924 (BG, MO).
MASAYA:Laguna de Apoyo, 24 Oct 1980 (d), Moreno
3990 (MO); ParqueNacional VolcanMasaya, 13 Dec 1977
(9), Neill 3094 (MO, U), 21 May 1980 (9 fr), Stevens et
al. 17212 (MO). MATAGALPA:20 km NE of El Tuma, be-
tween Rio Wasaka& Rio Bulbul,9 Oct 1982 (9 fr), Grijalva
et al. 1355 (MO); rd. Muy Muy-Equipulas,ca. km 19, 30
Jul 1978 (c), Stevens 9556 (MO), (9 fl-fr), Stevens 9557
(MO). NUEVASEGOVIA:8 km NE of Jicaro, 2 Aug 1980
(d), Moreno 1707 (BG, MO); ca. 0.5-4 km N of San Fer-
nando, 13 Aug 1977 (9 fl-fr), Stevens3341 (BG, MO); ca.
1.6 km W of Murra,rd. to El Jicaro,27 Jun 1980 (c), Ste-
vens et al. 17599 (BG, MO). Rio RIVAS:Volcan Maderas,
above Balgiue,Isla de Ometepe, 15 Sep 1983 (9 fr), Nee et
al. 28120 (BG, MO, NY); nr. Moyogalpa, 16 Sep 1983 (9
fl-fr),Nee et al. 28174 (BG, MO, NY); Isla Ometepe,Volcan
Maderas,27 Apr 1984 (9 fl), Robleto 401 (BG, MO). ZE-
LAYA:Between El Muelle de los Bueyes & Villa Somoza,
6 Apr 1961 (9 fl), Bunting et al. 1052 (NY, S); PuertoCa-
bezas, 28 Feb 1971 (9 fr), Little 25025 (MO, US); Mun.
Siuna, El Hormiguero,30 Jun 1984 (6), Ort(z 1979 (MO);
CornIs., 4-6 Apr 1981 (9 fr), Stevens19926 (MEXU,MO).
COSTA RICA. ALAJUELA: Alajuela, Mar 1896 (6 +
9 fl), Donnell Smith6772 (BM, G, GH, K, NY, US); nr.Los
Chiles, Rio Frio, 7 Aug 1949 (9 fr), Holm et al. 950 (A, G,
MO, P). GUANACASTE: Pozo Azul, 14 May 1992 (9 fl),
Biesmeijer et al. 188 (U); 6 km SE of Santa Cruz, 24 Jul
1964 (9 fl-fr), Ferreyra 15939 (USM); Inter-American
Hwy., Rio Lagartos, s.d., Khan et al. 1155 (BM); Nicoya,
Mar 1900 (9 fl-fr), Tonduz13870 (BM, K, NY, P, US).
PUNTARENAS:Rio Terraba,nr. PalmarNorte, 27 Aug 1952
(d fl-fr), Allen 6543 (EAP); Monteverde,rd. Santa Elena-
San Luis-Lagarto, 10 Jul 1990 (9 fl-fr), Haber 9970 (MO);
nr.Cascajal,25 km ESE of Puntarenas,3 Jul 1949 (6), Holm
et al. 234 (G, MO, P, U); between Lagarto & Botuca, 24
Aug 1965 (9 fl), A. JimenezM. 3473 (NY, PMA, US); Rin-
c6n de Osa, 11 Feb 1974 (6), Liesner 2066 (BG, MO);
Santo Domingo de Golfo Dulce, Mar 1896 (6), Tonduz
10055 in herb. Donnell Smith 7164 (US). SAN JosE: Be-
tween SantaAna & Villa Col6n, 21 Jul 1971 (9 fr), Burger
et al. 7611 (BM, MO, U).
PANAMA. CHIRIQUi:Puerto Armuelles, 23 Jul 1967
(d), Hladik497 (P); 6 mi W of David, 29 Aug 1962 (9 fr),
Tyson929 (MO); 6 mi N of David, 25 Apr 1969 (9 fl-fr),
Lazor 2824 (MO); 10 mi N of David, 25 Apr 1969 (9 fl),
Lazor2825 (MO, PMA). COCLE:El Cope, 1 Sep 1977 (6),
Berg et al. 397 (BG), (9 fl-fr),Berg et al. 398 (BG); 0.5 km
N of El Cope, 10 Sep 1977 (6), Folsom 5226 (MO, PMA,
U). COL6N:Nr. RIo Piedras,rd. to PuertoBello, 16 Jul 1966
(6), Blum et al. 2515 (MO), (9 fr), Blumet al. 2529 (MO);
Coco Solo, 13 Aug 1967 (9 fr), Elias et al. 1606 (GH,
MICH, MO); Salud, 3 Aug 1971 (9), Lao et al. 203 (MO,
PMA); Col6n Hwy., bridge crossing Rio Chagres, 1 May
1969 (9 fl), Lazoret al. 3066 (MO); 3 km E of BuenaVista,
28 Oct 1973 (9 fr), Nee 7652 (MO); Fort Sherman,28 Jun
FLORA NEOTROPICA
1967 (9 fl-fr), Stimson5230 (NY); GatunRailway Station,
5 Mar 1966 (6), Tyson3524 (MO). DARIEN:Rfo Pirre,2-5
mi above El Real, 23 Jun 1962 (9 fr), Duke4904 (GH,MO);
23 km SE of Jaque,23 Jan 1981 ( 9 fr), Garwood1162 (MO,
PMA, U); nr.El Real, 22 May 1969 (9 fl), Lazoret al. 3439
(MO); Rio Tuquesa,mining camp CharcoChiva,5 Jul 1975
(9 fl-fr), Mori 6999 (BG, MO). Los SANTOS:12 mi S of
Macaracas,22 Feb 1966 (9 fl-fr), Blum et al. 2176 (MO).
PANAMA: BarroColoradoIs., 27 Apr 1968 (6), Croat5131
(MO), 25 Sep 1968 (9 fl), Croat 6452 (MO); rd. to Cerro
Azul, 4 mi from Inter-AmericanHwy., 26 Jul 1970 (d),
Croat 11496 (MO); SummitGardens,6 Aug 1970 (9 fl-fr),
Croat 11749 (MO); El Llano, 14-19 Oct 1962 (9 fl-fr),
Duke 5846 (US); San Miguelito, 9 Aug 1963 (9 fr), Duke
et al. 6645 (BG); San Jose Is., 9 Oct 1944 (d), Johnston78
(GH, US), 15 Oct 1944 (9 fl-fr), Johnston 154 (MO), (6),
Johnston 155 (GH, US), 12 Nov 1944 (9 fl-fr), Johnston
471 (GH); La Capitana,Chepo, 9 Jun 1971 (6), Lao 122
(MO); rd. to Cerro Campana, 1-3 mi from Pan-American
Hwy., 3 Aug 1983 (6), J. S. Miller et al. 953 (MO); 6 km
S of Arraijan,28 Aug 1973 (6), Nee 6659 (MO); 2 km W
of Gamboa,21 Oct 1973 (9 fl-fr), Nee 7533 (LL, MO, US);
rd. to Cerro Azul, 23 Apr 1972 (6), Tyson 6205 (PMA).
SAN BLAS: Nusagandi, 18 Aug 1989 (9 fl-fr), Fisher 26
(BG), (6), Fisher 27 (BG); rd. Suedi-Kariadi, Rio Ailin-
gandi, 10 Dec 1989 (9 fl-fr), Herrera et al. 644 (MO); El
Llano-Cartird., continentaldivide, 25 Aug 1984 (9 fl-fr),
Nevers3753 (BG, MO). VERAGUAS:Ca. 20 mi ESE of Gua-
bala, 3 Jul 1966 (9 fl-fr), Blumet al. 2443 (MO);Islas Con-
treras, Isla Brincaneo, 20 Jul 1984 (9 fl), Churchill5720
(BG, MO).
JAMAICA. Parishof St. Andrew,groundsof University
of the West Indies, 2 Jul 1966 (9 fl-fr),Andersonet al. 3000
(GH, LL, MICH,MO, US); Parishof St. Andrew,rd.Gordon
Town-Guava Ridge, 3 Jul 1966 (9 fl-fr), Anderson et al.
3001 (MICH, US), (6), Andersonet al. 3010 (GH, MICH,
US); Parish of Portland,rd. Section-Hardwaregap, 3 Jul
1966 (6), Andersonet al. 3020 (MICH,US); CockpitCoun-
try, Troy, 13-18 Sep 1906 (9 fl-fr), Brittonet al. 508 (NY);
nr.Nigril, 9-12 Mar 1908 (9 fl-fr),Brittonet al. 2048 (NY);
Nary Is., 26 Jul 1897 (9 fl), Fredholm3266 (NY); Parish
of St. Ann, ca. 1 km N of Hollymount,Mt. Diablo, 6-7 Aug
1965 (9 fr), Hespenheideet al. 874 (GH, LL, MICH,MO);
Parishof St. James,Irwin, 30 Apr 1952 (6), Johnsonet al.
22J181 (IJ); Parish of St. Elizabeth,Giddy Hall, Apr 1926
(9 fl), Maxwell s.n. (BM); Arntully,26 Sep 1927 (9 fl-fr),
Orcutt 3030 (G, GH, K, S, US); Parish of St. Mary, nr.
Hampstead,1954 (9 fl-fr), Proctor 8692 (IJ).
GRENADINES. Becquia, (d fl), D. Joseph (in H. H.
Smithet al.) 13100 (K).
BARBADOS. St. John, Bath Woods, Mar 1940 (6),
Gooding 331 (BM).
TRINIDAD. Roxborough-BloodyBay rd., 1 Mar 1981
(juv), Baksh et al. 321 (NY); Caura-Royalrd., 15 Feb 1979
(6), Berg 978 (U) & 979 (AAU, BG, U); Arima-Blanchis-
seuse rd., ca. km 6 km NNW of Arima, 15 Feb 1979 (9 fl-
fr), Berg 984 (BG, U); Aripo Savanna, 19 Aug 1951 (st),
Richardson683 (NY); Maraval, 1891-1892 (st), Warming
s.n. (C).
SYSTEMATICTREATMENT
COLOMBIA. ANTIOQUIA: Mun. Taraza,Vrda. Bana-
blanco, source of Rfo Puri, 24 Apr 1993 (d), Callejas et al.
10879 (HUA); Rfo Sucio, nr. Uramita,4 Oct 1961 (9 fl-fr),
Cuatrecasas et al. 26213 (COL, US), (d), Cuatrecasas et
al. 26214 (P, US); Rfo Cauca, between Puerto Valdivia &
Valdivia,550-1 100 m, 19 Feb 1942 (6), Cuatrecasaset al.
30096 (COL, F, US); 1 km W of Turbo,31 Mar 1962 (9 fl-
fr), Feddema2112 (MICH,US); Mun. San Luis, Rfo Claro,
25 Mar 1994 (i), Franco et al. 4583 (BG, HUA), (d),
Franco et al. 4584 (BG, HUA); Mun. Frontino,nr. Cgto.
Nutibara,1200 m, 5 Mar 1995 (c), Franco et al. 5561 (BG,
COL, HUA, LP, MO, NY, QCA); Mun. Anorf,Vrda.El Ro-
ble, 1400 m, 8 Mar 1995 (9 fl-fr), Franco et al. 5566 (BG,
COL, HUA, LP, MO, US); betweenVilla Artega& Rio Mu-
tata, 20 Nov 1948 (d), Johnson et al. 18C240 (MEDEL);
Mun. Jeric6, 1200 m, 5 Jul 1983 (c), Ortiz S. et al. 110
(MEDEL);Vrda. La Rubiela,rd. Segovia-Antioquia, 19 Jul
1979 (9 fl-fr), Renterfaet al. 1688 (COL, NY); Mun. Fre-
donia, Cgto. Marsella, 14 Apr 1992 (9 fl-fr), D. SdnchezS.
1760 (MEDEL); Mun. Zaragoza, Cgto. de Providencia,9
Feb 1971 (6), Soejartoet al. 2662 (COL,GH);nr.Medellfn,
10 Sep 1927 (9 fl-fr), Toro586 (NY); Mun. Caucasia,rd.
to Nechf, 17 km from rd. Caucasia-PlanetaRica, 22 Mar
1987 (9 fl-fr), Zarucchi et al. 4915 (BG, COL, MO). AR-
CHIPELAGODE SAN ANDRES Y PROVIDENCIA:Isla de Prov-
idencia, Mar-Jul 1977 (6), Freeman 35 (MEDEL).
ATLANTICO: Nr. Barranquilla,Aug 1928 (9 fl-fr), Hno.
Elias 578 (US), Jul 1934 (9 fr), Hno. Elias 1230 (F, MICH,
US). BOLiVAR: Mun. Turbaco,8 Jun 1982 (6), CuadrosV
1372 (MO), (9 fl), CuadrosV 1383 (MO); nr. Turbaco,29
Jul 1953 ( 9 fr), Dugand et al. 3357 (COL,F); nr.SanMartfn
de Loba, Apr-May 1916 (6), Curran65 (GH, S, US); Mun.
San Juan Nepomuceno, Loma de Los Colorados,ca. 2 km
S of San Juan, 7 May 1987 (st), Gentry et al. 57488 (BG,
MO); Torrercilla,nr.Turbaco,7-19 Nov 1926 (st), Killip et
al. 14243 (GH, NY, US); Mun. Cartagena,7 km SW of
Arroyo Grande,31 Jul 1985 (9 fl-fr), Zarucchiet al. 3884
(BG, MO, NY). CALDAS: Nr. La Dorada,1-2 Feb 1946 (6),
Duque-Jaramillo2590 (NY); QuebradaYeguas,20 km N of
Honda, 7 Mar 1977 (9 fl-fr), Gentry et al. 18194 (COL,
MO); Pan-AmericanHwy., nr. Marmatoturnoff,NE of Rfo
Sucio, 27 Mar 1980 (6), Gentry28801 (AAU, MO, U, US).
CESAR:Rd. La Paz-Manaure,26 Apr 1987 (6), CuadrosV
et al. 3402 (BG); Sierrade Perija,E of Manaure,El Podrido,
1550-1600 m, 15 Nov 1959 (6), Cuatrecasas25356 (US);
rd. San Martfn-Ocaina,Quebradadel Gobernador,27 Sep
1969 (d), Cuatrecasaset al. 27965 (COL, NY, US), (9 fl-
fr), Cuatrecasas et al. 27968 (COL, NY, US); 5 km W of
Manaure, 13 Jan 1988 (st), Gentry et al. 60723 (MO).
CHOcO:Cerrosdel Cuchillo, Rosa Marfa, 16 Oct 1987 (9
fl), Cdrdenas679 (JAUM,MO); Mun. Riosucio, Cerrosdel
Cuchillo, rd. Cuchillo Negro-Punta de las Flores, 18 May
1988 (9 fl-fr), Cardenas1934 (JAUM,MO); Mun. Pizarro,
rd. Pie de Pep6-Berrecul,km 21-24, (9 fl-fr), Espina 1920
(COL, MO); nr. Bahfa Solano, 4 Apr 1990 (9 fl-fr), Franco
et al. 3001 (COL); Rfo San Juan, above Istmina, 14 Aug
1976 (6), Gentryet al. 17664 (AAU, BG, MO, US); Mun.
Acandf, Cgto. de Ungufa, Rfo Cuti, 27 Jul 1957 (6),
Romero-Castanteda6449 (COL, MO, NY); Rio Jurad6,4
147
Oct 1940 (6), SneidernA.246 (MICH,NY, S, UA); Parque
Nacional NaturalLos Katios, Sautata,3 May 1983 (9 fl),
Zuluaga R. 637 (COL). C6RDOBA: PlanetaRica, rd. Mon-
teria-Planeta,El Charco,24 Oct 1987 (9 fl-fr), Leguizamo
493 (MO), (6), Leguizamo494 (MO). CUNDINAMARCA: 9
km E of Melgar, on rd to Fusagasuga, 8 Jul 1972 (9 fl),
Barclay et al. 3557 (COL,US); Mun. Melgar,Rio Sumapaz,
nr. La Nariz del Diablo, 12 Mar 1993 (9 fl), Franco et al.
4346 (COL,MO), (d), Francoet al. 4347 (BG, COL).GUA-
JIRA: Between Cuestecita & Carraipia,Rio Cesar, 30 Nov
1959 (d), Cuatrecasaset al. 25522 (COL), (9 fl-fr), Cua-
trecasas et al. 25535 (COL, US); 15 km S of Mingueo, trail
to Pueblo Viejo, 21 Aug 1986 (st), Gentry et al. 55425
(MO); SerraniaLa Macuira,Cerro Manzano, 5 Mar 1963
(juv), Saravia T 2419 (COL,US). HUILA: Between Gigante
& Potrerillo, 1 Nov 1926 (9), Juzepczuk6710 (LE); nr.
Neiva, 7 Apr 1944 (9 fl-fr), Little et al. 7595 (COL, F, P);
between Neiva & Campoalegre,19 Mar 1940 (9 fr), Perez
Arbelaez et al. 8301 (COL, F). MAGDALENA: Rd. to Rio-
hacha, km 62, 17 Apr 1982 (9 fl), CuadrosV 1352 (COL,
MO); SierraNevada de Santa Marta,Rio Donachuf,below
Donachui, 1230-1350 m, 24 Sep 1959 (9 fl-fr),Cuatrecasas
et al. 24419 (COL, US); Sierra de Perija, Manaure,Rio
Manaure,4 Nov 1959 (d), Cuatrecasaset al. 25020 (COL);
Rio Buritaca,Alto de Mira, 1100 m, 20 Jul 1989 (6>),Mad-
riniinet al. 406 (MO); Santa Marta,6 Apr 1899 (d), H. H.
Smith2105 (BM, COL, E, G, GH, K, LE, LL, MO, NY, P,
S, U, US); 13 km SE of SantaMarta,8 May 1977 (6), White
et al. 438 (COL, HUA, MO, NY). NORTE DE SANTANDER:
Between Gramalote& Peralonso,24 Jul 1940 (9 fr), Cua-
trecasas 10140 (COL, F); Rio Pamplonita,between Cdcuta
& Pamplona,25 Jul 1940 (9 fr), Cuatrecasas10178 (COL,
F). SANTANDER: 5 km S of Bucaramanga,14 Jul 1968 (9
fl-fr), Barkley38C212 (GH); Mun. Piedecuesta,Cgto. Pes-
cadero, Vrda. Mesetas, Rio Umpala, 12 Jul 1996 (9 fl-fr),
Cadena et al. 3008 (COL);RIo Chicamocha,between Cap-
itanejo & Enciso, 1200-1300 m, 18 Jul 1940 (9 fr), Cua-
trecasas 9843 (COL, F); Rio Pienta, between El Limite &
Encino, 2000 m, 4 Aug 1940 (9 fl), Cuatrecasas 10437
(COL, F), (6), Cuatrecasas 10438 (COL, F); Mun. Pie de
Cuesta,La Mesa de los Santos, 1460-1700 m, 16 May 1969
(6), Garcia-Barriga 19686 (COL, NY); between Puerto
Wilches & Puerto Santos, 29 Nov 1926 (6), Killip et al.
14892 (A, GH, NY, US); Mesa de los Santos, 1000-1500
m, 11-15 Dec 1926 (9 fl-fr), Killip et al. 15010 (A, GH,
NY, US); 12 mi SE of BarrancaBermeja, 15 Sep 1954 (9
fl-fr), Romero-Castaneda4847 (COL, US). TOLIMA: Be-
tween Mariquita& Guayabal,7 May 1940 (9 fl-fr), Cua-
trecasas 9410 (COL, F); Mun. Mariquita,rd. Mariquita-
Honda, 17 May 1990 (9 fl), Franco et al. 2968 (COL,MO);
rd. Espinal-Ibague,nr. Rio Coello, 6 Feb 1955 (6), Franco
et al. 4646 (BG, COL); Mun. Piedras, rd. Piedras-Doima,
km 5, 7 Feb 1995 (9 fl-fr), Franco et al. 4649 (COL); 11
km NE of Melgar,Rfo Sumapaz,8 Jan 1974 (9 fl-fr),Gentry
et al. 8958 (COL, GH, MO); between Colache & Coyaima,
19 Oct 1926 (6), Juzepczuk6928 (LE). VALLE:Buenaven-
tura, 16 Mar 1994 (9 fl-fr), Franco et al. 4536 (BG); rd. to
Bajo Calima, Bahia Malaga, 16 Mar 1994 (6), Franco et
al. 4542 (BG, TULV), (6), Franco et al. 4543 (BG, TULV).
148
VENEZUELA. AMAZONAS: Depto. Atabapo, Rfo
Ocamo, nr. Raudal Arata, Jan 1990 (d), Ferndndez6533
(BG). ANZOATEGUI: Cantaura, 19 Apr 1950 (d), F D.
Smith 141 (US). ARAGUA: ParqueNacional Henri Pittier,
Rancho Grande, 11 Aug 1953 (Y fr), Little 15451 (NY,
VEN), (6), Little 15452 (NY, VEN); nr. Maracay,2 Apr
1926 (6), Pittier 12143 (G, NY, US); rd. San Casimiro-
Camatagua,Aug 1966 (6), Veldsquez86 (US, VEN); Ran-
cho Grande,Aug 1966 (Y fl-fr), Veldsquez94 (US), (6),
Vela'squez95 (US, VEN); Guamites, 12 May 1938 (Y fr),
Ll. Williams10074 (F, VEN). BARINAS: Mun. Pedreza,40
km SE of CiudadBolivia, 24 Apr 1953 (Y fl), Little 15119
(VEN), 25 Apr 1953 (d), Little 15120 (VEN); Mun. Cruz
Paredes, Dtto. Obispos, 20 km NW of Barrancas,26 Mar
1932 (6), Marcano Berti et al. 2963 (VEN); rd. Merida-
Barinas, between La Yuca & Barinitas,21 May 1980 (6),
Marcano Berti et al. 109-980 (G, U, VEN); rd. M6rida-
Barinas,between La Yuca & Barinitas, 11 May 1980 (Y fl-
fr), Marcano Berti et al. 113-980 (G, U, US); rd. Barinas-
San Crist6bal, Canagua, 21 May 1980 (Y fl-fr), Marcano
Berti et al. 114-980 (G, U, US, VEN); rd. Barinas-SanCris-
tobal, between Socop6 & Capitanejo,22 May 1988 (6),
MarcanoBerti 115-980 (G, U, VEN); rd.Barinas-Acarigua,
Dec 1966 (9 fl-fr), Veldsquez102 (US, VEN). BOLiVAR:
Dtto. Cedefno,nr. Corozal, 13 Apr 1986 (6), Boom et al.
6493 (NY); Dtto. Roscio, San Martin de Turumban,Aug
1979 (9 fl-fr), Delascio et al. 7845 (VEN); Mun. Raul
Leoni, 86 km E of San Francisco, Jun 1989 (9 fl-fr), Del-
gado 241 (BG, MO, NY, VEN); 20-35 km SW of El Man-
teco, rd. to San Pedro de las Dos Bocas, 1-3 Aug 1978 (9
fl-fr), Liesner et al. 5815 (MO, U, VEN); Reserva Forestal
Imataca, nr. confluence of Rio Botanamo & Rio Corumo,
19-20 Jan 1983 (9 fl), Stergios et al. 5372 (US); Rio Asa,
above RaudalCotua, S of La Paragua,1 Aug 1960 (9 fl-fr),
Steyermark86723 (NY, US); Rio Caura,Salto Para, 15 Jan
1977 (6), Steyermarket al. 113020 (F, K, MO, U, VEN);
Rio Uairen, Gran Sabana, Santa Elena, Feb 1946 (6), Ta-
mayo 2981 (F, G, S, VEN); Reserva Forestal del Imataca,
Aug 1967 (9 fl-fr), Veldsquez252 (US); El Palmar,25 Apr
1940 (9 fl-fr), Ll. Williams12893 (F, K, S, US, VEN). Co-
JEDES: Dtto. Giradot,Hato Piniero,rd. to CharcoAzul, 6 Jul
1990 (9 fl-fr), Aristeguietaet al. 1482 (VEN); El Mapuey,
26 Sep 1975 (9 fr), Delascio et al. 3446 (VEN). COLON:
San Crist6bal,5 Jan-22 Feb 1923 (d), Broadway259 (GH,
NY, US). DELTA AMACURO: Rd. Tucupita-LosGiiires, 17
Apr 1973 (9), Agostini et al. 1624 (U); LowerRio Orinoco,
Manoa, May 1896 (6), Rusby et al. 99 (A, BM, F, G, K,
MO, NY, US, VEN); Rio Cuyubini,between La Paloma &
Moron, 20 Nov 1960 (d), Steyermark87693 (VEN). Dis-
TRITOFEDERAL:Rd. Caracas-LaGuaira,22 Mar 1922 (d),
Pittier 10266 (VEN); Curucuti,27 Jul 1922 (9 fl), Pittier
10418 (G, GH, US, VEN); Caracas,BotanicalGarden,(d),
Veldsquez80 (US, VEN), (9 fr), Veldsquez81 (US, VEN).
FALCON: Dtto. Zamora,CerroMampostal,4 Aug 1977 (9
fl-fr), A. Gonzdlez 1012 (MO, VEN); Dtto. Acosta, Mun.
Jacura,Cerro La Mina, 14 Nov 1979 (9 fl-fr), Marcano
Berti 456-979 (U, VEN). GUARICO: Rd. La Encrucijada-
Calabozo,Jul 1954 ( 9 fl-fr),Aristeguietaet al. 2308 (VEN);
FLORA NEOTROPICA
rd. El Sombrero-Calabozo,8 Aug 1966 (Y fl-fr), Veldsquez
87 (US, VEN), (6), Veldsquez88 (US, VEN). MERIDA: Be-
tween Ejido & Mesa de los Indios, 1200 m, 23 Apr 1953
(6), Bernardi416 (NY); Dtto. Justo Briceilo, between La
Panamaricana& Las Virtudes, 11 May 1978 (6), Bunting
et al. 6341 (U); rd. Merida-Tovar,between Ejido & Lagun-
illas, 850 m, 10 Apr 1969 (6), Oberwinkleret al. 15607
(VEN); nr. Merida, Feb 1968 (6), Veldsquez263 (US,
VEN). MIRANDA: Dtto. Paez, rd. San Juan-Montevideo,7
Sep 1977 (Y fr), A. Gonzales et al. 1405 (MO, VEN); Alto
de Guayabo,rd. Caracas-Cua,km 17, 17 Apr 1924 (Y fl),
Pittier 11511 (A, G, K, NY, P, US, VEN); ParqueNacional
de Guatopo,SantaCruz,between SantaTeresa& Altagracia
de Orituco, 23 Nov 1961 (Y fl-fr, Steyermark89976 (NY,
US, VEN); El Guapo,Aug 1965 (? fl-fr), Veldsquez78 (US,
VEN), (6), Veldsquez79 (US, VEN); ParqueNacional Gua-
topo, Oct 1966 (6), Veldsquez97 (US, VEN); Panaquire,
Jul 1966 (Y fl-fr), Velasquez247 (US, VEN). MONAGAS:
Rd. Jusepin-Caicarade Maturfn,22 Aug 1970 (6), Ariste-
guieta et al. 7531 (VEN); Reserva Forestalde Guarapiche,
22 Feb 1978 (Y fl), Castillo 711 (NY, VEN); Dtto. Caripe,
Cerro de Los Rastrojos,QuebradaLas Majaguas, 13 May
1982 (Y fr), 0. Huber6348 (NY, U); 2 km SSW of Jusepin,
5 Mar 1967 (d), Pursell et al. 8241 (VEN); Caicaro, 12 Jun
1940 ( fr), Ll. Williams13300 (F, US). NUEVAESPARTA:
Isla de Margarita,Jul 1984 (Y fr), Delascio et al. 12225
(MO, VEN); Isla Margarita,CerroCopey, 25 Apr 1983 (Y
fl-fr),Sugden1182 (K, MO, VEN), 2 May 1983 (d), Sugden
1204 (K, MO). PORTUGUESA: Dtto. Guanare,grounds of
UNELLEZ, 21 Jan 1982 (9 fr), Aymardet al. 811 (MO);
Dtto. Sucre, Caserfo Villa Rosa, 20 km E of Buscucuy,
1200-1500 m, 16 Jun 1985 (9 fl), Aymardet al. 3655 (US);
Dtto. Ospino, El Corozo, Rfo Guache, 22 May 1983 (d),
Delgado 20 (MYF). SUCRE:Penfnsulade Paria, Cerro Pa-
tao, 6 Sep 1984 (d) Millikenet al. 159 (MO); Penfnsulade
Paria,CerroPatao,NE of Giuiria,23 Jul 1962 (9 fl-fr),Stey-
ermark et al. 91272 (US, VEN); rd. Rfo Caribe-Tunapui,
Feb 1967 (9 fl-fr), Veldsquez98 (US), VEN). TACHIRA: 7
km W of Rubio, 18 Mar 1981 (d), Liesneret al. 10737 (MO,
U, VEN); Rfo Frio, between jct. of Rio Quinimari& Rfo
Frio, between (village) Rfo Negro & La Laguna,30 Jul 1979
(d), Steyermarket al. 119082 (MO, U, VEN); Quebrada
Colorada,between San Juande Col6n & Las Cruces, at La
Pocina, 14 Nov 1979 (6), Steyermarket al. 120308 (MO,
NY, U). YARACUY: Rfo Yurubf,13 Sep 1974 (6), Delascio
et al. 2578 (VEN); Salom, 18 Feb 1954 (9 fl), Little 16215
(VEN). ZULIA:Dtto. Col6n, between Casigua El Cubo &
km 8 on rd to Palmira,28 Apr 1979 (9 fl), Bunting et al.
7323 (U); Dtto. Bolfvar, between Quir6s-El Pensado &
Cerro Socope, 5-8 Aug 1980 (9 fl-fr), Bunting 9853 (NY,
U, VEN); Dtto. Perija, nr. Estaci6n Hidrol6gicaAricuaisa-
Pie de Monte, 1-3 1982 (9 fl-fr),Buntinget al. 11458 (NY,
U); Mun. Santa Barbara,Dtto. Col6n, 17 Feb 1981 (9 fl-
fr), MarcanoBerti et al. 146-981 (G, U, VEN); Rio del Pal-
mar,San Martin, 15 Oct 1922 (9 fr), Pittier 10518 (G, NY,
US); Dtto. Mara,2.5 km W of CorpozuliaCampamentoCur-
ichuano, 3 Jun 1980 (9 fl-fr), Steyermarket al. 123221
(MO, NY, U, VEN); Sierrade Perija,rd. La Matera-ElPar-
SYSTEMATICTREATMENT
amito, San Jose de Los Altos, 8 Jul 1975 (9 fr), Zambrano
222 (VEN).
GUYANA. DemeraraR., 7 Apr 1987 (d), Boom 7190
(BG, NY, U); nr. Bartica,30 Jan 1951 (6), Fanshawe3033
= FD 6363 (K, NY, U); EssequiboIls-W. DemeraraRegion,
NaamryckCanal,ca. 3.5 km SW of Parika, 14 Apr 1989 (9
fr), Gillespie 995 (BG, U); U. Takatu-U.EssequiboRegion,
ca. 15 km S of Sand Creek Vill., 21 Jun 1989 (9 fl-fr),
Gillespie et al. 1729 (BG, NY); Potaro-Siparuniregion,Pak-
araimaMtns., E slope of Malakwalai-Tipu,1100 m, 10 Jul
1994 (d), Henkel et al. 5542 (BG); KanukuMtns., Mai-
paima, 17 Nov 1987 (9 fl-fr),Jansen-Jacobset al. 971 (BG,
K, MO, NY, U); RupununiDistr., KusadMtns., 27 Sep 1992
(9 fl-fr), Jansen-Jacobset al. 2658 (BG, MO, NY, U); Ber-
bice R., S of New Dageraad,6 Oct 1981 (9 fl), Maas et al.
5571 (F, K, MO, NY, U); KanukuMtns., Moku-mokuCr.
31 Mar-16 Apr 1938 (9 fl-fr), A. C. Smith 3426 (A, F, G,
K, MO, NY, P, S, U, US).
SURINAME. Wilhelmina Mtns., Kayser Airstrip, 4.5
km above confluence of Zuid R. & Lucie R., 29 Sep 1963
(9 fl), Irwin et al. 57681 (US); Waneweg, 30 km S of
Paramaribo,9 Dec 1960 (d), Krameret al. 2317 (NY, U);
Coppename R., Nature Reserve Voltzberg, 25 Feb 1977
(9 fl-fr), Lindeman& Mennegaet al. 150 (K, MG, NY, RB,
U); Sipaliwini savanna,nr. Brazilian frontier,25 Sep 1970
(st), Oldenburgeret al. 1362 (U); Zanderij,3 Jul 1916 (d),
Samuels 506 (F, GH, K, NY, P); Afobaka rd. 10 km
S of Paranam,1 May 1970 (6), Teunissen(LBB) 12739 (U).
BRAZIL. PARA:Parque Nacional do Tumucumaque,
Rio Parudo Oeste, Missao Tiriyo, 16 Feb 1970 (d), Cav-
alcante 2403 (NY, U, US), 20 Feb 1970 (9 fl-fr),Cavalcante
2475 (NY, U, US). RORAIMA: Ilha de Maraca, 12-26 Jul
1986 (st), Campbell et al. 15832 (BG); Ilha de Maraca,
SEMA Ecological Reserve, 7 Jul 1987 (6), Milliken 408
(BG, E, K).
In Jamaica,some collections have trichilia,others
have vestigial ones, and in severalcollections they are
absent.Materialwithouttrichiliaor vestigial ones can
be distinguishedfrom Cecropiaschreberianavar.an-
tillarum (found in other Caribbeanislands) by the
scarce and white pith in the intemodes and the ab-
sence of relativelylong and soft hairs on the veins of
the lamina beneath. Trichilia are present in the col-
lection from the Swan Islands and in all collections
made elsewhere. Several collections, e.g., Jansen-
Jacobs et al. 971 and 1260 from Guyana, and Fer-
ndndez 6533 from Venezuela(Amazonas),have long
white hairs in the trichilia;they resemble C. metensis
in this respect,but lack the dense arachnoidindumen-
tum on the petiole.
The species has been introducedinto severalcoun-
tries in Africa, Asia, and the Pacific, and also to the
BermudaIslands (Brownet al. 1683, NY). It has be-
come naturalizedin several countriesin West Africa,
at least in Cameroun (R. Letouzey 12474 and 14828,
149
P; S. A. Thompsonet al. 1398, MO), Ghana (H. H.
Schmidtet al. 2032, MO), Ivory Coast (E. Merklen
s.n., US; Leeuwenberg4504, MO; Miege & AkeAssi
3960, G, P; Tere HGT 2155, G), and Senegal
(Berhaut 984, P); also French Polynesia (Makatea,
Raiatea, Rapa, Tahaa,and Tahiti, accordingto Flor-
ence [1997: 24-26, fig. 1]), for the firsttime collected
in 1927; and Malaysia (Mat Asri FRI 25549, A; see
also Putz & Holbrook,1988). In Cameroun,it is com-
peting with Musanga cecropioides (McKey, 1988a).
The species has been or is still present in some bo-
tanicalgardens:China(Guangzhou,accordingto Liao
[1991, 1992], and Xishuangbanna),Gabon (Libre-
ville, 1900, C. Chalot 48, P), Singapore(1930, C. X.
Furtados.n., A), Indonesia(Bogor, 1952, Anonymous
s.n., A), and Taiwan (Shiaping Tropical Botanical
Garden,Chishan,accordingto Liao [1991, 1995]; in-
troduced from El Salvador and with trichilia, see
Liao, 1991: 131, fig. 5, photo. 1; and Liao, 1995: 134,
fig. 5, photo. 1). For the material in the Botanical
Gardenof Bogor, Brazil was indicatedas the prove-
nance. Accordingto Putz & Holbrook(1988) the ma-
terial in Malaysia is introducedfrom Indonesia(Bo-
gor). If the annotationon the Bogor material about
the provenanceis correct,the materialshould belong
to Cecropiapachystachya,but the collections exam-
ined do not show some of the characteristicfeatures
of that species. Moreover,C. pachystachyahas never
been encounteredwithout trichilia. Thus, the infor-
mation about origin or the identity of this materialis
dubious. In materialfrom Africa trichilia are absent
or poorly developed. In French Polynesia they may
often be lacking. In Malaysianmaterialexaminedby
Putz & Holbrook (1988), 80% of the specimens had
fully (fused) developedtrichilia,8%poorly (separate)
developed ones, and 12%none. The intriguingques-
tion is whether all this materialhas as its origin Ja-
maica, the only areain which C. peltata is found with
and withouttrichilia,or whetherloss of the ability to
form (well-developed) trichilia was developed after
introductionelsewhere (in two species?). The latter
case would confirm the hypothesis of dissolution of
the mutualismbetween Cecropia and Azteca due to
the absence of the latterpartnerand manifest in the
absence of trichilia (Janzen,1973).
Vernacular names. Mexico: kooch le' or
xkoochle (Maya, Quintana Roo); chancarro(Vera-
cruz). Belize: cho-otz (Maya);guarumo,trumpettree.
Panama:trampy(Jamaican);nig-la (Kuna,San Blas).
Jamaica:snake wood, trumpet.Colombia: guarumo
or yarumo blanco. Venezuela: guargiuero(Barinas);
gueremo, orumo (Zulia). Guayana:congo pump.
150
42. Cecropia pittieri Robinson,in Stewart,Proc. Ca-
lif. Acad. Sci., ser. 4, 1: 389. 1912; Burger,Fiel-
diana, Bot. 40: 127. 1977. Type. Costa Rica. Co-
cos Is., Jan 1902 (d), Pittier 16237 (holotype:
GH).
Tree, to 20 m tall. Leafy twigs 2.5-4 cm thick,
(sub)hispudulouswith straighthairs. Lamina charta-
ceous, ca. 30 X 30 cm to 60 X 60 cm, the segments
10-11, the incisions down to 2/10-3/10; apices
rounded to subacuminate;upper surface scabrous,
hispidulous;lower surfacehirtellousto subhirsute(to
subtomentose)on the veins, with arachnoidindumen-
tum in the areoles and on the reticulum;lateralveins
in the free part of the midsegment ca. 10-12 pairs,
submarginally to marginally loop-connected, often
branched;petiole 20-50 cm long, with arachnoidin-
dumentum,at base and apex hirtellousto subhirsute;
trichilia lacking; stipules 8-17 cm long, color un-
known, hirtellous to subhirsuteand with arachnoid
indumentumoutside, sericeous inside. Staminatein-
florescences in pairs (?), the peduncle erect and the
spikes pendulous (?); peduncle 8-9 cm long, puber-
ulous to hirtellous and with arachnoidindumentum;
spathe ca. 15 cm long, color unknown,hirtellous to
strigose to hirsute outside, sparsely hairy inside;
spikes ca. 10-12, ca. 10-12 X ca. 0.3 cm, with stipes
0.6-0.8 cm long and sparselyhairy;rachishairy.Sta-
minate flowers: perianth tubular, 1-1.5 mm long,
sparsely minutely puberulous below the apex, the
apex almost plane; anthers 0.5-0.6 mm long, de-
tached at anthesis (?). Pistillate inflorescences in
pairs, pendulous; peduncle 5-10 cm long, sparsely
puberulousto hirtellous and also with arachnoidin-
dumentum;spathe not seen; spikes 4, to 21 X 1-1.2
cm in fruit,sessile; rachisglabrous.Pistillateflowers:
perianthca. 2 mm long, with arachnoidindumentum
below the apex outside, the apex convex, muriculate;
stigma comose. Fruit ellipsoid to ovoid, ca. 2 mm
long, slightly tuberculate,darkbrown.
Distribution (Fig. 8.5). Endemicof Cocos Island.
Specimensexamined.COSTARICA.Cocos ISLAND:
Bahfade Chatham,13 Aug 1973(Y fr), Carrasquilla 362
(MO,NY), 13Aug 1973(9 fl-fr),Dressler4460(NY,PMA,
311 (NY),13Apr1965
US), 8 Mar1964(9 fl-fr),Fournier
3184 (COL,MO, U); Bahfa de Wafer,12
(9 fl-fr), Jimenez
Oct 1994(9 fl-fr),Queseda1064(BG),Bahiade Chatham,
1905-1906 (juv), Stewart291 (GH).
This species is interestingbecause of its limited
distributionand the absence of trichilia. It is not in-
habited by ants and is probablyrelated to Cecropia
longipes.
FLORANEOTROPICA
43. Cecropia plicata Cuatrecasas,RevistaAcad. Col-
omb. Ci. Exact. 6(22/23): 280, t. 1. 1945. Type.
Colombia. Valle: Rfo Sanquinini, La Laguna,
1250-1400 m, 10-20 Dec 1943 (9), Cuatrecasas
15469 (holotype:COL; isotypes: F, US).
CecropiadiguensisCuatrecasas,
RevistaAcad.Colomb.
Ci. Exact.9(36/37):332. 1956. Type.Colombia.
Valle:RioSanJuan,belowQueremal, 1300-1500m,
24 Mar1947(d), Cuatrecasas23967(holotype:
US;
isotype:F).
Tree,to 20(-30) m tall. Leafy twigs 2-4 cm thick,
dark green to brown or white sparsely puberulous,
also with ? dense pluricellularhairs and ? dense
arachnoidindumentum.Laminacoriaceousand often
? plicate, ca. 30 X 30 cm to 75 X 75 cm, the seg-
ments 8-10(-l 1), the free partsof the upperones el-
liptic to oblong to subobovate,the incisions down to
6/10-8/1 0; apices rounded(or acuminate);uppersur-
face smooth, sparsely strigillose (on the main veins)
or subglabrous;lower surface sparsely minutely pu-
berulousand with ? dense brownpluricellularhairs,
with arachnoid indumentum lacking or sparse (in
minute patches) in the small areoles (surroundedby
thick veinlets); lateral veins in the free part of the
midsegment 15-24 pairs, submarginally loop-
connected, unbranchedor some branched;petiole ca.
25-70 cm long, sparsely minutely puberulous and
with ? dense brown pluricellularhairs and rather
sparsearachnoidindumentum;trichiliafused (or sep-
arate),the brown indumentumintermixedwith short
whitishhairs;stipules 12-33 cm long, darkred-brown
to purplish, puberulous to hirtellous and with ?
dense brownpluricellulartrichomesoutside,glabrous
or (sub)sericeous inside. Staminate inflorescences
solitary or in in pairs, erect; peduncle 5-13 cm long,
darkred to red-brown,(sparsely)puberulousor to hir-
tellous, also with brownpluricellularhairsand arach-
noid indumentum;spathe 12-21 cm long, darkred to
red-brown, ? sparselypuberulousto strigose to hir-
tellous and with dense brown pluricellular hairs
outside, glabrousor in the upperpart (sub)sericeous
inside; spikes 5-9, 11-18 X 0.7-1.5 cm, with stipes
0.5-2.5 cm long, red, and appressed-puberulousor
glabrous; rachis hairy. Staminate flowers short-
pedicellate;perianthtubular,2-2.5 mm long, puber-
ulous (or also with arachnoidindumentum)below the
apex and on the pedicel, the apex convex to plane,
hispidulous; filaments swollen; anthers0.6-0.8 mm
long, appendiculate,detached at anthesis, reattached
to the marginsof the apertureby the appendagesafter
anthesis.Pistillate inflorescencesin pairs or solitary,
erect, deflexed in fruit; peduncle 4-8 cm long, dark
red or partly whitish, puberulousto subhirsute,also
SYSTEMATICTREATMENT
with brown pluricellularhairs and arachnoidindu-
mentum; spathe 9-11 cm long, the color and indu-
mentum as in the staminateinflorescence;spikes 2-
3(4), 4.5-11 X 0.6-1.2 cm, to 25 X 3 cm in fruit,
(sub)sessile; rachis hairy. Pistillate flowers: perianth
2-3 mm long, with arachnoidindumentumbelow the
apex outside, absent inside, the apex convex, muri-
culate to hispidulous; style ratherlong, muriculate;
stigma comose. Fruit subovoid, ca. 3 mm long,
smooth, pale brown.
Distribution (Fig. 11.1). Colombia, from Antio-
quia to Valle, in (sub)montaneforest, at 1000-2300
M.
Representativespecimensexamined.COLOMBIA.
ANTIOQUIA:DonMatfas,
2100m,27 Mar1994(d), Franco
et al. 4592 (BG,HUA);nr.Yarumal, 2050m, 27 Mar1994
(Y fl-fr),Francoet al. 4593 (BG, HUA);Mun. Urrao,
ParqueNacionalNaturalLasOrquideas, Quebrada LaAgu-
delo, 1300-1380m, 6 Jul 1992 (st), Ram[rezet al. 4151,
4184, 4242 (JAUM).CHocO:Mun.San Josedel Palmar,
Vrda.El Sinai, 1650 m, 19 Mar 1994 (d), Francoet al.
4563 (BG,HUA);Mun.SanJosedel Palmar,ca. 1 kmE of
SanJosedelPalmar,1250m, 19Mar1994(? fl-fr),Franco
et al. 4570 (BG,HUA);Mun.Frontino,Cgto.Nutibara, rd.
Nutibara-La Blanquita, Murriregion,1900m, 4 Mar1995
(Y fl-fr), Franco et al. 5549 (BG,COL,HUA),1500m, 4
Mar 1995 (d), Franco et al. 5554 (BG, COL, HUA). RIs-
ARALDA: Mun. PuebloRico, rd. to CerroMonctezuma,
1350m, 15Mar1986(Y fl-fr),Bernalet al. 997(COL,MO,
NY). VALLE: Mun.Darien,LagoCalima,1400m, 15 1994
( fl-fr),Franco et al. 4529 (BG,TULV).
This species appearsto be very closely relatedto
Cecropia maxima,from which it differs in generalin
the smaller dimensions of severalparts (such as lam-
ina, stipules, spikes of pistillate inflorescences) and
the denser indumentumof brown pluricellularhairs.
Cecropiaplicata can be distinguishedfrom C. maxi-
ma by the smaller numberof segments of the lamina
(8-10 vs. 10-15) and by the arachnoidindumentum
on the lamina beneath,this being confined to the ar-
eoles and not present on the main veins. Staminate
specimens may be consistently distinct in the pres-
ence/length of the stipes of the spikes. The natureof
the differences are such that they could justify sepa-
rationonly at the subspecificlevel. The two taxa ap-
pear to be allopatric.Cecropiaplicata also resembles
C. insignis, particularlyspecimenswith 8 laminaseg-
ments. Such specimensdifferfrom those of C. plicata
in the presence of arachnoid indumentum on the
smaller veins and at least initially also on the main
veins on the laminabeneath.Pistillate specimens can
be distinguishedby the difference in the numberof
spikes: usually 2-3 in C. plicata and mostly 5 in C.
151
insignis. Overlapof the elevationalranges of the two
species has not yet been found in Colombia.Ants are
often present.
44. Cecropia polystachya Trecul, Ann. Sci. Nat.
Bot., Ser. 3. 8: 80. 1847 (Aug). Type. Peru.With-
out locality, (5), Ruiz & Pavon s.n. (holotype:P;
isotypes: B, G). Fig. 39
Cecropiaflagellifera Trecul,Ann, Sci. Nat. Bot., Ser. 3,
8: 81. 1847 (Aug); Miquel in Martius,Fl. Bras.4(1):
151. 1853; Snethlage, Notizbl. Bot. Gart. Berlin-
Dahlem 8: 369. 1923. Type. Peru. Withoutlocality,
(Y), Ruiz & Pav6n s.n. (holotype:FI-W,photograph
seen; isotypes: B, G, GH).
Cecropiascabra Klotzsch, Linnaea20: 531. 1847 (Oct),
non Martius (1841). Type. Peru. Without locality,
(5), Ruiz & Pav6n s.n. (holotype:B, destroyed,pho-
tographsex B in F, MO).
Cecropia ruiziana Klotzsch, Linnaea 20: 532. 1847
(Oct). Type. Peru. Withoutlocality, (5 + Y), Ruiz
& Pavon s.n. (syntypes:B, destroyed,photographin
F).
Cecropia nivea Poeppig ex Klotzsch, Linnaea 20: 532.
1847 (Oct). Type. Peru. Withoutlocality (or Huan-
uco: Cuchero),(5), Poeppigs.n. or 1593? (holotype:
B, destroyed,photographsin F, MO; isotype?:LE).
Cecropiapinnatiloba Klotzsch, Linnaea 20: 533. 1847
(Oct); Miquel in Martius,FH.Bras. 4(1): 153. 1853.
Type.Peru.Withoutlocality,(d + Y), Ruiz & Pavon
s.n. (syntypes:B, destroyed,photographin F; Y, K).
Cecropialeucophaea Poeppig ex Miquel, in Martius,Fl.
Bras. 4(1): 151, t. 50, f. 1. 1853. Type. Peru. Huan-
uco: Cuchero, (5), Poeppig s.n. or 1593 (W, de-
stroyed,fragmentin F, photographsin F, MO).
CecropiaklotzschianaMiquel, in Martius,Fl. Bras.4(1):
152. 1853, as synonym of C. scabra Klotzsch.
CecropiafrancisciSnethlage,Notizbl. Bot. Gart.Berlin-
Dahlem 8: 369. 1923; Berg, Acta Amazonica 8(2):
167. 1978. Type. Brazil. Acre: Seringal Sao Fran-
cisco, Oct 1911 (Y), Ule 9312 (holotype: B, de-
stroyed;isotypes: G, K, MG).
Cecropiaboliviana Cuatrecasas,RevistaAcad. Colomb.
Ci. Exact. 9(36/37): 335. 1956. Type. Bolivia. Santa
Cruz:Prov.Sara,Buenavista,3 Oct 1925 (Y), Stein-
bach 7267 (holotype:F; isotypes: A, G, GH, K, MO,
S).
Tree,to 20 m tall. Leafy twigs 3-6 cm thick, green
to red-brownwith conspicuouslenticels, hispidulous.
Lamina subcoriaceous,ca. 40 X 40 cm to 85 x 85
cm (to 100 X 100 cm), the segments 9-11(-12), the
free parts of the upper segments elliptic to oblong to
subobovate,the incisions down to ca. 5/10-7/10, the
largerlobes (especially in subjuvenilematerial)pin-
nately lobed (with the lobes pointingupwardin fresh
material); apices obtuse; upper surface smooth (to
152 FLORA NEOTROPICA

I 1~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~

[VQw1I
10 7 ~

FIG. 39. Cecropiapolystachya. 1. Lamina, reduced (Berg 1696). 2. Apex of lamina and venation (Fisher 178). 3.
Stipules and bases of petioles with trichilia (Nee 37514). 4. Stipules, young leaf, and base of petiole with trichilium(Berg
et al. 1641). 5. Pair of staminate inflorescences with spathes and base of petiole with trichilium (Berg et al. 1638). 6.
Staminateinflorescenceat anthesis and base of petiole with trichilium.7. Staminateflower and stamen (Berg 1696). 8. Pair
of pistillate inflorescenceswith spathesand base of petiole with trichilium(Berg et al. 1641). 9. Pistillateinflorescenceafter
anthesis and base of petiole with trichilium(D. N. Smithet al. 3123). 10. Pistillate flower and fruit (Beck 3793).
SYSTEMATICTREATMENT
scabridulous), the "umbilicus"white-hirsute;lower
surface on the main veins sparsely hirtellous to pu-
berulous to hispidulous (mainly with uncinate to
curvedhairs)and also with sparsebrownpluricellular
hairs, with arachnoidindumentumin the areoles and
on partof the reticulum;lateralveins in the free part
of the midsegment 15-25 pairs, submarginallyloop-
connected, most of them branched;petiole ca. 35-70
cm long, grayish, sparsely to densely hirtellous to
subhispidulous,with brown pluricellularhairs, and
with ? dense arachnoidindumentum, + conspicu-
ously lenticellate (especially in the lower part); tri-
chilia fused, the brown indument intermixed with
long white hairs; stipules 20-35(-45) cm long, ca-
ducous, red-brownto darkred (or greenish),but often
? grayish due to the indumentum,with ? dense
arachnoidindumentum,brownpluricellularhairs,and
simple whitish hairs outside, (sub)glabrous inside;
terminalbuds often curved. Staminateinflorescences
in pairs, erect to subpendulous;peduncle 8-14 cm
long, hirsute to hispidulous to puberulous,and with
arachnoidindumentum;spathe ca. 10-25 cm long,
red-brownto darkred or greenish to pinkish, but ?
grayish due to the indumentum, + dense arachnoid
indumentum),with sparse brown pluricellularhairs,
and scatteredlong white unicellularhairsoutside,gla-
brous inside; spikes ca. 15-45, 7-18 X 0.2-0.4 cm,
with stipes to 1.5 cm long and glabrous; rachis
sparselyhairy.Staminateflowers:perianthtubular,1-
1.5 mm long, glabrous,the apex ? distinctly2-lobed;
filaments flat; anthers ca. 0.6-0.8 mm long, appen-
diculate or not, remainingattachedto the filamentat
anthesis.Pistillate inflorescencesin pairs, erect, later
on the peduncle curved downward and the spikes
curved upward;peduncle 6-18(-26) cm long, with
indumentumlike that of the staminateinflorescence;
spathe ca. 8-12 cm long, the color and indumentum
as in the staminateinflorescence;spikes (2-)4-8, 5-
11 X ca. 0.4-0.8 cm, to 18 X 1.2 cm in fruit, some-
times initially tortuose, subsessile; rachis sparsely
hairy. Pistillate flowers: perianth ca. 1.5 mm long,
with arachnoidindumentbelow the apex outside, and
often also below the style channel inside, the apex
convex, muriculate, the aperture slit-shaped; style
short; stigma penicillate. Fruit oblongoid, ca. 2 mm
long, slightly tuberculate.
Distribution (Fig. 18.4). FromPeruto Bolivia and
Brazil (Acre and Rondonia),in forest and secondary
growth, at elevations to 1400(-1800) m.
Representative specimens examined. PERU. Cuzco:
Prov. La Convenci6n, Illapani, 1 Feb 1969 (d), ChavezA.
463, in herb. VargasC. 21274 (US); Prov. La Convenci6n,
Quiteni, 6 Jan 1976 (Y fl-fr),ChavezA. 3389 (MO); Prov.
153
Paucartambo,rd. Pilcopata-Atalaya,halfway, 24 Oct 1984
(Y fl), Maas et al. 6161 U); Prov. La Convenci6n, rd.
Cuzco-Quillabamba,ca. 150 km, 1100-1300 m, 14 Nov
1987 (Y fl), Ntiuez 8578 (MO); Prov. La Convenci6n,Rfo
Manguriari,2 Feb 1991 (9 fr), Nuinezet al. 12793 (BG).
HUANUCO: Nr. Tingo Marfa, 23 Jul 1940 (6), Asplund
12446 (S); rd. Huanuco-TingoMarfa,below Chinchao,ca.
1800 m, 23 Nov 1997 (6), Berg et al. 1735 (BG, COL,
MOL); rd. Huanuco-Tingo Marfa, nr. Tres Estrellas, ca.
1100 m, 24 Nov 1997 (9 fl-fr), Berg et al. 1740 (BG, COL,
MOL); ca. 90 km N of Tingo Marfa,rd. to Tocachi, 4 Feb
1984 (9 fl), Gentry et al. 44934 (NY); Prov. Pachitea,ca.
26 km S of PuertoInca, 18 Aug 1988 (6), Johannet al. 4/5-
18888 (BG), 15 Nov 1988 (9 fl-fr), Johann 2/32-151188
(BG); ca. 90 km N of Tingo Marfa,rd. to Tocachi, 4 Feb
1984 (9 fl-fr), Gentryet al. 44934 (MO, NY). JUNiN:Prov.
Tarma, nr. La Merced, 22 Jan 1946 (9 fl), Ferreyra398
(USM); between San Ram6n& La Merced,27 Feb 1982 (9
fl-fr), Gentryet al. 35775 (BG, USM); Prov.Chanchamayo,
Cataratadel Tirol, 3 Feb 1999 (d), NuiiiezL 85 (BG), (9 fl-
fr), NtiuiezI. 86 (BG); Prov. Satipo, Dtto. Satipo, Zapallal,
31 May 1982 (6), Reynel 480 (MOL); nr. San Ram6n,
Funda La Genova, 1300 m, 16 Apr 1989 (9 fl), Reynel et
al. 4016 (MO), 9 Sep 1989 (d), Reynel et al. 4417 (MO).
LORETO:Prov. Alto Amazonas, rd. to San Ram6n, 22 Sep
1954 (d), Ferreyra10186 (US, USM). MADREDE Dios:
Prov.Manu,nr.Salvaci6n,31 Jul 1988 (d), Berg etal. 1612
(BG, Centro de Medicina Andina, Cuzco); Prov. Tahua-
manu,Rfo Tahuamanu,16 Nov 1973 (9 fl), ChdvezA.1721
(MO); Prov.Tambopata,15 km ENE of PuertoMaldonado,
1991 (9 fl-fr),Fisher 178 (BG); ParqueNacionalManu,Rfo
Manu, Tayakome, 29 Sep 1986 (d), Foster et al. 11526
(US); between Shintuya & Salvaci6n, 31 Oct 1978 (9 fl),
Gentry et al. 27376 (BG); Prov. Tambopata,ReservaTam-
bopata,22 Aug 1990 (6), Reynelet al. 5396 (BG, MO); 39
km SW of Puerto Maldonado,above confluence of Rfo La
Torre & Rfo Tambopata,6 Oct 1985 (9 fl), S. E Smith et
al. 785 (NY, US); Prov. Tambopata,Las Piedras, Albergue
Cuzco Amaz6nico,26 Feb 1991 (9 fl-fr), Timand1539 (BG,
MO); Prov.Tambopata,Las Piedras,QuebradaGamitana,6
Mar 1991 (6), Timandet al. 1595 (BG, MO, NY). PASCO:
Nr. Pozuzo, 5 Feb 1983 (st), Gentryet al. 40079 (BG); Prov.
Oxapampa,Cedropampa,12 km S of Villa Rica, 1350 m, 18
Jan 1983 (9 fl-fr), D. N. Smithet al. 3123 (BG, MO). SAN
MARTiN:Prov.Lamas,rd. Lamas-Pamashto,ca. halfway,4
Dec 1997 (9 fl-fr), Berg et al. 1769 (BG, COL, MOL), (6),
Berg et al. 1770 (BG, COL, MOL); Prov. San Martfn,nr.
Tarapoto, 31 Aug 1968 (9 fl-fr), Ferreyra 17423 (US,
USM); rd. Tarapoto-Juanjui,km 34, 7 Oct 1984 (d), Maas
et al. 5981 (BG, U); Prov.MariscalCaceres,TocacheNuevo,
23 Jul 1974 (6), Schunke V 7694 (PMA, U). UCAYALI:
Nueva Requena, 11 km from Monte Verde,at km 33 on rd.
Pucallpa-Lima, 11 Aug 1988 (d), Berg et al. 1638 (BG,
COL, MOL); San Miquel de Semuya, S of Campo Verde,
12 Aug 1988 (9 fl-fr), Berg et al. 1641 (BG, COL, K,
MOL); jct. of Rio Pachitea & Rio Yuyapichis, SE of Pu-
callpa, 13 Oct 1985 (9 fl), Morawetzet al. 18-131085 (BG),
21 Oct 1985 (d), Morawetzetal. 11-211085 (BG); Semuya,
15 Mar 1988 (9 fl-fr), Rfos T 1801 (K).
154
BRAZIL. ACRE:Between Igarapedo Cuyubim& Igar-
ape Jocamin,FazendaBom Sossego, 27 Sep-7 Oct 1985 (st),
Campbellet al. 9720 (BG); Rio Branco, 18 Jun 1980 (5),
L. Coelho et al. 1795 (INPA, NY); nr. Sena Madureira,27
Sep 1980 (Y fl-fr), C. A. Cid Ferreiraet al. 2542A (INPA,
MG, NY); nr. mouthof Rio Macaua,5 Aug 1933 (5), Kru-
koff 5307 (A, BM, F, G, K, MICH, MO, NY, S, U, US).
ROND6NIA: Ji-Parana,Ouro Preto, 23 Jun 1981 (Y fl),
Fearnside1019 (INPA);rd. Ariquemes-OuroPreto,km 136,
21 Oct 1979 (d), Vieiraet al. 554 (INPA, NY, U).
BOLIVIA. BENI: Prov. Ballividn, rd. Caranavi-San
Borja, Serranfadel Pil6n Lajas, 30 Oct 1989 (5), D. N.
Smith 13772 (BG, MO); between Trinidad & Misiones
Guarayos,Sep 1926 (? fl), Werdermann2588 (LPB, MO,
S); Prov.Ballivian,Rio Beni, nr.Rurrenabaque,15 Sep 1989
(Y fl), D. E. Williams987 (BG). COCHABAMBA:Prov.Cha-
pare, Villa Tunari,30 Dec 1982 (Y fl), FerndndezC. et al.
7907 (G, NY); Prov.Chapare,Villa Tunari,24 Nov 1981 (Y
fl-fr), Va'zquezAvila 425 (U). LA PAZ: Prov. Larecaja,rd.
Caranavi-Guanay,km 26.8, 28 Nov 1980 (Y fl-fr), Beck et
al. 3793 (BG, LPB); Prov. Nor Yungas,rd. Chuspipata-Ya-
losa, km 14, 1150 m, 6 Aug 1988 (5), Beck 13913 (BG,
LPB, MO); Prov. Sud Yungas,nr. Sapecho, 26 Feb 1994 (Y
fl-fr), Berg 1696 (BG, LPB); Prov. Sud Yungas, Santa Ana
de los Mosetenes, 23 Mar 1990 (9 fl-fr), Hinojosa et al.
1104 (LPB); Prov. Sud Yungas, Rio Bopi, San Bartolome,
1-22 Jul 1939 (9 fl-fr), Krukoff10305 (A, G, K, MICH,
MO, NY, S, US), (5), Krukoff10465 (A, F, G, K, MICH,
MO, NY, S, U); Prov.SudYungas,nr. Sapecho,23 Oct 1993
(5), Seidel 7394 (BG, LPB); Prov. Nor Yungas,0.3 km N
of Yolosa on rd. to Caranavi,1200 m, 6 Oct 1984 (d), Sol-
omon et al. 12471 (AAU, BG); Prov. Nor Yungas, rd. Yo-
losa-Chuspipata,km 5.4, 1280 m, 6 May 1990 (5), Solo-
mon et al. 18965 (BG, MO). PANDO: Prov.Nicolas Suarez,
nr. Cobija, 7 Jan 1983 (9 fl-fr), Ferna'ndezC. et al. 8052
(G, NY); Prov. Nicolas Sudrez, Campo Ana, 14 Jan 1983
(5), FerndndezC. etal. 8241 (G, NY); Prov.Nicolas Suarez,
San Luis, 15 Aug 1990 (st), G. Gonzdlez12 (BG, LPB); nr.
Cobija, Oct 1911 (5), Ule 9311 (MG). SANTA CRUZ: Prov.
Andres Ibafiez,San Jose, 13 Nov 1994 (9 fl-fr),Mostacedo
et al. 2530 (NY, USZ); Prov. Andres Ibainez,Buena Vista,
18 Dec 1994 (9 fl-fr), Mostacedo et al. 2610 (NY, USZ);
Prov. Andres lbafiez, W of Santa Cruz, 16 Jan 1988 (9 fl),
Nee et al. 35924 (BG, LPB, NY, USZ); Prov.AndresIbdfiez,
Botanical Garden,E of Santa Cruz, 4 Jan 1989 (d fl), Nee
37513 (LPB, NY), (5), Nee 37514 (BG, LPB, MO, NY,
TEX), (9 fr), Nee 37515 (BG, LPB, NY); Prov. Andres
Ibafiez,SW of SantaCruz,Villa Fatima,4 Nov 1990 (9 fl),
Nee 39708 (BG, LPB, NY, USZ); Prov. Ichilo, 0-2 km SW
of El Carmen,ParqueNacional Ambor6, 9 Nov 1990 (d),
Nee 39842 (LPB, NY, USZ); Prov. Ichilo, ParqueNacional
Ambor6, Rio Saguayo, nr. mouth of QuebradaYapoje, 15
Jun 1991 (9 fl-fr), Nee 41051 (LPB, MO, NY, USZ); Prov.
Ichilo, 4 km WSW of Hondo, 14 Feb 1994 (9 fl-fr), Nee et
al. 44967 (NY, USZ); Nuflo de Chdvez, ReservasVida Sil-
vestre Rios Blanco y Negro, 14 Apr 1992 (9 fl-fr), Saldias
et al. 2409 (USZ); between Angostura& Samaipata,30 Oct
1980 (9 fl), Zuloaga et al. 1552 (U).
FLORANEOTROPICA
Young trees of this species usually have pinnately
lobed segments, especially in the upper part of the
lamina;the lobes point upward.In adult (flowering)
trees, pinnatelylobed leaf segments are less common
or the lobes areless pronounced.At the southernlimit
of the species distributionin Santa Cruz (Bolivia),
nearthe transitionof semideciduoussubtropicalforest
to chaco scrub forest, the tree shape is usually con-
spicuously different from the normal (lowland Ce-
cropia) tree shape in being more robust and usually
with distinctumbrella-shapedcrowns, shortandthick
trunks,and relatively short internodes.Moreover,in
this area the species is predominantlyrepresentedby
a green morph (with whitish stipules). The common
red morph, with reddish-greyishstipules occurring
exclusively elsewhere in the range of the species, is
far less common in the region indicated.The append-
ages at the bases of the thecae varies from well-
developed (to 0.1 mm long) to absent. Cecropiapo-
lystachya is essentially a lowland species, but in
HuainucoandJunin,Peru,it can be found at elevations
to 1800 m. Materialof this species collected by Ruiz
and Pavoncomprises specimenswith the segmentsof
the laminadistinctlylobed andleaves with (sub)entire
segments. Both Trecul (1847) and Klotzsch (1847)
described the two morphs as distinct species. Ruiz
and Pav6n collected both staminateand pistillate in-
florescencesof this species. Some sheets containboth
types of inflorescences(like those on which Klotzsch
described as C. pinnatiloba and C. ruiziana), others
only one of them. The names Cecropia leucophaea
and C. nivea are probablybased on the same collec-
tion made by Poeppig.
The species is (or has been) in cultivationin sev-
eral places in Brazil: Minas Gerais (Vicosa) and Sao
Paulo (Campinas,and Piracicaba),and it may have
become naturalizedin Mogi-Guacu.
BRAZIL. SAOPAULO: Campinas,Dtto. BaraoGeraldo,
CidadaUniversitaria,16Oct1978(6), Gabrielliet al. 8758
(UEC),(d + ), Gabrielliet al. 8759(UEC),Mun.Mogi-
Guaqu,ReservaBiol6gicade Mogi-Guaqu, 19 Nov 1992
(6), Godoiet al. 269 (SP),(9), Godoiet al. 270 (SP).
It has also been plantedin the Botanical Gardens
of Durban (South Africa: 1967, A. P Mills 22, A)
and Singapore (1967, Sidek bin Kiah S.60, A, C;
1979, J. E Maxwell 79-59, AAU). The species is
planted as an ornamentaltree along an avenue in
SantaCruz de la Sierra(Bolivia: 1994, Mostacedo et
al. 2391, NY).
Vernacular names and use. Peru:toroc (Cuzco);
yungul (Junin).Brazil: imbaubabranca (Rondonia).
Bolivia: ambaiboblanco (Beni, Santa Cruz).
SYSTEMATICTREATMENT
The fruiting spikes are sold in markets in Santa
Cruz de la Sierra.
45. Cecropia purpurascens C. C. Berg, Acta Ama-
zonica 7: 185. 1977, Acta Amazonica 8(2): 179.
1978. Type. Brazil. Amazonas:Nr. Manaus,Berg
264 (holotype: INPA; isotypes: MO, NY, P, RB,
U, US) = P 18808 (isotypes: COL, K, MG, P,
VEN, Z). Fig. 40
Tree,to 15 m tall. Leafy twigs 1-5 cm thick,green,
hispidulous to hirtellous, with curved to uncinate
hairs. Lamina (sub)coriaceous,ca. 20 X 20 cm to 65
X 65 cm, the segments (5-)6-7, the free partsof the
upper segments ovate, the incisions down to 4/10-5/
10; apices obtuse; upper surface scabrous, hispidu-
lous; lower surfaceminutelypuberulouson the veins,
on the larger veins also sparsely hirtellous with un-
cinate hairs to subtomentose, with arachnoidindu-
mentumin the areoles or sometimes also on the main
veins; lateralveins in the free partof the midsegment
8-10 pairs, submarginally loop-connected, usually
unbranched;petiole 20-50 cm long, puberulous to
hirtellous, or sometimes also with arachnoidindu-
mentum; trichilia fused, the brown indumentumin-
termixed with short white hairs; stipules 8-16 cm
long, pale red-brown,hirsuteoutside,subsericeousin-
side. Staminate inflorescences in pairs, with the pe-
duncle erect and the spikes pendulous;peduncle 6-
9(-15) cm long, hispidulous to puberulous;spathe
12-15 cm long, grayish red to orange-red,densely
hirtellous outside, glabrous inside; spikes 15-20, 7-
13 X 0.3-0.4 cm, with stipes 1.2-1.5 cm long and
minutelypuberulous.Staminateflowers connate;per-
ianth tubular,ca. 1.5 mm long, glabrous, the apex
plane; filaments ? swollen; anthersca. 0.5 mm long,
appendiculate,detachedat anthesis (?), reattachedto
the margins of the apertureby the appendages (?).
Pistillate inflorescences in pairs, erect, deflexed in
fruit; peduncle 5-10 cm long, hispidulous to puber-
ulous; spathe 8-12 cm long, the color and indumen-
tum as in the staminateinflorescence;spikes 4, 8-10
X 0.4 cm, to 13 X 1 cm in fruit;subsessile or stipes
to 0.5 cm long and minutelypuberulous;rachishairy.
Pistillateflowers: perianthca. 2 mm long, with arach-
noid indumentumbelow the apex outside, also in the
style channel inside, the apex truncate,muriculate;
style long, ? S-shaped,minutelypuberulous;stigma
comose. Fruit oblongoid, ca. 2.5 mm long, ? tuber-
ulate.
Distribution (Fig. 10.1). The middle Amazon ba-
sin, near and northof Manaus,in non-inundatedfor-
est.
155
specimensexamined.BRAZIL.AMA-
Representative
ZONAS: Rd. BR.174, Reserva Campina-INPA,7 Aug 1979
(9 fl-fr), Benson 10345 (BM, K, NY, RB, UEC); San Al-
berto,confluenceof Rio Negro & Rio Branco,25 Aug 1924
(9 fl-fr), Bequaert 3 (GH); rd. Manaus-Caracarai,km 65,
21 Sep 1973 (6), Berg 281 (K, NY, U); Distr.Agropecuario,
2?24-25'S, 59043-45'W, 18 Nov 1988 (9 fl-fr),Boom et al.
8533 (BG, MO, NY); Mun. Manaus, Reserva Florestal
Ducke, 28 Sep 1962 (d), Duarte 7188 (RB), 21 Jun 1994
(9 fl-fr), Hopkins et al. 1422 (BG, INPA); Distr. Agrope-
cuario, 2?24-25'S, 59043-45'W, 15 Nov 1988 (6), Mori et
al. 19817 (BG, INPA, NY); Mun. Manaus,ca. 80 km N of
Manaus,Distr.Agropecuarioda SUFRAMA,25 Jun 192 (9
fl-fr), Nee 42895 (INPA, K); Rio Negro, Camanau, 11 Jul
1995 (9 fl), Oliveira2771 (SPF);Manaus,Igarapedo Binda,
26 Jun 1961 (9 fl-fr), Rodrigues et al. 2090 (INPA); rd.
Manaus-Itacoatiara,km 66, 18 Aug 1970 (9 fl-fr), Ro-
drigues 8908 (INPA, MO).
This species is exceptional in the connate peri-
anths of the staminateflowers. It shows affinities to
Cecropiaficifolia and C. obtusa.
Vernacular name. Brazil: imbauba roxa (Ama-
zonas).
46. Cecropia putumayonis Cuatrecasas, Revista
Acad. Colomb. Ci. Exact. 6(22/23): 278. 1945;
Berg & FrancoRosselli, Fl. Ecuador48: 45. 1993.
Type. Colombia. Putumayo: Rio Putumayo, be-
tween Puerto Asis & Umbria, 24 Dec 1940 (9),
Cuatrecasas 11263 (holotype: COL). Fig. 41
Cecropia trilobata Cuatrecasas,Revista Acad. Colomb.
Ci. Exact. 9(36/37): 336. 1956. Type. Colombia.Ca-
queta: 97 km NW of Las Guacamayas,ca. 1200 m,
24 Apr 1944 (6), Little 7766 (holotype:F; isotypes:
COL, US).
Tree,to 10(-20) m tall. Leafy twigs ca. 1.5-3 cm
thick, green, puberulous.Laminachartaceous,ca. 25
X 25 cm to 60 X 60 cm, with 8-10 radiatingprimary
veins, only in the upper part distinctly incised down
to 3/10-5/10, separatinginto 3 lobes, the lower part
faintly lobed to subentire;apices acuminate;upper
surface ? scabrous,hispidulousand with very sparse
arachnoidindumentum,lower surface on the veins
puberulous,with arachnoidindumentumusually con-
fined to the margin, sometimes also present on the
main veins; lateral veins in the free part of the mid-
segment 7-11 pairs, submarginallyloop-connected,
most of them branched;petiole ca. 25-50 cm long,
with dense arachnoidindumentum;trichiliafused, the
brown pluricellular hairs intermixed with (rather)
sparse short hairs; stipules ca. 5-15(-20) cm long,
bright to dark red, puberulous outside, sparsely to
156 FLORA NEOTROPICA

-K>~~~~~~~~~~~~~~~2

6 '3

FIG.40. Cecropiapurpurascens.1. Leaf and stipules. 2. Apex of laminaand venation(Berg s.n.). 3. Pairof staminate
inflorescenceswith spathes and base of petiole with trichilium(Berg 281). 4. Staminateinflorescence at anthesis and base
of petiole with trichilium (Mori et al. 19817). 5. Staminate flower and stamen (Berg 281). 6. Pistillate inflorescence at
anthesisand base of petiole with trichilium(Pereiraet al. 2303.2165). 7. Pistillate flower.8. Fruitwith persistentstyle (Berg
et al. P 18808).
SYSTEMATIC TREATMENT 157

J~~~~~~~~~~~~~~~~~~~~~

10 cm.
i TnS ij j

FIG. 41. Cecropiaputumayonis.1. Leaf, stipules and base of petiole with trichilium.2. Apex of laminaand venation
(Gentry et al. 37903). 3. Stipules, staminateinflorescence with spathe, open prostoma,and bases of petioles with trichilia
(Gudino 586). 4. Staminateinflorescences with spathes and bases of petioles with trichilia (Palacios 1892). 5. Staminate
inflorescence at anthesis and base of petiole with trichilium (Lugo 4825). 6. Staminate flower. 7. Stamen (Franco et al.
4509). 8. Pistillate inflorescencein fruit and base of petiole with trichilium(Berg et al. 1093). 9. Pistillateflower. 10. Pistil.
11. Fruit(Franco et al 4678)
158 FLORA NEOTROPICA

densely sericeous inside. Staminateinflorescencesin QCNE). NAPO:Tena, 14 Oct 1939 (Y fl-fr), Asplund9301
pairs, pendulous;peduncle ca. 15-25 cm long, min- (S); rd. Coca-Auca, ca. km 50, 28 Feb 1980 ( fl-fr), Berg
utely puberulousand with sparse to dense arachnoid et al. 1093 (AAU, BG, COL, NY, QCA, TUR, U); Reserva
Biol6gica JatunSacha, 8 km E of Misahualli,22 Oct 1988
indumentum;spathe ca. 25-35 cm long, yellowish
(Y fl-fr), Cer6n et al. 5429 (MO, QAME, QCNE); Cant6n
brown,puberulousto hirtellousand usually also with
Archidona,rd. Hollin-Loreto, Rfo Huataraco,23-30 Aug
sparse to dense arachnoidindumentumoutside, gla- 1989 (st), Ceron et al. 7633 (MO, QCNE);San Jose de Pay-
brousinside; spikes 4-5, 20-26 X ca. 0.4 cm, sessile; amino, 40 km W of Coca, 3 Apr 1984 (st), Irvine 728 (F);
rachis hairy. Staminateflowers: perianthtubular,1- Codo Alto, between Santa Rosa & Cascada de San Rafael,
1.2 mm long, with short straighthairs on the margin ca. 1000 m, 14 Sep 1990 (d), Jaramilloetal. 12711 (QCA);
of the apex and lower down ? dense arachnoidin- ParqueNacional Yasuni, rd. Pompeya Sur-Iro, km 76, 13
dumentum,the apex plane; anthers 1-1.2 mm long, Aug 1994 (Y fl), Jaramillo et al. 16961 (QCA); rd. Coca-
not appendiculate,remainingattachedto the filament Tiputini, 5 Aug 1975 (d), Little et al. 61 (Q, QAME,
by a bundle of stretched spiral thickenings of tra- QCNE); Puyopungu, 15 Sep 1976 (6), Lugo 4825 (BG,
GB); Reserva Biol6gica JatunSacha, 1-15 Sep 1987 (d),
cheary elements at anthesis. Pistillate inflorescences
Palacios 1892 (BG, NY, QAME, QCNE);Cant6nEl Chaco,
in pairs, pendulous; peduncle 25-45 cm long, min-
Rio Quijos, Codo Sinclair, 16-20 Sep 1990 (d), Palacios
utely puberulousand with sparse to dense arachnoid 5741 (BG, QCNE); Afiangu, nr. mouth of Rio Afiangu,
indumentum;spathe ca. 30-35 cm, whitish, puberu- ParqueNacionalYasuni,30 Jun-9 Jul 1982 (d), SEF 10245
lous to hirtellous and usually also with dense (to (AAU, NY, QCA, U). PASTAZA: Between Mera& Moravia,
sparse) arachnoidindumentumoutside, almost gla- ca. 1000 m, 17 Dec 1955 (9 fl-fr),Asplund18877 (S); Can-
brous inside; spikes 4, ca. 30 X 0.5 cm, to ca. 50 X t6n Pastaza,Pozo petroleroCorrientes,1-31 Aug 1990 (6),
1.2 cm in fruit,sessile; rachishairy.Pistillateflowers: Gudino 519 (BG, MO, QCNE); 1 km S of Mera, ca. 1000
perianth ca. 2-2.5 mm long, with arachnoidindu- m, 6 Mar 1985 (9 fl-fr), Neill et al. 6003 (MO, QAME).
mentumbelow the apex outside,also in the upperpart SUCUMBiOS: Rd. Lago Agrio-Tarapoa, 1 Aug 1975 (6),
Little et al. 6 (Q, QAME, QCNE); Lumbaqui,ca. 1000 m,
of the style channel inside, the apex convex, finely
13 Aug 1975 (9 fl-fr), Little et al. 169, 178 (Q, QAME,
muriculate;style long, S-shaped, sparsely minutely
QCNE). ZAMORA-CHINCHIPE: Rd. Zamora-Gualaquiza,
puberulous;stigma penicillate to comose. Fruit ob- betweenYantzatza& Cumbaratza,4 Jan 1991 (d fl-fr),Berg
longoid, ca. 1.8 mm long, smooth. et al. 1660 (BG, LOJA,QCA).
Distribution (Fig. 18.5). From Amazonian Co- PERU. AMAZONAS: Rfo Cenepa, nr. Huampami,ca. 5
km E of ChdvezValdivia, 11 Jul 1978 (6), Ancuash 1020
lombia(Caqueta,Meta,andPutumayo)to Peru(Ama-
(BG), 25 Jul 1978 (d), Ancuash 1094 (BG, MO); Rio Ce-
zonas and Huanuco), mostly in forest in non-
nepa, QuebradaHuampani,ca. 5 km E of Chavez Valdivia,
inundatedplaces, at elevations to 1000(-1350) m. 11 Jul 1978 (9 fl-fr), Ancuash 1020 (BG); Prov. Bagua,
Representative specimens examined. COLOMBIA. Dtto. Imaza,Rio Marafi6n,Kampaenza,7 Oct 1994 (d), N.
CAQUETA: Mun. Florencia,rd. to Gabinete,850 m, 18 Oct Jaramilloet al. 517 (MO); Prov.Bagua,Yamayakat,11 Oct
1993 (d), Franco et al. 4509 (BG), 1000 m, 18 Oct 1993 1995 (9 fl-fr), N. Jaramillo et al. 851 (MO); Rio Cenepa,
(d), Francoet al. 4514 (BG); trailLas Guacamayas-Ramos, QuebradaHuampani,15 Feb 1973 (9 fl-fr), Kayap 368 (F,
km 59, 97 km NW of Las Guacamayas,ca. 1200 m, 24 Apr GH, MO, USM); Prov. Bagua, Dtto. Imaza, Rio Marafi6n,
1944 (Y fl-fr),Little 7765 (COL, F, US). META: Mun. Vi- Kampaenza,Feb 1995 (9 fl-fr), R. Vdsquezet al. 19532
llavicencio, rd. Villavicencio-Guayabetal,900-1100 m, 13 (MO); Prov. Bagua, Rio Marafi6n,opposite QuebradaMir-
Feb 1995 (Y fl-fr),Franco et al. 4678 (BG, COL,HUA, NY, and, 16 Sep 1962 (9 fl-fr), Wurdack2025 (F, NY, US). LOR-
ETO: Mouth of Rio Santiago, (9 fl-fr), Tessmann4033
US); Mun. Cubarral,Vrda. Agua Claras, 19 Nov 1995 (Y
fl-fr), Morales et al. 647 & 648 (COL). PUTUMAYO: Rio (NY). SANMARTiN: Prov. San Martin,rd. Tarapoto-Yuri-
San Miguel, between Rio Bermeja & Rio Conejo, 13 Dec maguas, ca. km 22, ca. 900 m, 5 Dec 1997 (9 fr), Berg et
km 20-27 km, 21 Jul 1982 (6),
1940 (d), Cuatrecasas11053 (COL);Mun. Mocoa, rd. Mo- al. 1774 (BG, COL, MOL),
Gentry et al. 37903 (BG, MO, USM).
coa-Villa Garzon,El Pepino, 18 Feb 1995 (Y fl-fr),Franco
et al. 4689 (BG, COL);Vrda.La Campucana,FincaLa Mar- This species shows clear affintiesto Cecropia ut-
iposa, 1350 m, 28 Apr 1994 (st), Franco et al. 5464 (BG); cubambana. They are similar in their usual habit:
Mocoa, between La Campucana& San Antonio, 1000 m, 2 slendertrees with relativelysmall and narrowcrowns
May 1994 (6 fl-fr), Franco et al. 5513 (BG).
and inflorescenceswith very long peduncles.The lat-
ECUADOR.MORONA-SANTIAGO: Rio Blanco, be-
tween Macas & Sucua,26 Jan 1981 (Y fr), Berg 1215 (AAU,
ter species has more deeply incised laminas,but such
BG, COL,GB, K, MO, NY, QCA, TUR, U); rd.Gualaquiza- a difference may occur in the same species (as in C.
San JuanBosco, km 15, 1200 m, 16 Feb 1994 (d fr), Berg andina and C. pastasana). However, the staminate
et al. 1694 (BG, QCNE); Rio Cuyes, between Bomboiza & flowersaredistinctlydifferent.This species occasion-
Gualaquiza, 1 Nov 1982 (d), Palacios 1447 (BG, QAME, ally shows anotherhabit:large trees with thick stems
SYSTEMATICTREATMENT
and broad crowns (Franco et al. 4678), collected at
the limit of its geographicaland elevationalrange.
Vernacular names. Ecuador:yura tsitica dundu
(Quichua, Napo). Peru: tseke, yawg tseke (Ama-
zonas).
47. Cecropia reticulata Cuatrecasas,Revista Acad.
Colombia Ci. Exact. 6(22/23): 279. 1945; Berg &
Franco Rosselli, Fl. Ecuador48: 47. 1993. Type.
Colombia. Valle: Rio Digua, Piedra de Moler,
900-1180 m, 19-28 Aug 1943 (Y), Cuatrecasas
15117 (holotype: COL; isotypes: F, US). Fig. 42
RevistaAcad.Colomb.
CecropiabracteataCuatrecasas,
Ci. Exact.9(36/37):331. 1956. Type.Colombia.
Valle:Quebrada de SanJuan,belowQueremal, 1350
m, 8 Nov 1946(Y), Cuatrecasas 22763 (holotype:
US;isotypes:F,VALLE).
Cecropia reticulata Cuatrecasas forma alboreticulata
Cuatrecasas,RevistaAcad.Colomb.Ci.Exact.9(36/
37): 329. 1956.Type.Colombia.Valle:Rio Dfgua,
betweenQueremal & LaElsa,1160-1200m,29 Mar
1947 (Y), Cuatrecacas24008 (holotype:US; iso-
type:F).
CecropiascutataCuatrecasas,RevistaAcad.Colomb.
Ci.Exact.9(36/37):334. 1956.Type.Colombia.An-
tioquia,nr.Dabeiba,21 Mar1948(Y), W Johnson
& E A. Barkley127 (holotype:US; isotypes:COL,
F, MEDEL,VALLE).
Tree,to 20 m tall. Leafy twigs 1.5-2 cm thick,grey
to blackish, with dense to sparse (filiform) brown
pluricellularhairs, glabrescent.Lamina chartaceous,
ca. 40 X 40 cm to 75 X 75 cm (to 90 X 90 cm), the
segments 7-9(-10), the free parts of upper segments
ovate to elliptic, the incisions down to 3/10-5/10(-7/
10) in the upperpartof the lamina, down to ca. 3/10
or less in the lower partof the lamina; apices obtuse
to rounded; upper surface scabridulous to almost
smooth, minutely hispidulous or partly puberulous,
sometimes ? bullate; lower surface puberulousand
also with brown pluricellularhairs, with (sometimes
very sparse) arachnoid indumentumin the areoles
and on the smaller veins, often (initially) also on the
main veins, or absent;lateralveins in the free partof
the midsegment 10-15 pairs, submarginallyloop-
connected, unbranched(or rarely branched);petiole
ca. 20-65(-80) cm long, sparselypuberulousandwith
sparseto dense arachnoidindumentum,mostly on an
abaxiallybulging (subscrotiform)base of the petiole;
trichiliafused, large, mostly extendedwith lateralas-
cending lobes, the brown indumentum intermixed
with dense long (curvedto straight)white hairs;stip-
ules ca. 10-20(-25) cm long, red-brown(to whitish),
159
caducous,densely sericeous and/orwith dense arach-
noid indumentumoutside, very sparselyhairy to gla-
brous inside. Staminateinflorescencesin pairs, sub-
tended by caducous (to subpersistent)bracts, to 7.5
cm long, the peduncleerect and the spikes erect to ?
spreading;peduncle 2-7 cm long, tomentose to stri-
gose, often also with sparse arachnoidindumentum,
or subglabrous;spathe 8-14 cm long, brown to red-
brown,with dense brownpluricellularhairsandoften
also with (sparseto ratherdense) arachnoidindumen-
tum and/or sparsely sericeous outside, (sub)glabrous
inside; spikes 4-12, 6-12 X 0.4-0.8 cm, sessile; ra-
chis hairy or glabrous. Staminateflowers sessile or
with pedicels to 0.6 mm long; perianthtubular,1.2-
1.5 mm long, glabrous,the apex convex to plane; fil-
aments flat; anthers0.6-0.8 cm long, appendiculate,
detachedat anthesis (?), reattachedto the marginsof
the apertureby the appendages(?). Pistillate inflores-
cences in pairs,erect, subtendedby caducous(to sub-
persistent) bracts, to 3 cm long; peduncle 3-5 cm
long, densely hirtellousto hispid;spathe5-9 cm long,
the color and indumentumas in the staminateinflo-
rescence;spikes 4-5, 4-7 X ca. 0.5 cm, to 1 cm diam.
in fruit, sessile; rachis hairy. Pistillateflowers: peri-
anth 1.5-2 mm long, with arachnoidindumentumbe-
low the apex outside, also in the lower partof the style
channel inside, the apex convex, finely granulateto
muriculate;style short;stigma penicillateto comose.
Fruit ellipsoid, ca. 2 mm long, ? tuberculate.
Distribution (Fig. 18.5). Western Ecuador and
Colombiaand adjacentPanama,in forest and second-
ary growth,at elevationsto ca. 1800 m, mostly above
1000 m.
Representative specimens examined. PANAMA.
DARIEN: Cerro Pirre, 1200 m, 16 Nov 1989 (Y fr), Fisher
57 (BG); N of Cerro Pirre, between Cerro Pirre & Rancho
Pastico, 1200-1400, 14 Nov 1977 (st), Folsom et al. 6351
(BG); Summit of Cerro Pirre, 1000-1400 m, 29 Dec 1972
(d), Gentryet al. 7022 (BG, MO).
COLOMBIA. ANTIOQUIA:Mun. Anorn,Vrda. El Car-
men, km 12-16 NW of Anorn,940-1100 m, 18 Nov 1989
(6), Callejas et al. 8757 (BG, HUA, NY); Yarumal,1725
m, 27 Mar 1994 (6), Franco et al. 4595 (BG HUA); Mun.
Frontino,Cgto. Nutibara,rd. Nutibara-LaBlanquita,Murri
region, 1650-1700 m, 4 Mar 1995 ( fl-fr), Franco et al.
5551 (BG, COL, HUA, LP, US), (6), Franco et al. 5552
(BG, COL, HUA, LP,US), 1250 m, 4 Mar 1995 (6), Franco
et al. 5557 (BG, COL, HUA, LP); Mun. Frontino, Cgto.
Nutibara,source of Rio Cuevas, 1630 m, 15 Jan 1984 (Y fl-
fr), D. SdnchezS. et al. 990 (BG, MEDEL). CAUCA: W of
Rio San Joaquin,1400-1500 m, 29-30 Jun 1922 (Y fl), Kil-
lip 7873 (GH, NY, US). CHoCO:Mun. San Jose del Palmar,
Rio Torito, Finca Los Guadales, 1 Mar 1980 (Y fl), Forero
et al. 6480 (COL, US); Mun. San Jose del Palmar,Vrda.El
160 FLORA NEOTROPICA

~~~~~ 1~~~~~~I0

FIG.42. Cecropia reticulata. 1. Leaf, reduced.2. Apex of lamina and venation. 3. Stipules and young leaf. 4. Base
of petiole with trichilium.5. Pairof staminateinflorescenceswith subtendingbracts(Berg 1244). 6. Staminateinflorescence
with spathe (Dodson et al. 12747). 7. Staminate inflorescence at anthesis (Berg 1244). 8. Staminate flower and stamen
(Franco et al. 4696). 9. Pair of pistillate inflorescences with spathes (0llgaard et al. 57707). 10. Pistillate inflorescenceat
anthesis and base of petiole with trichilium(Dodson 6668). 11. Staminateflower. 12. Fruit(Franco et al. 5551).
SYSTEMATICTREATMENT
Sinai, 1800 m, 19 Mar 1994 (Y fl-fr), Franco et al. 4560
(BG, HUA). NARINO:Mun. Barbacoas, El Diviso, 7 Oct
1993 (st), Franco et al. 4462 (BG); Mun. Ricaurte, rd.
Ricaurte-Altaquer,1300 m, 8 Oct 1994 (6), Franco et al.
4474 (BG); nr. Altaquer,1150 m, 21 Feb 1995 (6), Franco
et al. 4696 (BG, COL, HUA, LP, US); Mun. Ricaurte,Res-
guardo Indigena Nulpe Medio, QuebradaLa Conga, 8 Jan
1996 ( fl-fr),M. S. Gonzdlezet al. 1619 (COL);Mun. Nar-
bacoas, Resguardo El Sabalo, Rio Cangapi, 21 Aug 1995
(6), M. S. Gonzalez et al. 8200 (COL). RISARALDA:Mun.
Mistrat6, rd. San Antonio de Chami-Mistrat6, Quebrada
Sutu, 27 Apr 1992 (9 fl-fr), Alonso et al. 10193 (COL).
VALLE: Rio San Juan,below Queremal, 1300-1700 m, 19
Mar 1947 (6), Cuatrecasas23883 (US, VALLE);Mun.Cal-
ima, Vrda. Campo Alegre, 1000 m, 4 Oct 1983 (6), Devia
396 (TULV).
ECUADOR. AZUAY: Nr. Molleturo, 25-27 Jan 1991
(6), L. Ortiz 203 (QCA, QCNE). CARCHI: Cerro Golon-
drinas, Upper Rio Pablo drainage, 1730-1760 m, 24 Apr
1993 (st), Boyle et al. 1730 (BG); San Marcos de los Coai-
queres, trail Chical-TobarDonoso, 10 Feb 1985 (9 fl-fr),
0llgaard et al. 57707 (AAU, QCA); Chical, 1700-1900 m,
20-29 Jul 1991 (9 fl-fr), Quelal et al. 273 (BG, QCNE);
Cant6nTulcan, Reserva IndigenaAwa, San Marcos, 25 km
NW of Chical, 1500 m, 16-30 Nov 1990 (6), Rubio et al.
978 (BG, QCNE); TobarDonoso, 19-28 Jun 1992 (6), Ti-
paz et al. 1480 (BG, QCNE). COTOPAXI: Rd. Quevedo-
Latacunga, nr. El Palmar, 17 Feb 1981 (6), Berg 1280
(AAU, BG, COL, GB, MO, QCA, U); rd. Quevedo-Lata-
cunga, km 52-53, Tenefuerte,Rio Pilal6, 750-1300 m, 21
Feb 1982 (d), Dodson et al. 12724 (MO, QCNE);rd. Pilal6-
Quevedo, between Macuchi & La Mana, 29 Jul 1980 (6),
Holm-Nielsenet al. 24705 (AAU); Rio Guapara,20 km NW
of El Coraz6n,24 Jun 1967 (6), Sparre17358 (S). EL ORO:
Between La Vega & Rivera, Rio Chilola, 19 Nov 1994 (9
fl), Cornejoet al. 3578 (BG); rd. Pifias-SantaRosa, km 12.5,
7 Oct 1979 (6), Dodson et al. 8959 (MO, QCNE), (9 fr),
Dodson et al. 8960 (MO, QCNE). ESMERALDAS: Mun.
Quininde,Bilsa Biological Station,Mache Mtns., 35 km W
of Quininde,Fila de Bilsa, 7 km E of San Jose de Bilsa, ca.
80 km SW of Esmeraldas,4 Oct 1994 (9 fl), J. L. Clarket
al. 140 (BG, QCNE), 30 Jan 1991 (6), Gentryet al. 72939
(QCNE). IMBABURA:Nr. Lita, 7 Feb 1981 (6), Berg 1244
(AAU, BG, COL, GB, K, MO, QCA, U, WIS). Los Rios:
Rio Palenque Biological Station, rd. Quevedo-Santo Do-
mingo de los Colorados,km 56, 2 Oct 1976 (9 fl-fr), Dod-
son 6451 (F, MO, QCA, US), 5-14 Mar 1977 (9 fl-fr),Dod-
son 6668 (AAU, MO, QCA). Los RiOS/PICHINCHA: El
Centinela, at crest of Montaniasde Ila, rd. PatriciaPilar-24
de Mayo, km 12, 27 Nov 1978 (6), Dodson 7304 (BG, MO,
QCNE, U). MANABf: MachalillaNational Park,San Sebas-
tian, 20 Jan 1991 (6), Gentry et al. 72460 (MO, QCNE).
PICHINCHA: Nr. Santo Domingo de los Colorados, Hda.
Gloria Maria, 18 May 1955 (9 fl-fr), Asplund 16402 (S);
rd. Tandapi-SantoDomingo de los Colorados, km 20, 13
Sep 1977 (6), Berg et al. 422 (AAU, BG, COL, MO, QCA,
U); Rio Guajalito,ca. 4 km N of Palmeras,km 59 on rd.
Quito-San Juan-Chiriboga-Empalme,1800 m, 15 Dec 1990
(6), Berg et al. 1647 (BG, QCA); rd. Nono-Rio Yacuambi,
161
5-10 km above Nanegalito, 1700 m, 21 Jul 1980 (d), Holm-
Nielsen et al. 24410 (AAU); Rio Guajalito,ca. 4 km N of
Palmeras, km 59 on rd. Quito-San Juan-Chiriboga-Em-
palme, 1800 m, 29 Feb 1992 (d), Jaramillo et al. 14657
(NY, QCA).
The arachnoid indumentum on various parts varies
from dense to almost lacking, the latter state found in
particular in the northern part of the distribution range
(Antioquia). Moreover, in the northern part of the spe-
cies range, the incisions are often less deep (down to
ca. 3/10) and the characteristic shape of the trichilia
on a subscrotiform base of the petiole wanting. This
northern type has been described as Cecropia scutata
and might represent a distinct infraspecific entity, oc-
curring in Antioquia at elevations higher than the
more typical one with the lamina more deeply incised
(down to ca. 5/10), subscrotiform bases of the peti-
oles, and trichilia with ascending lateral lobes.
From some distance the leaves (and therefore the
trees) often look whitish in spite of the absence of
arachnoid indumentum on the upper surface of the
lanmina. The bracts subtending the peduncle are con-
spicuously present in fresh material, but as they are
mostly caducous, usually absent in herbarium mate-
rial.
Vernacular name. Colombia: cosedera (Narifio).
48. Cecropia sararensis Cuatrecasas, Revista Acad.
Colomb. Ci. Exact. 6(22/23): 292. 1945. Type. Co-
lombia. Norte de Santander: Sarare region, Rfo
Cubugon, El Indio, 13 Nov 1941 (9), Cuatrecasas
13098 (holotype: COL, isotype: F).
Cecropia cobariana Cuatrecasas, Revista Acad. Col-
omb. Ci. Exact. 6(22/23): 278. 1945. Type. Colom-
bia. Norte de Santander:SarareRegion, confluence
of Rio Cubug6nand Rio Cobaria,El Banco, 15 Nov
1941 (9), Cuatrecasas13197 (holotype: COL; iso-
type: F).
Cecropia libradensis Cuatrecasas,Revista Acad. Col-
omb. Ci. Exact.9(36/37): 337. 1956;Velasquez,Acta
Bot. Venez. 6: 43, t. 7. 1971. Type. Colombia.Norte
de Santander:SarareRegion, Quebradade la China,
SantaLibrada,El Reposo, 800 m, 20 Nov 1941 (d),
Cuatrecasas13355(holotype:F; isotype: COL).
Tree, to 15 m. Leafy twigs 1.5-4 cm thick, green
to brownish with conspicuous lenticels, sparsely to
rather densely hispidulous with curved hairs to gla-
brous. Lamina chartaceous to subcoriaceous, ca. 25
X 25 cm to 70 X 70 cm, the segments 8-10(-13), the
free parts elliptic to oblong, the incisions down to 6/
10-8/10; apices subacute to subacuminate or to ob-
tuse; upper surface smooth or ? scabrous, subgla-
162
brous to hispidulous;lower surface sparselyto rather
densely (minutely) puberulousand with longer unci-
nate hairsto subglabrouson the veins, with arachnoid
indumentumin the areoles and on the smaller veins;
lateral veins in the free part of the midsegment 11-
18 or 18-24 pairs, marginally loop-connected, un-
branchedor (often only the lower ones) branched;pet-
iole 15-45 cm long, sparsely (to densely) puberulous
to subglabrous,sometimes with sparse arachnoidin-
dumentum,the base of the petiole sometimes ? bulg-
ing (subscrotiform);trichilia fused, the brown indu-
mentum intermixed with short white to brownish
hairs; stipules 7-18 cm long, greenish to yellowish
brown or red(dish), with only brown pluricellular
trichomes or also sparsely strigillose or sparsely to
densely whitish to brownish hirtellous to subhirsute
to sericeous, sometimesalso with sparse(to ? dense)
arachnoidindumentumoutside, (densely) yellowish
sericeous to villous inside. Staminate inflorescences
in pairs, (sometimes?)subtendedby caducousbracts,
to 3 cm long, the peduncle erect to deflexed and the
spikes spreading to pendulous; peduncle 4-17 cm
long, (sub)glabrousor sparselypuberulous,hirtellous
or subhirsute,sometimes in the lower partwith sparse
arachnoidindumentum;spathe9-19 cm long, green-
ish, yellowish (brown),red(dish)or whitish,with only
brown pluricellular trichomes or also sparsely to
ratherdensely whitish to brownishhirtellousto sub-
villous, sometimes sparse to ? dense arachnoidin-
dumentumoutside, densely whitish to yellowish to
brownishsericeous to villous (or subglabrous)inside;
spikes 4-18, 6-17 X 0.3-0.5 cm, with stipes 0.2-1
(-2) cm long and sparsely hirtellousor minutely pu-
berulous;rachisglabrous.Staminateflowers:perianth
tubular,1-1.5 mm long, glabrous or sparsely puber-
ulous below the apex, the apex plane; filamentsflat;
anthersca. 0.6 mm long, appendiculate,detachedand
reattachedto the margins of the apertureby the ap-
pendages.Pistillate inflorescencesin pairsor solitary,
erect to (at least in fruit) pendulous, (sometimes?)
subtendedby caducousbracts,to 2 cm long; peduncle
7-19 cm long, subglabrousor sparsely (to densely)
puberulousto hirtellous to subhirsute,sometimes in
the lower part with sparse arachnoidindumentum;
spathe 8-15 cm long, the color and indumentumas
in the staminateinflorescence;spikes (3 or) 4, 6-13
X ca. 0.5 cm, to 33 X 1 cm in fruit, (sub)sessile or
with stipes to 0.8 cm long and glabrous; rachis
(sub)glabrous.Pistillateflowers free or connateat the
base; perianth ca. 1.5 mm long, with arachnoidin-
dumentumbelow the apex or to nearthe apertureout-
side, absent (or present in the style channel) inside,
the apex plane, muriculateor smooth;stigmatruncate
FLORA NEOTROPICA
andpenicillateto peltate.Fruitoblongoidto ellipsoid,
ca. 2 mm long, smooth, blackish.
Distribution (Fig. 18.6). Andean region of Vene-
zuela and in Colombiain northernpartof the eastern
slopes of the eastern Cordillera,southwardto Meta,
in forest and secondary growth, at 300-1400 m, in
Venezuelato 2300 m.
Specimens examined. COLOMBIA: ARAUCA: Mun.
Fortul,Vrda.Palmarito,26 Jan 1995 (d), Franco et al. 4632
(BG, COL, HUA, US), (Y fl-fr), Franco et al. 4633 (BG,
COL,HUA,US);Mun.Tame,rd.to Sacama,26 Jan1995
(Y fl-fr), Franco et al. 4634 (BG, COL, HUA, US).
BOYACA: Mun. Pajerito, between Corinto & Peniadel Gallo,
1200 m, 29 Jan 1995 (Y fl-fr), Franco et al. 4642 (COL,
HUA,MO,US). CASANARE: Rd.to Sacama, 29 Jan1995
(6), Franco et al. 4635 (COL), (Y fl-fr), Franco et al. 4636
(COL); Sacama, 29 Jan 1995 (d), Franco et al. 4637 (BG,
COL). CUNDINAMARCA: Rd. Guayabetal-Villavicencio,
1035-1150 m, 29 May 1993 (d), Franco et al. 4459 (BG,
COL).META:Above Rio Negro,ca. 20 kmW of Villavi-
cencio, 1100 m, 23 Feb 1972 (Y fl-fr), A. S. Barclay et al.
3197 (COL,US); nr. Villavicencio,10 Feb 1995 (Y fl),
Francoet al. 4655 (BG, COL), (6), Francoet al. 4656 (BG,
COL), (d), Franco et al. 4663 (BG, COL); rd. Villavicen-
cio-Guayabetal,900-1 00 m, 13 Feb 1995 ( Y fl-fr),Franco
et al. 4672 (COL), (6), Franco et al. 4673 (BG, COL), (d),
Franco et al. 4674 (BG, COL), (Y fl-fr), Franco et al. 4675
(BG, COL); Llanos de San Martin,Rio Humadea,Guamal,
10 Sep 1958 (Y fl-fr),R. Jaramillo-Mejiaet al. 1017 (COL).
NORTE DE SANTANDER:Mun. Toledo, Cgto. San Bernardo,
1250 m, 23 Jan 1995 (d), Franco et al. 4623 (BG, COL);
Mun. Toledo, rd. La Mesa-Samore, 1150 m, 23 Jan 1995
(d), Franco et al. 4626 (BG, COL), (Y fl-fr), Franco et al.
4627 (BG, COL), (d), Franco et al. 4628 (BG, COL);Mun.
Toledo, Samore, 25 Jan 1995 (Y fl-fr), Franco et al. 4629
(BG, COL);nr.Chinacota,1400 m, 18 Mar 1927 (6), Killip
et al. 20852 (A, GH, NY, US).
VENEZUELA. BARINAS: Mun. Pedraza, Alto del
Aguada, 1250-1300 m, 22 Feb 1955 (Y fr), Bernardi2031
(NY); rd. Barinitas-ElCacao, 7 Jul 1981 (Y fl-fr),Marcano
Berti et al. 277-981 (U, US); rd. Barinas-San Crist6bal,El
Corozo, Dec 1966 (d), Veldsquez101 (VEN). FALC6N:
Cerro Galicia, Dtto. Petit, 15 Dec 1977 (juv), T Ruiz et al.
2806 (VEN). LARA: Dtto. Jim6nez, Parque Nacional Ya-
cambu, QuebradaNegra, 24 Oct 1982 (d), Davidse 20921
(MO, VEN). MERIDA: Dtto. Tovar, rd. Tovar-Zea, 900-
1000 m, 1 Jul 1954 (Y fl-fr), Bernardi1313 (NY); between
Estanquez& La Coloradas,12 May 1982 (Y fl-fr),Marcano
Berti et al. 982-013 (G, U, US, VEN); between Merida &
Santo Domingo, 500-2500 m, Feb 1968 (d), Velasquez261
(US, VEN), (Y fl-fr), Velasquez262 (US, VEN). PORTU-
GUESA: Concepci6n, 10 Jan 1967 (d), R. E Smith V526
(VEN). TACHIRA: 10 km E of La Fundaci6n,nr. Represa
Dorada,600-1000 m, 1-13 Mar 1981 (9 fl), Liesner et al.
10437 (MO,VEN); CerroLas Minas, 18-20 km SE of Santa
Ana, 1150-1250 m (9), Steyermnark et al. 119050 (MO,U).
TRUJILLO: Dtto. Bocon6, La Morita, above Rio Saguras,S
SYSTEMATICTREATMENT
of CampoElias,2300m,4 Jun1988(Y fl),Dorret al. 5367
(BG,NY).
The stigma varies from penicillate to peltate, and
linked to this difference, the arachnoidindumentum
varies from present only below the apex of the peri-
anth to presentnear the apertureof the perianth.The
indumentumof the stipules and spathes varies from
brown to white subhirsuteto subvillous to subgla-
brous (only with brownpluricellularhairspresent)to
(sometimes) with ? dense arachnoidindumentum.A
mixture of the various types of indumentumis most
common in the middle part of the range of this spe-
cies. The numberand the length of the spikes of the
staminateinflorescencevariesfromnumerousandrel-
atively shortto few andrelativelylong. In the northern
partof the range (Venezuela),the materialwith more
numerouslateralveins (18-24 pairs) in the free part
of the midsegment,with spathesbeing (sub)glabrous
outside, with staminateinflorescences with few and
long spikes, and with comose(-penicillate) stigmas
appearsto be predominant.In this part of the range,
Cecropiasararensis reaches to elevations of 2300 m.
The type with these features includes the type of C.
libradensis. Cecropiasararensis, as it is delimitedin
the present treatment, is rather heterogeneous and
may prove to comprise two entities, parallel to the
two subspecies of C. schreberiana, showing differ-
ences rathersimilar to those between the two types
circumscribedabove. However,with the materialand
data availableat present, it is not possible to find ac-
ceptable delimitations of these types. Moreover, C.
sararensis as a whole can barely be distinguished
from C. peltata and C. metensis,the neighboringtaxa
of the C. peltata group. Cecropia sararensis can be
distinguished from C. metensis by the absence of
dense white arachnoidindumentumon the stipules,
spathes, and (also) the petioles, and from the former
by subglabrousstipules, spathesand leafy twigs or by
the relativelylong brownishhairson the stipules(and
spathes).
49. Cecropia saxatilis Snethlage,Notizbl. Bot. Gart.
Berlin-Dahlem8: 360. 1923; Berg & Carauta,Al-
bertoa 1(1): 10. 1986. Type.Brazil. Syntypes.Bra-
zil, Piuai, Serra Branca, Jan 1907 (d), Ule 7174
(B, destroyed, K, only an inflorescence),(?) Ule
7174 (B, destroyed), specimen with pistillate in-
florescence(s) at K here designated as lectotype
(K; isolectotype: G).
Tree,to 10 m tall. Leafy twigs 2-4 cm thick, hir-
tellous with curved to uncinate hairs or also with
163
sparseto ratherdense arachnoidindumentum.Lamina
subcoriaceousto coriaceous, ca. 15 X 15 cm to 50 X
50 cm, the segments 6-10, the free partsof the upper
ones (broadly)obovate, the incisions down to ca. 5/
10-7/10; apices rounded;upper surface scabrousto
almost smooth, sparsely to densely hispidulous or
subvelutinous,sometimes with (ratherdense) arach-
noid indumentum;lower surfaceon the (main) veins
subtomentoseto hirtellous,with arachnoidindumen-
tum almost confinedto the areoles or extendedto the
main veins; lateral veins in the free part of the mid-
segment 8-10 pairs, submarginally to marginally
loop-connected, branched;petiole 10-35 cm long,
reddish, puberulousand with sparse to dense arach-
noid indumentum;trichilia fused, the brown indu-
mentum intermixed with short and ratherlong (?
bristle-like) white hairs; stipules 8-15 cm long,
reddish, with ? dense arachnoidindumentumand
hirtellousoutside, sparselyhairy inside. Staminatein-
florescences in pairs, erect or the peduncle erect and
the spikes pendulous(?); peduncle3-9 cm long, sub-
hispidulous to hirtellous and with arachnoid indu-
mentum; spathe 8-18 cm long, red to purplish,
sparsely puberulous to densely hirtellous and with
sparse to dense arachnoidindumentumoutside, gla-
brous inside; spikes 4-8, 7-17 X 0.3-0.8 cm, with
stipes 0.4-1.5 cm long and puberulousand/or with
arachnoidindumentum;rachis (sub)glabrous.Stami-
nateflowers: perianthtubular,ca. 1.5 mm long, with
dense arachnoid indumentumbelow the apex, the
apex muriculateto smooth, the aperturesurrounded
by a low rim; filamentsflat;anthers0.8-1.2 mm long,
appendiculate,detachedat anthesisbut remainingat-
tachedto the filamentby stretchedspiralthickenings.
Pistillate inflorescencesin pairs, erect to pendulous
(?); peduncle4-23 cm long, subhispidulousto hirtel-
lous and with arachnoid indumentumor glabrous;
spathe 6-8 cm, the color and indumentumas in the
staminateinflorescence;spikes4, 6-15 X 0.5-0.8 cm,
to 28 X 1.5 cm in fruit, (sub)sessile; rachisglabrous.
Pistillate flowers: perianth ca. 1.5 mm long, with
arachnoidindumentumbelow the apex outside, also
in the style channel inside, the apex slightly convex,
muriculateto smooth; style short;stigma penicillate.
Fruit oblongoid to subovoid, ca. 2.5 mm long, tuber-
culate, darkbrown.
Distribution (Fig. 8.6). CentralBrazil andeastern
Bolivia (Santa Cruz), in cerrado areas, usually on
sandstoneor limestone rocks, in Bolivia in grassy sa-
vanna, at elevations to 1400 m.
Representativespecimensexamined.BRAZIL.BA-
HIA: Espigao Mestre, 34 km W of Barreiras,2 Mar 1972
164
(9 fl-fr),Andersonet al. 36458 (BG, NY, US); Mun. Gentio
do Ouro,nr.SantoInacio, rd. to Xique-Xique,Serrado Acu-
rua, 27 Nov 1992 (d), Arbo et al. 5346 (BG, K, SPF); nr.
Barreiras,3 Jun 1991 (9 fl-fr), Brito et al. 333 (GUA, NY),
(d), Brito et al. 336 (NY); Mun. Oliveira dos Brejinhos,
Serra de Agua Quente, 16 Apr 1999 (9 fl-fr), Forza et al.
1242 (NY); valley of Rio das Ondas, Espigao Mestre, ca. 5
km NW of Barreiras,4 Mar 1971 ( 9 fl-fr),Irwinetal. 31509
(BG, NY, UEC, US). DISTRITO FEDERAL: Rd. Campo
Grande-Cuiaba,BR.163, 30 Jan 1979 (d + 9 fl), Heringer
et al. 997 (NY, UEC, US). GoiAs: Chapadados Veadeiros,
6-7 km E of Alto Paraiso,ca. 1400 m, 7 Mar 1973 (9 fr),
Andersonet al. 6552 (MO, NY, U, US); Chapadados Vead-
eiros, S of Terezina,ca. 1000 m, 19 Mar 1973 (9 fr), An-
derson 7495 (MO, NY, U, US); Vila Boa de Goias, 22 Jan
1969 (9 fr), Carauta 723 (GUA, U); Mun. Alto Paraisode
Goias, rd. Terra-NovaRoma, 6 km E of Alto Paraiso de
Goias, 1200 m, 13 Mar 1995 (9 fl-fr), Cavalcantietal. 1294
(SPF); Campos Anayas, Mar 1840 (9 fl-fr), Gardner3981
(BM, K) Serra Dourada,21 Jan 1966 (5), Heringer 1096
(US); SerraDourada,20 km SE of Goias Velho, 16 Jan 1966
(5), Irwin et al. 11799 (G, NY, RB, S, SP, US); SerraGeral
do Parana,ca. 20 km S of Sao Joao de Alianqa,ca. 1000 m,
17 Mar 1971 (9 fl-fr), Irwin et al. 32099 (BG, MOL, NY,
QCA, US); Mun. Alto Paraiso, rd. to Cavalcanti,Chapada
dos Veadeiros, 1200-1300 m, 13 Feb 1990 (9 fl), Hatsch-
bach et al. 53946 (SP); Serra Dourada, 21 Jan 1966 (d),
Heringer 10961 (NY), 22 Jan 1978 (5 + 9 fl-fr), Rizzini
(RB) 205206 (GUA, RB). MATO GROSSO: Chapadados
Guimaraes, nr. Portao do Inferno, 27 Jan 1989 (9 fl-fr),
Carauta5776 (GUA, NY), (d), Carauta5777 (GUA, NY);
Mun. Rondon6polis,rd. BR.163, km 15, 12 Nov 1975 (d),
Hatschbach37457 (U); CampoGrande-Cuiaba,30 Jan 1979
(9 fl-fr),Heringeret al. 997 (GUA); nr.Diamantina,23 Apr
1983 (9 fl-fr),Moraes et al. 279 (MG);jct. of rds.to Cuiaba,
Santarem& PortoVelho, 4 Feb 1979 (9 fl), M. G. Silva et
al. 4441 (MG, NY, U). MINASGERAIS: 15 km W of Janu-
aria, 20 Apr 1973 (9 fl-fr), Anderson 9258 (MO, NY, U,
US); Mun. Januaria,Dtto. de Fabiao,24 Jan 1997 (5), Lom-
bardi et al. 1757 (BHCB), 16 Feb 1998 (9 fl), Lombardiet
al. 2214 (BG, BHCB). PUAi: Sao RaimundoNonato, Serra
da Capivara,13 Dec 1978 (st), Freires.n. (GUA).
BOLIVIA. SANTACRUZ:Prov.Velasco, rd. SantaRosa
de la Roca-Concepci6n, km 45, 9 Dec 1994 (9 fl-fr), Guil-
ln et al. 2686 (USZ); Prov.Velasco, EstanciaFlor de Oro,
W side of Rio Guapor6(= Rio Itenez), 22 Jun 1991 (d),
Nee 41225 (LPB, MO, NY), (9 fl-fr), Nee 41226 (K, LPB,
MO, NY, USZ); Prov. Velasco, Parque Nacional Noel
Kempff M., PampaToledo, 1 Jul 1993 (9 fl-fr), Saldias et
al. 2891 (USZ).
The leaves of Cecropia saxatilis resemble those of
C. obtusa, but the lateral veins in the free parts of the
leaf segments are branched, whereas they are mostly
unbranched in C. obtusa. The pistillate inflorescences
of the two species are also quite similar. However, the
staminate flowers are very different. Those of C. ob-
tusa have anthers of the most common type, but those
FLORANEOTROPICA
of C. saxatilis have anthersof the less usual type with
long anthersremainingafterabscissionattachedto the
filament by spiral thickenings, as found in C. gla-
ziovii, C. palmata, and C. sciadophylla. Anderson
9258 differs from all other collections by the rela-
tively narrowand subobovatefree partsof the lamina
segments in which the lateral veins are not much
branched.
Vernacular name. Brazil: imbadbabranca (Ba-
hia).
50. Cecropia schreberiana Miquel, in Martius,Fl.
Bras. 4(1): 150. 1853. Type. Probablyfrom one of
the Frenchislands of the Lesser Antilles, certainly
not from "Guyane"as indicatedon the label, (d),
Schreber s.n. (holotype: U; isotype: M, photo-
graphin F). Fig. 43
Tree, to 15 m tall. Leafy twigs 1.5-3 cm thick,
green,glabrous,sparselypuberulous(sometimesonly
on the stipularscars) or densely hirtellousto subhir-
sute to subvillousor to hispidulous;internodespartly
filled with brownpith. Laminasubcoriaceousto cor-
iaceous, sometimes ? plicate, ca. 25 X 25 cm to 50
X 50 cm (to 70 X 70 cm), the segments 8-10, the
free partsof the upperones ovate to elliptic to oblong,
the incisions down to 4/10-7/10; apices roundedto
subacute;uppersurface smooth and (sub)glabrousor
? scabrousand hispidulousto (densely) puberulous
to hirtellous,often also with sparseto dense arachnoid
indumentum; lower surface glabrous, with rather
dense brown pluricellulartrichomes or hirtellous to
subhirsuteon the veins, with arachnoidindumentum
in the areoles and on smaller veins, at least initially
often extendingto the main veins; lateralveins in the
free part of the midsegment 10-16 pairs, marginally
loop-connected, usually branched; petiole 10-35
(-45) cm long, glabrousor hirsuteto hirtellousor to
subvillous, often only at the base, often also with
sparse to dense arachnoidindumentum;trichilia ab-
sent, occasionallypresent,but then usuallypoorlyde-
veloped; stipules 7-20 cm long, green, glabrous or
sparselyto densely (sub)hirsuteto (sub)strigose,often
also with arachnoidindumentumoutside, sparselyto
ratherdensely (sub)sericeous(or subglabrous)inside.
Staminateinflorescencessolitary or in pairs, the pe-
duncle erect and the spikes erect to pendulous (?);
peduncle 4-10(-13) cm long, glabrous or hirtellous
to subhirsute,often also with arachnoidindumentum;
spathe4-8(-10) cm long, green or grayishgreen,gla-
brousor hirtellousto subhirsuteto strigose, often also
with arachnoidindumentumoutside, glabrousinside;
SYSTEMATIC TREATMENT 165

L~~~~~~~

FIG. 43. Cecropia schreberiana subsp. schreberiana. 1. Leafy twig with staminate inflorescences with spathes. 2.
Staminate inflorescence and part of spike (Wagner 566). 3. Staminate flower and stamen (Berg 1526). 4. Pistillate inflores-
cence at anthesis (Beard 180). 5. Pistillate flower and style. 6. Fruit (Berg 1533). C. schreberiana subsp. antillarum. 7.
Pistillate flower (Maas et al. 6420). (By T. Schipper and Hendrieke Berg.)
166
spikes ca. 10-20, 1.5-6(-9.5) X 0.2-0.3 cm, with sti-
pes 0.3-1.5 cm long and glabrous or hairy; rachis
hairy. Staminateflowers: perianthtubular,ca. 1-1.2
mm long, glabrous,the apex convex to almost plane;
filamentsflat;anthersca. 0.5 mm long, appendiculate,
detachedandreattachedto the marginsof the aperture
by the appendages at anthesis. Pistillate inflores-
cences often solitary, erect; peduncle 2-8(-1 1) cm
long, glabrous or hirtellous to subhirsute,often also
with arachnoidindumentum;spathe3-8 cm long, the
color and indumentumas in the staminateinflores-
cence; spikes 4-6, 1.5-6 X ca. 0.5 cm, to 13 X 1.3
cm in fruit, sessile or stipes to 0.4 cm long, glabrous
or hairy. Pistillate flowers: perianth ca. 1.5-2 mm
long, with arachnoidindumentumbelow the apex or
extending to near the apertureoutside, absent inside,
the apex slightly convex to plane, muriculateto min-
utely puberulousor smooth; style short;stigma ligu-
late to subpeltateto peltate. Fruit ellipsoid to oblon-
goid, ca. 1.5-2.5 mm long, tuberculate,darkbrown.
In the presence of ample brown pith in the inter-
nodes, the absent or poorly developed trichilia, and
the smooth leafy twigs and uppersurfaceof the lam-
ina (in subsp. schreberiana),this species shows sim-
ilarities to several Andean species. The presence of
peltate stigmas and arachnoid indumentumon the
apex of pistillateflowersin subsp.antillarumindicate
affinities to the C. peltata-group, in particularto C.
sararensis, which show parallel variation in the in-
dumentumand featuresof the staminateflowers.Sub-
juvenile specimens of C. schreberianacan hardlyor
cannot be distinguishedfrom material of C. peltata
without trichilia (from Jamaica).Even if trichiliaare
sometimes more or less developed, then they do not
containMullerianbodies (Rickson, 1977), or only ab-
normal(?) ones (Wheeler, 1942: 73). In many studies
on the genus in the Caribbean,e.g., those by Janzen
(1973) and Rickson (1977), this species was not re-
gardedas distinct from C. peltata, in spite of the fact
that Snethlage (1923) established several names for
this species and that Wheeler (1942) noted that the
Cecropia species in PuertoRico did not belong to C.
peltata. Two subspecies can be recognized.The tran-
sition of charactersis gradualand intermediatesoccur
in Hispaniola and Puerto Rico. Juvenile and subju-
venile specimens of the subspecies are quite similar.
Charactersof the subjuvenile state can be retained
until flowering starts.
Key to the subspecies
1. Laminawith the uppersurface smooth,
glabrous,or only with arachnoidindumentum;
lower surface with only arachnoidindumentum
FLORA NEOTROPICA
on the main veins, this often initially covering
the main veins.................. a. subsp. schreberiana
1. Laminawith the uppersurface ? scabrous,
hispidulousto hirtellous;lower surface
hirtellousto subhirsuteor often also with
arachnoidindumentumon the main veins
............................................ b. subsp. antillarum
50a. Cecropia schreberiana Miquel subsp. schre-
beriana Fig. 43.1-6
Cecropia urbaniana Snethlage, Notizbl. Bot. Gart.
Berlin-Dahlem 8: 366. 1923. Type. Guadeloupe.
Camp-Jacob,Matouba, 1896 (Y), Matouba, Duss
3621 (holotype:B, destroyed;isotypes: NY, P, US).
Leafy twigs (usually) glabrous or only with brown
arachnoid trichomes or white hairs at the stipular
scars. Lamina with the upper surface (usually)
smooth, glabrous or only with arachnoid indumen-
tum, lower surface only with arachnoid indumentum,
initially often extending to the main veins. Perianth
of the pistillate flower without arachnoid indumentum
at the apex; stigma ligulate to subpeltate.
Distribution (Fig. 18.3). Lesser Antilles, Puerto
Rico, and Hispaniola, in forest and secondary growth,
at elevations to 1100 m.
Representative specimens examined. HAITI. Tortue
Is., nr. La Vallee, 28 Dec 1928-9 Jan 1929 (?), Leonardet
al. 11496 (A, GH, US); betweenPlaisance& Limbe,28 Aug
1903 (5), Nash 889 (NY, U).
PUERTO RICO. CaribbeanNational Forest, Luquillo
Mtns., 27 Sep 1989 (5), AcevedoR. 2956 (NY, US); Maun-
abo, Cuchillade las Panduras,24 Feb 1991 (? fl-fr),Axelrod
et al. 2062 (UPRRP);Jayuya,Toro Negro Forest Reserve,
17 Jan 1992 (? fl-fr), Axelrodet al. 3772 (UPRRP);Carri-
bean NationalPark,El Junque,31 Mar 1985 (5), Berg 1526
(AAU, BG, U); between Arecibo & Rio AbajoForest,2 Apr
1985 (st), Berg 1527 (BG); Rio Abajo Forest, 2 Apr 1985
(Y fl-fr), Berg 1528 (BG); Marici Forest, 3 Apr 1985 (Y fl-
fr), Berg 1533 (BG, U); CaribbeanNational Forest, rd. to
summitof El Yunque,4 Jan 1987 (Y fl-fr),Boom 6894 (NY,
UPRRP); Sierra de Luquillo, between Mameyes & Juncos,
Sep-Oct 1966 (5), Byer 66-960 (UPRRP); Mun. Cayey,
Bosque Nacional Carite, 5 Apr 1992 (5), Escobar et al.
9935 (UPRRP);LuquilloNationalForest,El Yunque,1 Mar
1985 (5), Gentry et al. 50369 (BG, NY); Mun. Patillas,
Reserva Forestal Carite, 19 Nov 1981 (5), Hansen et al.
9158 (US); Rio Piedras, Finca Bueno Consejo, May 1914
(5), Bro. Hioram s.n. (NY, US); Cerro Maravillaarea, 30
Jan1979 ( fl), Liogier et al. 28274 (NY, US); Quebradillas,
26 Feb 1986 (st), Liogier et al. 35988 (NY); CaribbeanNa-
tional Forest, 25 Jun 1950 (5), Little 13103 (NY, US), (9
fr), Little 13014 (US); Las Piedras,22 May 1989 (9 fl), Loiz
et al. 37 (UPRRP);Mun. Cayey, rd. to Barrio Farrall6n,9
Feb 1991 (5), G. S. Miller et al. 6000 (UPRRP);Mun. Rio
Grande, rd. to El Verde, 9 Dec 1988 (st), Ortega 128
SYSTEMATICTREATMENT
(UPRRP);Mun. Caguas, Sierrade Cacey, CariteForestRe-
serve, 2 Feb 1986 (6), Proctor 41467 (IJ);MaricoaInsular
Forest, 25 Jun 1965 (9 fl), Stimson 1337 (MICH); Mun.
Orocovis, rd. 564, km 7.3, 18 Aug 1966 (9 fl-fr), Stimson
4057 (MICH);Mun. Aibonito, rd. 162, between rd. 1 & rd.
716, 8 Apr 1989 (9 fl-fr), Tayloret al. 8823 (UPRRP);Sa-
bana rd., nr. Palmer,2 Feb 1968 (9 fl-fr), Wagner1321 (A,
U) Barranquitas,30 Jul 1979 (9 fl-fr), Woodburyet al. s.n.
(NY, US); El Yunque,23 Aug 1979 (9 fl-fr + juv), Wood-
bury s.n. (NY, US).
MONTSERRAT. Olveston area, 19 Mar-16 Apr 1979
(9 fl-fr), Howard et al. 18982 (A, BM, NY, US); Gorge of
Gage's, Soufriere stream, 12 Feb 1959 (6), Proctor 19153
(A, BM, IJ, US); FergusMtns., 30 Jan 1907 (6), Shafer342
(NY, U, US).
SABA. The Mountain,24 Jul 1906 (st), Boldingh 2158
(U); top of The Mountain, 11 Aug 1953 (st), Stoffers4230
(A, K, NY, U).
ST. EUSTATIUS. The Quill, 7 Jun 1906 (st), Boldingh
254 (U); Bottom of the Quill, 17 Jun 1906 (st), Boldingh
904 (U), 21 Jun 1906 (st), Boldingh 955 (U).
ST. KITT'S. MolyneauxEstate, 8 Sep-5 Oct 1901 (st),
Britton et al. 572 (NY).
GUADELOUPE. Bains-Jaunes, 1892-1893 (? fl-fr),
Duss 2860 (NY); Bains-Jaunes,Mar 1905 (st), Duss 4207
(NY); Marie Galante, Les Balisiers distr., 1 Jun 1960 (d),
Proctor 21135 (A).
DOMINICA. Morne Colla Anglais, 10-23 Aug 1938
(st), Hodge 632 (GH, NY, U, US); nr. Calibishie, 23 Apr
1940 (9 fl-fr), Hodge et al. 3168 (GH); La Chaudiere,10-
14 Mar 1940 (? fl-fr), Hodge et al. 3663 (BM, GH, US);
St. Joseph Parish, Layou Park Estate, 22 Nov 1964 (9 fl),
Nicholson 2037 (BM, GH, U, US); Laudat,24 Apr 1988 (9
fl-fr), Ramages.n. (BM, K); withoutlocality,Mar 1942 (d),
D. Taylor125 (GH); rd. to Trou Cochon, 9 Oct 1983 (6),
Whitefoord3958 (BM, US); Baiac, 18 Sep 1984 (d), White-
foord 4135 (A, BM, NY, US).
MARTINIQUE. Mome-Rouge, Fonds St. Denis, 1883,
Duss 1405 (NY, US); Bois de Caleluze (?), Dec 1867 (st),
Hahn 210 (BM, G, K); La Chapelle, Apr 1873 (? fl-fr),
Hahn s.n. (G); Pitons Boucher, Lacroix et Dumauze, ca.
1000 m, 13 Apr 1968 (6), Oldemanet al. M.58 (P); Morne
Mitan,4 Apr 1980 (st), Sastre 6912 (U); Piton Dumauze, 11
Apr 1980 (9 fl-fr), Sastre 6922 (U); La Soufriere,between
Baines Jaunes & Savane a Mulets, 1000-1100 m, 17 May
1980 (st), Sastre 6939 (U).
ST. VINCENT. Lodge R., Jan 1890 (d), Eggers 6829
(A, P, US); ChateaulairR., 16-25 Apr 1947 (st), Morton
5419 (US); CumberlandR., 2-3 May 1947 (st), Morton5571
(US); Parish of St. Andrew,Upper BuccamentR. valley, 7
Mar 1965 (6), Proctor 26152 (IJ).
ST. LUCIA. Quiless, 14 Dec 1943 (6), Beard 180 (A,
K, NY, U, US); SE of Piton Troumass6e,11-12 May 1958
(? fl-fr), Proctor 17949 (A, BM, IJ, US); Piton Flore, (d
+ 9 fl-fr), Sturrock714 (A).
Brokaw (1998) presented an account of this sub-
species, particularly in relation to its role as a colo-
nizer, e.g., after hurricanes.
167
Vernacular names. Dominica and St. Lucia:bois
canot. Montserrat:trumpetertree. St. Vincent:trum-
pet. FrenchAntilles: bois trompette.
50b. Cecropia schreberiana Miquel subsp. antil-
larum (Snethlage)C. C. Berg & P. Franco,comb.
et stat. nov. Fig. 43.7
Cecropia antillarum Snethlage,Notizbl. Bot. Gart.
Berlin-Dahlem 8: 364. 1923.Type.DominicanRe-
public.Nr. Barahona,Jun 1910 (i), Fuertes 78 (ho-
lotype:B, destroyed;isotypes:BM,G, K, NY,U).
Cecropia sericea Snethlage,Notizbl.Bot.Gart.Berlin-
Dahlem 8: 368. 1923. Type. Haiti. Nr. Terreneufe,
Buch 99 (holotype:B, destroyed).
Leafy twigs hirtellous to hirsute to subvillous or
to hispidulous. Lamina with the upper surface +
scabrous,hispidulousto hirtellous;lower surfacehir-
tellous to subhirsuteon the main veins and initially
often also with arachnoid indumentum. Perianth of
the pistillate flower with arachnoidhairs on the apex
to near the aperture;stigma subpeltateto peltate (or
ligulate).
Distribution (Fig. 18.3). Cuba,Hispaniola,Puerto
Rico, and the Virgin Islands, in forest and secondary
growth, at low elevations.
Representativespecimensexamined.CUBA.CAMA-
GUEY: Nr. La Gloria, 8 Feb 1909 (st), Shafer325 (NY, US).
LAS VILLAS: Trinidad
Mtns.,TopedeCollantes,17Jul1957
(9 fl-fr), (Bro. Alain) Liogier 6432 (US), (i), (Bro. Alain)
Liogier 6433 (US). MATANZAS: Valley of the St. Augustine
R., 2 Sep 1903 (9 fr), Brittonet al. 258 (NY); San Miquel
de los Banios,Apr 1951 (9 fl), Dahlgren 51/102 (US); be-
tween Mantanzas& Varadero,21 Jun 1988 (9 fl), Fernandez
C. et al. 10690 (NY). ORIENTE: Bayate, nr. Rio Jagua, 5
Nov 1915 (st), Ekman6548 (NY, S); Los Pifiales,SE of Paso
Estancia, 1-2 May 1909 (9 fl-fr), Shafer1726 (A, NY, US);
Monteverde,Jul 1859 (9 fl-fr), Wright1440 (G, GH, K, P,
S). PINAR DEL Rio: Nueva Gerona,May 1904 (6), Curtiss
s.n. (NY); Vinales, 11 Dec 1930 (9 fr), Killip 13571 (US);
nr. Cabafias,20 May 1900 (st), Palmer et al. 768 (US); nr.
Pinar del Rio, 19 May 1941 (9), Rutten-Pekelharing 436
(U). SANTA CLARA: Cienfuegos Distr.,Cieneguita, 23 Mar
1895 (9 fl-fr), Combs 111 (GH, NY, P); Cienfuegos, Sole-
dad, 30 Mar 1926 (9 fl), Jack 4485 (S); Cienfuegos, Sole-
dad, Limones, 11 Aug 1927 (6), Jack 5271 (P, US); Cien-
fuegos, Soledad, 8 Oct 1927 (9 fl-fr), Jack 5558 (S);
Cienfuegos, Soledad, Belmonte, 17 Jul 1929 (9 fl-fr), Jack
7481 (A, NY, P, US).
HAITI. Massif de la Selle, Papette,28 Nov 1927 (9 fl-
fr), Ekman9376 (NY, S, US); Soliette R., Fonds Verrettes,
21 Mar 1942 (6), Holdridge 1061 (MICH,NY, US); nr.
Petionville, 15-29 Jun 1920 (9 fl-fr), Leonard4875 (NY,
US); nr. St. Michel de I'Atalaye, 3 Jan 1926 (9 fl-fr), Leon-
ard 8532 (NY, US); Dept. du Nord, nr. Dondon, 7 Jan 1926
(6), Leonard8655 (US); Dept. de l'Artibonite,18 Feb 1926
168
(st), Leonard9990 (US), 23 Feb 1926 (d), Leonard10026b
(GH, US); TortueIs., nr. La Vallee, 28 Dec-9 Jan 1929 (9
fr), Leonard 11496 (A, GH); nr. Basse Terre,23 Mar 1929
(d6),Leonardet al. 13998 (US), 21-29 Mar 1929 (9 fl-fr),
Leonardet al. 14084 (A, GH, NY, US); Dept. Sud, Bouzy,
Dabo, 20 Jun 1980 (st, juv), Sastre et al. 6950 (P).
DOMINICAN REPUBLIC. BARAHONA:3-4 kmN of
Paraiso, on rd. to Las Auyamas, nr. Nizaito, 11 Apr 1985
(? fl), Gentry et al. 50717 (BG, MO, NY); El Arroyo, nr.
Barahona,25 Mar 1985 (d), Maas et al. 6414 (BG, U); La
Filipina, 6.3 km from Ocean Hwy., 25 Mar 1985 (9 fl),
Maas et al. 6420 (BG,U). DISTIUTo NACIONAL:SantoDo-
mingo, 30 Apr 1929 (9 fl-fr), Ekman 12324 (GH, S, US);
Santo Domingo, JardinBotdnico Nacional, 2 Jun 1980 (9
fl-fr), Mejfa et al. 6732 (NY), (6), Mejia et al. 6733 (NY);
SantoDomingo, 24-25 Mar 1923 (6), Rose et al. 4138 (NY,
US), 8 Jul 1981 (9 fl-fr),Zanoniet al. 15282 (NY). MONTE
CRISTI: SabanetaDistr., La Ceiba, 17 Oct 1930 (? fl-fr),
Valeur480 (C, G, MICH,US). MONTE PLATA: Rd. Ya-
masa-Maim6n, Rio Verde,28 Mar 1987 (6), Zanoni et al.
38796 (NY). SAN CRISTOBAL: Rd. CambitaGarabita (de
San Crist6bal)-ArroyoMarfa,4 Oct 1987 (6), Zanoniet al.
40487 (NY).
PUERTO RICO. Yabucoa, Punta Guayanes, old rd.
from S end of Palmas del Mar, 12 May 1991 (9 fl-fr), Ax-
elrod et al. 2427 (NY, UPRRP), 18 Aug 1991 (9 fl-fr),Ax-
elrod et al. 2787 (NY, UPRRP);Vieques, Monte Pirata,23
Jul 1992 (9 fl), Axelrod et al. 4921 (NY, UPRRP); Mun.
Isabela, Bo. Arenales Altos, 8 Nov 1993 (9), Nee 44158
(NY); Bayam6n 31 Mar 1885 (d), Sintenis 250b (G, GH,
K, NY, P, S, US); nr. San. German,9 Mar 1886 (9 fl-fr),
Sintenis 3971 (BM, G, GH, K, NY, P); nr. Penluelas,Talba-
boa, 8 Aug 1886 (6), Sintenis 4805 (BM, G, GH, K, NY, P,
S, US); Vieques Is., between Isabel Segunda & Martineau,
31 Jan 1914 (d), Shafer2643 (NY, US); S of Cayay, 15 Mar
1932 (6), J. S. Miller 1604 (US).
ST. CROIX. CaledoniaGut, 8 Jul 1873 (9 fl-fr),Eggers
s.n. (C); Jolly Hill, 29 Jan1906, Raunkiaers.n. (C); Lebanon
Hill, 24 Jun 1896 (9 fl-fr), Ricksecker449 (NY); without
locality, (9), Wests.n. (C).
ST. JOHN. Cruz Bay, Center Line rd., 1 Jul 1989 (9
fl), Acevedo 2845 (NY, UPRRP,US).
TORTOLA. Rd. to High Bush, 13-17 Feb 1913 (6),
Brittonet al. 715 (NY, US).
Due to the (sub)peltatestigmas, the white arach-
noid indumentumalso largely covering the apex of
the perianthpistillateflower,and some featuresof the
indumentumof the vegetativeparts,materialreferred
to this subspecieslooks quite similarto Cecropiapel-
tata without trichilia. This applies even more so to
material from the Virgin Islands and Vieques Island,
as the lamina has a less coriaceous texture than else-
where. Cecropia schreberiana subsp. antillarum has
been introducedin Madagascarand known underthe
vernacularname "Taigisy"or "Taingisa."According
to Capuron23568 (P) the (sub)species is planted in
(Est) Faragangana.Materialcollected by Capuron(15
FLORA NEOTROPICA
Oct 1964) has staminateinflorescences.Anothercol-
lection in P, collected by RakotoariveloAndrianjar-
afeno (undernumber459, 20 Oct 1961) is made near
Vangaindranahas pistillate inflorescences.The label
data of the latter collection suggest that this
(sub)species is becoming naturalizedin Madagascar.
51. Cecropia sciadophylla Martius,Flora 24, Beibl.
2: 93. 1841; Velasquez,Acta Bot. Venez. 6: 60, t.
14; Berg, Acta Amazonica 8(2): 181. 1978. Am-
baiba sciadophylla (Martius)Kuntze,Revis. Gen.
P1.2: 624. 1891. Type. Colombia.Amazonas:Rfo
Caqueta',Puerto Mirania,Martius 630(1) (holo-
type: M, fragmentin U).
CecropiajuranyianaV. Richter,Biblioth.Bot. 43: 13.
1897. Cecropiasciadophyllavar.juranyiana(V.
Richter) Snethiage, Notizbl. Bot. Gart. Berlin-
Dahlem 8: 358. 1923. Type. French Guiana. Acar-
ouany, 1856 (i), Sagot 861 (holotype: P; isotype:
BM,K).
Cecropiasciadophyllavar.decurrensSnethlage,No-
tizbl. Bot. Gart.Berlin-Dahlem8: 358. 1923. Type.
Brazil. Amazonas: Nr. Manaus (?), Ule 5512 (syn-
type: B, destroyed) and Guyana.Barakara,nr. Kar-
tabo, 15-18 Jul 1920 (st), nr. Kartabo,Bailey 12
(syntype:B, destroyed;GH).
Cecropiasciadophylla var.pedroa Cuatrecasas,Revista
Acad. Colomb.Ci. Exact. 6(22/23): 299. 1945. Type.
Colombia. Putumayo: Between Umbria & Puerto
Asfs, 10 Nov 1940 (d), Cuatrecasas 10551 (holo-
type: COL).
Cecropiasciadophyllavar.guamuesensis
Cuatrecasas,
Revista Acad. Colomb. Ci. Exact. 6(22/23): 299.
1945. Type. Colombia. Putumayo: Rfo Guamues,
San Antonio, 19 Dec 1940 (6), Cuatrecasas11203
(holotype:COL).
Cecropiasciadophylla Re-
var.subsessilisCuatrecasas,
vista Acad. Colomb. Ci. Exact. 6(22/23): 299. 1945.
Type. Colombia. Caqueta:Sucre, 4 Apr 11940 (6),
Cuatrecasas9106 (holotype:COL).
Cecropia inchuensis Cuatrecasas,Revista Acad. Col-
omb. Ci. Exact. 9(36/37): 329. 1956. Type. Peru:
Ucayali: Prov.CoronelPortillo,Inchua,2 Aug 1946,
Soukup3018(F,mixedcollectionconsistingof a pis-
tillate inflorescenceof uncertainidentity[possibly C.
ficifolia,see Soukup3017] andseparateleaf parts,
the latterelement(s) designatedas lectotype, Berg &
FrancoRosselli, Fl. Ecuador48: 48. 1993, F).
Tree,to 30 m tall. Leafytwigs 1.5-5 cm thick,
green to brownish, glabrous or puberulous.Lamina
coriaceous, ca. 20 X 20 cm to 100 x 100 cm, the
segments (7-)10-15, lanceolate, the incisions down
to the petiole andthe segmentsusuallywith petiolules
1-7 cm long; apices acuminateto acute;uppersurface
SYSTEMATICTREATMENT 169

smooth and glabrous; lower surface with sparse Barriga 14012 (COL, US), 20 Aug 1951 (9 fr), Schulteset
brown pluricellularhairs on the (main) veins, with al. 13679 (COL, U, US), 15 Sep 1951 (6), Schultes et al.
short and sparse arachnoidindumentumin the are- 13990 (COL, GH, U, US). CAQUETA: Mun. Florencia,rd.
to Gabinete, 17 Oct 1993 (9 fl-fr), Franco et al. 4507 (BG,
oles; lateralveins in the midsegment(25-)30-40(-45)
COL); nr. Las Guacamayas,27 Apr 1944 (9 fl-fr), Little
pairs, submarginally loop-connected, unbranched;
7749 (US), (st), Little 7750B (US); Solano, Rfo Caqueta,8
petiole ca. 20-90 cm long, glabrous;trichiliaabsent, km SE of Tres Esquinas,below mouthof Rfo Ortequeza,13
sometimes a few long white hairs insteadof trichilia; Mar 1945 (6), Little et al. 9785 (COL, GH, NY, US); Ar-
stipules 13-50 cm long, red-brown(or greenish),gla- aracuara,25 Jan 1993 (9 fl-fr), Vesteret al. 725 (BG). CAS-
brous or sparsely puberulous outside, (densely) to- ANARE: Hato Corozal, Vrda. Las Tapias, 30 Oct 1981 (st),
mentose to subsericeous inside. Staminate inflores- Mahecha et al. 3572 (UDBC). META:Mun. PuertoL6pez,
cences in pairs, erect; peduncle 7-15 cm long, rd. to Puerto Gaitan, 10 Feb 1995 (d fl-fr), Franco et al.
minutely puberulous;spathe 8-18 cm, red-brown(or 4659 (BG, COL, HUA, MO, US); nr. La Macarena,11 Feb
greenish to yellowish), puberulousoutside, glabrous 1995 (9 fl-fr), Franco et al. 4666 (BG, COL, HUA, MO,
US); Cord.La Macarena,between Rio Guejar& CanloGua-
or with sparse arachnoidindumentuminside; spikes
payita,CanloYerli,20-28 Dec 1950 ( 9 fr), Idroboet al. 792
ca. 8-15, 7-12 X 0.3-0.8 cm, with stipes to 4 cm (COL, GH, U, US); Mun. Puerto L6pez, SE of Cabuyaro,
long and subglabrous;rachis hairy. Staminateflow- nr. Laguna de Yurimena, 16 Sep 1958 (6), Jaramillo M.
ers: perianthtubular,ca. 1.5-2.5 mm long, with dense 1237 (COL); Serraniade Menegua, 17 Sep 1958 (6), Jar-
arachnoid indumentum below the apex, the apex amillo M. et al. 1261 (COL, NY); Mun. San Juan Arama,
slightly convex to plane muriculate; filaments ? Rio Siejas, 1971 (9 fr), Mahecha s.n. (UDBC 8093); Mun.
swollen, sometimes (?) connate at the base; anthers Vista Hermosa, cacerfo Pifialito, Nov 1973 (st), Mosquera
ca. 1-1.5 mm long, appendiculate,detached,remain- s.n. (UDBC 10855); Sierrade La Macarena,SerraniaCha-
musa, ParqueNacional NaturalTinigua,Jan 1991 (6), Ste-
ing attachedto the filamentby a bundle of stretched
venson 226 (COL). PUTUMAYO: Mocoa, Jul 1989 (st), Ma-
spiral thickenings of trachearyelements at anthesis.
hecha et al. 5779 (UDBC). VAUPES:Miti, 10 Sep 1956 (9
Pistillate inflorescencesin pairs or solitary, erect to fr), A. S. Barclay et al. 623 (COL, GH, NY); nr. Miraflores,
pendulous;peduncle 3-10 cm long, minutely puber- 8 Feb 1944 (9 fr), Gutierrez V et al. 769 (COL, GH,
ulous; spathe 8-12 cm long, the color and indumen- MEDEL).
tum as in the staminate inflorescence; spikes 3-6 VENEZUELA. AMAZONAS: Alto Orinoco, Raudal de
(-10), (2-)5-10 X 0.8-1 cm, to 26 X 1.5(-2.5) cm los Guaharibos,Salto Salas, 18 Aug 1951 (6), Croizat501
in fruit, (sub)sessile or with stipes to 0.5 cm long and (F, NY, US); Rio Orinoco, 5 Jan 1952 (st), Cruxent 158
glabrous; rachis hairy with stiff short hairs and/or (NY); Dtto. Atabapo,Rio Ventuari,Salto Yureba,24 Oct-4
Nov 1981 (9 fl-fr), Delascio et al. 11050 (VEN); Rio Ca-
arachnoid indumentum.Pistillate flowers: perianth
siquiare,RIo Padamo,Jan-Feb 1969 (st), Fariniaset al. 689
ca. 2-3 mm long, with arachnoidindumentumbelow
(MO, NY, VEN); 14 km NE of San Carlos de Rio Negro,
the apex outside, also in the lower part of the style 24 Jan 1980 (9 fr), Liesner 8664 (MO, NY, VEN); Tayari,
channel(or absent?)inside, the apex convex to almost 7 May 1975 (9 fl-fr), Lissot 75/35 (VEN); San Carlos de
plane, muriculate; style rather long, straight, with Rio Negro, 17-18 Apr 1970 (6), Steyermarket al. 102718
short hairs;stigma penicillate.Fruit oblongoid to su- (NY, US, VEN). BOLiVAR:Uairen region, ApradaTepui,
bobovoid, 2-3 mm long, tuberculate. 1200 m, 19 Aug 1953 (d), Bernardi 826 (NY); rd. Pica
Caicaradel Orinoco-San Juande Manapiare,Rio Suapure,
Distribution (Fig. 11.1). Amazon basin, the Gui- Mar 1975 (9 fl-fr), Delascio et al. 2782 (VEN); Dtto. Ced-
ana region (FrenchGuianato easternVenezuela),and eno, Rio Tabaro,Research Station Dedmai, 11 Aug 1992
the llanos region in Colombia.In primaryuplandand (st), Knab et al. 196 (MICH);Rio Caura,rd. Campamento
gallery forest and in secondary growth, in non- Las Pavas-El Play6n, 11 May 1982 (6'), Morillo et al. 9154
inundatedplaces, at elevations to ca. 1300 m. (MO, VEN); Rio Canaracuni,13-26 Apr 1988 (9 fl-fr),
Stergios 11784 (BG, NY); Quebrada O-paru-ma, below
Representativespecimensexamined.COLOMBIA. SantaTeresitade Kavanayen,915-1065 m, 25 Nov 1944 (9
AMAZONAS: Rfo Igara-Parand,La Chorrera,9 May 1975 fr), Steyermark60540 (F, NY, VEN); Chimanta Massif,
(st), Idrobo 8086 (COL); Santa Izabel, Reserva Indfgena Chimanta-tepui,above Rfo Tirica, 1000-1400 m, 15 May
Miratia,5 Nov 1984 (Y fr), La Rottaet al. 543 (COL);Mun. 1953 (9 fr), Steyermark75387 (F, NY,VEN); nr.Venezuela/
Leticia, ParqueNacional NaturalAmacayacu,nr. Quebrada Brazil border,NE of SerraniaPiua-soi, 5-6 Jan 1962 (6),
de Agua Pudre, 21-24 Mar 1992 (st), Rudas et al. 3558 Steyermark90614 (US); rd. El Dorado-Brazilianborder,km
(COL, MO); Rio Igara-Parana,nr. La Chorrera,4-10 Jun 88, Sep 1967 (9 fr), Velasquez258 (US, VEN); nr. El Pal-
1942 (st), Schultes 3964 (F, G); Trapecio Amaz6nico, Rio mar,25 Nov 1967 (9 fl-fr), WesselsBoer 2102 (U). DELTA
Loretoyacu, Oct 1946 (Y fl-fr), Schultes 8446 (F). AMA- AMACURO: Rio Grande,ENE of El Palmar,nr.Bolivarbor-
ZONAS/VAUPES: Rio Apaporis, between Rio Pacoa & Rio der, 19 Feb 1964 (9 fl-fr), MarcanoBerti 89 (GH, K, NY,
Kananari, Soratama, 1-15 Dec 1951 (Y fl-fr), Garcia- P, U, VEN); Sierra Imataca,Rimacuro,upstreamfrom San
170
Victor,past Rio Matapaima,up to Salto Quebradero,2 Nov
1960 (5), Steyermark87270 (BG, NY).
GUYANA. Northwest Distr., nr. Port Kaituma, Aug
1967 (st), Davis 247 (NY); EssequiboR., Rockstone, 15 Jul-
1 Aug 1921 (st), Gleason 663 (GH, K, NY, US); Mazaruni
Station, 19 Jun 1942 (Y fl), Fanshawe 731 FD 3467 (K);
NorthwestDistr.,WaunaMission, 11 May 1961 (5), Harris
Y29 (K); DemeraraR., Kamini R., PokoreroCr., Santa, 21
Apr 1923 (st), Hohenkerk115A (K); EssequiboR., Gunn's,
8 Aug 1989 (? fl-fr), Jansen-Jacobset al. 1542 (BG, NY,
U); PakaraimaMtns., N of ParuimaMission, 19 Oct 1981
(9 fl-fr), Maas et al. 5859 (K, NY, U); Boerasirie area, 5-
25 mi W of Georgetown,25 Oct 1955 (9 fr), Little 16954
(US); Soesdyke Hwy., 7 Nov 1982 (9 fl), Persaud243 (U);
MarudiMtns., nr. Aishalton, 12 Nov 1982 (9 fl-fr), Stoffers
et al. 317 (INPA, NY, U).
SURINAME. Sectie 0 (tree 531), 20 Oct 1915 (9 fl),
BW s.n. (U, US); S of Onverwacht,W of Paramaribo-Zan-
derij rd., 3 Jul 1977 (st), Heyde 710 (U); WilhelminaMtns.,
Lucie R., below confluence with Oost R., 10 Sep 1963 (5),
Irwin et al. 55577 (COL, K, MG, MO, NY, S, U, US); Cop-
penameR., nr.Raleigh Falls, 13 Sep 1933 (st), Lanjouw824
(RB, U); Mt. Nassau, 7 Mar 1949 (st), Lanjouwet al. 2485
(U); Lely Mtns., 29 Sep 1975 (st), Lindeman& Stofferset
al. 549 (K, NY, U); Mapane region, Jul 1970 (d), Roberts
(LBB) 12791 (U); between Lucie R. & WilhelminaMtns.,
17 Jul 1964 (5), Schulz (LPB) 10109 (COL, U); Zanderij,
Jan 1943 (3 fl-fr), Stahel (Woodherb.Suriname)179 (A, K,
MO, NY, S, U).
FRENCH GUIANA. Rd. St. Laurent-Cayenne,km 11,
1 Aug 1953 (st), BAFOG39M (U); Crique Margotregion,
9 Oct 1956 (st), BAFOG7561 (NY, P, U); Charvein,23 Dec
1913 (st), Benoist 442 (P); Mont Chauve,24 Apr 1997 (5),
Cremers et al. 15193 (BG); Taluwen, 28 Jan 1997 (st),
Fleury 1065 (BG); Lower Oyapock R., ArmontaboCr., 24
Feb 1981 (st), Granville4397 (P, U); Bassin de l'Oyapock,
Roche Touatou,20 May 1995 (5), Granville et al. 12995
(BG); Piste de St. Elie, Station ECEREX, 2 Nov 1990 (9
fl-fr), Hoff 6866 (BG); Crique Plomb, 12 Jul 1992 (st),
Loubry 1766 (CAY);Sail, 8 Nov 1990 (9 fl-fr), Mori et al.
21555 (NY); piste de St. Elie, 21 Sep 1979 (9 fl-fr),Prevost
801 (P, U), 7 Nov 1984 (9 fl-fr), Riera 905 (BG); basin of
ApprouageR., NoutaguesStation,11 Nov 1993 (9 fl), Riera
et al. 1943 (US).
ECUADOR. MORONA-SANTIAGO:PuertoMorona,Rfo
Morona, 30 Sep-2 Oct 1975 (9 fl-fr), Little et al. 572
(LOJA, Q); Cordillerade Cutucu, 5-10 km E of Logrofno,
1200-1500 m, 7-9 Oct 1975 (5), Little et al. 654 (LOJA,
QAME, QCNE); 56 km SE of Taisha,23-25 Sep 1976 (st),
Ortega U. 205 (Q); Pozo petroleroGarza,ca. 35 km NE of
Montalvo, 2-12 Jul 1989 (9 fl), Zak et al. 4756 (MO,
QCNE). NAPO: Tena, 13 Oct 1939 (9 fl-fr), Asplund9296
(S), 18 Oct 1939 (5), Asplund 9419 (S); ParqueNacional
Yasunf,Maxus rd., km 40, 19 Sep 1994 (st), Aulestia et al.
2826 (QCNE);ParqueNacionalYasunf,Pozo petroleroAmo
2, 14 Jan 1988 (9 fl), Ceron3332 (AAU, BG, MO, QAME);
Parque Nacional Yasunf, Pozo petrolero Daimi 2, 14 Jan
1988 (9 fl), Cer6net al. 3860 (BG, QCNE);rd. Auca-Coca,
km 22, 30 Apr 1982 (st), Chaguaro36 (QCA, QCNE);con-
FLORA NEOTROPICA
fluence of Rio Quiwado & Rio Tiwaeno, 11 Apr 1981 (9
fr), Davis et al. 921 (QCA, U); San Jose de Payamino, 20
Apr 1982 (st), Irvine 103 (F, QCA, QCNE); La Joya de los
Sachas, Maxus rd., km 1-5, 1-28 Sep 1992 (9 fl-fr), Gri-
jalva et al. 68 (NY, QCNE); CampamentoCosapo, 2 Aug
1975 (5l), Little et al. 19 (Q, QAME, QCNE);Rio Napo, 15
km W of Coca, 18-20 Apr 1985 (st), Neill et al. 6335 (BG,
QAME, QCNE); ReservaBiol6gica JatunSacha, 8 km E of
Misahualli, 1-15 Sep 1987 (9 fr), Palacios 1890 (BG, GB,
K, MO, NY, QAME, QCNE);Aniangu,ParqueNacionalYa-
suni, 30 May-21 Jun 1982 (5), SEF 8644 (AAU, MO, NY,
QCA), (9), SEF 9227 (AAU, MO, NY, QCA). PASTAZA:
Cant6nPastaza,Pozo petroleroDanta2, 50 km SSE of Cur-
aray, 1-20 Oct 1990 (9 fl-fr), Espinozaet al. 379 (BG, GB,
NY, QCNE);Cant6nPastaza,Pozo PetroleroNamoyacu,30
km S of Curaray,13-30 Nov 1990 (9 fr), Espinoza et al.
585 (BG, MO); Cant6nArajuno,25 km NW of Pozo Villano
2, 3-14 Sep 1998 (9 fl), E. Freire et al. 3260 (BG). Suc-
UMBiOS: Rio Cuyabeno,ca. 0O10'S,75?55'W, 19 Feb 1980
(st), Berg et al. 1067 (BG, MO, QCA, U); Cant6nGonzalo
Pizarro,30 km NW of Lago Agrio, 1050 m, 31 Mar 1990
(st), Cer6n et al. 9417 (BG); Sacha Lodge, 3 km NW of
Aniangu,5-13 Jun 1995 (5), J. L. Clarket al. 1004 (QCNE);
Rio Aguarico,between Dureno& Poblaci6nCofanes, 10 Jul
1980 (9 fl), Jaramillo et al. 3001 (AAU, QCA); Reserva
FaunisticaCuyabeno, nr. Laguna Grande, 1 Apr 1990 (st),
Valenciaet al. 67779 (QCA). ZAMORA-CHINCHIPE:27 km
NE of Zamora,1200-1500 m, 17 Sep 1975 (5), Littleet al.
430 (COL, LOJA,Q, QAME, QCNE).
PERU. AMAZONAS: Rio Cenepa, nr. Huampami,ca. 5
km E of Chavez Valdivia, 11 Jul 1978 (5), Berlin 2041
(MO);Rio Santiagovalley, 65 km N of Pinglo, nr.Caterpiza,
20 Nov 1979 (9 fl-fr), Huashikat1358 (MO); Rio Cenepa,
QuebradaYutuientsa, 22 Jan 1973 (9 fr), Kayap 213 (GH,
MO, USM); Prov. Bagua, Dtto. ImazamRio Marafi6n,Ya-
mayakat,Feb 1995 (5), R. Vdsquezet al. 19527 (MO), (st),
R. Vasquezet al. 19807 (MO). Cuzco: Prov.Paucartambo,
rd. Pilcopata-Patria,ca. km 5, 1 Aug 1988 (9 fl-fr), Berg
et al. 1621 (BG, Centrode MedicinaAndina,Cuzco); Prov.
Paucartambo,between Atalaya & Pilcopata, 16 Nov 1964
(9 fl-fr), VargasC. 15760 (US). HUANUCO: Tingo Maria,
23 Aug 1940 (9 fr), Asplund 13311 (S); Prov.Pachitea,ca.
26 km S of Puerto Inca, 3 Nov 1988 (9 fl), Johann 1/3-
31188 (BG); Prov. Leoncio Prado, Dtto. Jose Crespo, Cas-
tillo, CarreteraMarginalde la Selva, km 11, Aucayacu, 29
Mar 1968 (st), Rodriguezdel A. 23 (MOL);Prov. Pachitea,
ca. 26 km S of PuertoInca, 7 Oct 1988 (st), Wallnoefer19-
71088 (BG); Prov. Puero Inca, Dtto. Yuyapichis,DANTAS,
16-31 Oct 1990 (d), Tello 506 (G, NY), 16-31 Apr 1991
(9 fl), Tello 1841 (G, NY). JUNiN: Puerto Bermddez, 14-
17 Jul 1929 (5), Killip et al. 26499 (NY, US). LORETO:
Rio Corrientes,at Ecuadorborder,4 Apr 1977 (9 fr), Gen-
try et al. 18998 (F, MO); Prov.Requena,JenaroHerrera,23
Feb 1987 (st), Gentry et al. 56424 (MO); Prov. Alto Ama-
zonas, Puranchim,Rio Sinchiyacu,30 Mar-I Apr 1987 (d),
Lewis et al. 13413 (MO); Prov. Maynas, Rio Ampiyacu,
Puca Urquillo, 5 Apr 1977 (9 fr), Plowman et al. 6664 (F,
GH, K, US); nr. Indiana, Rio Napo, 1924 (5), Tessmann
3718 (G, NY, US); mouth of Rio Santiago, 1924 (5), Tess-
SYSTEMATICTREATMENT
mann 4717 (G, F, NY); Prov. Alto Amazonas, Capahuari
Norte, 7 Jun 1981 (5), R. Vasquezet al. 1995 (BG, MO);
Prov. Maynas, Mishana, Rio Nanay, 7 Oct 1982 (Y fr), R.
Vasquez et al. 3276 (BG, F, MO); Prov. Maynas, Puerto
Almendras,2 Jan 1985 (Y fl-fr),R. Vdsquezet al. 6074 (BG,
MO); Rio Amazonas, Caballo-Cocha,6 Aug 1929 (st), Ll.
Williams2130 (F, NY, US). MADRE DE DIos: ParqueNa-
cional Manu, Cocha Cashu Station, 7 Nov 1986 (st), Foster
et al. 12189 (BG, LPB, MOL);Prov.Tambopata,jct. of Rio
La Torre& Rio Tambopata,26 May 1987 (Y fl-fr), Gentry
et al. 57660 (BG, MO, NY); Prov. Manu, ParqueNacional
Manu,Cocha Cashu,20 Jul 1986 (Y fl-fr),Nuinez5461 (BG,
MO); Prov. Tambopata,Reserva Tambopata,14 Aug 1990
(6), Reynel et al. 5131 (BG, MO). PASCO:Pichis valley,
Puerto Bermudez, Dec 1980 (5), Foster et al. 8035 (U);
Prov. Oxapampa,Iscozacin, 15 Jun 1982 (5), D. N. Smith
1875 (MO, MOL, NY, USM); Prov.Oxapampa,Palcazuval-
ley, Cabeza de Mono, 5-6 km W of Iscozacin, 17-20 Apr
1983 (? fr), D. N. Smith3827 (AAU, MO). PUNO:Rfo Can-
damo, nr.mouthof Rio Guacamayo,25 May 1992 (st), Gen-
try et al. 77156 (MO). UCAYALI:Prov. Coronel Portillo,
Dtto. CampoVerde,rd. to Nueva Requena,km 11, 22 May
1988 (Y fl-fr), Arana 74 (K); Pucallpa,Rio Uyayali, 8 Apr
1953 (5), Ferreyra9035 (USM), (Y fl-fr), Ferreyra9042
(USM); Prov. Coronel Portillo, nr. Neshuya, between Pu-
callpa & Tingo Maria, 20 Dec 1968 (5), Ferreyra 17649
(US, USM); Prov. Coronel Portillo, rd. Pucallpa-Tingo
Maria,km, 86, Bosque Nacional de von Humboldt,8 Aug,
1980 (5), Gentryet al. 29516 (MO); Prov.CoronelPortillo,
Bosque Nacional de von Humboldt, 24 Jun 1981 (Y fr),
Younget al. 1052 (BG, MO, NY, TEX).
BRAZIL. ACRE: Upper Rio Moa, Fazenda Arizona,
24-30 Sep 1984 (st), Campbelletal. 8162 (BG); Mun.Bras-
ileia, ColocaqcoSao Jose, 25 km N of km 4 of rd. Brasileia-
Assis Brasil, 26 May 1991 (5), Daly et al. 6722 (BG, INPA,
NY); Mun. Xapuri, Rio Acre, 3 hr. downstreamfrom Xap-
uri, 5 Nov 1991 (Y fl-fr), Daly et al. 7143 (BG, NY); Mun.
MarechalTaumaturgo,Rio Jurua,FazendaParaguay,3 Apr
1993 (d), Daly et al. 7727 (NY); Hwy. Abuna-RioBranco,
km 242-246, nr. Campinas,20 Jul 1968 (Y fl-fr), Foreroet
al. 6415 (GH, INPA, K, MG, MO, NY, S, U, US); nr.mouth
of Rio Macaua,9 Aug 1933 (9 fl-fr), Krukoff5389 (A, BM,
F, G, GB, K, MICH, MO, NY, S, U, US); Serrade Moa, 21
Apr 1971 (9 fr), Prance et al. 12171 (F, INPA,NY, U, US),
29 Apr 1971 (5), Prance et al. 12603 (F, INPA,MICH,MO,
NY, P, S, U). AMAPA:Reserva INCRA Rio Falsino, 15-20
Aug 1985 (st), Campbellet al. 10334 (BG); Mun. Macand,
rd. Serra do Navio (vill.)-Agua Branca, km 8, 4 Jan 1985
(9 fl-fr),Mori et al. 17671 (NY); Mun. Mazagao,Morrodo
Felipe V, 1 Oct 1986 (juv), Pires et al. 1393 (BG); Rio Ar-
aguari, 101 tN, 52?8'W, 28 Sep 1961 (9 fl-fr), Pires et al.
51301 (MG, MO, NY, U, US); Mun. Macapa, rd. BR.210,
147 km NW of P6rto Grande,30 Dec 1984 (9 fr), Rabelo
3112 (BG, NY); 2 km SE of Clevelandia,2 Aug 1960 (9
fl-fr), Westra47315 (BG, MG). AMAZONAS: Mun. Presi-
dente Figueiredo,Rio Uatuma, Sep 1986 (9 fl-fr), Amaral
2082 (INPA);Rio Jurua,Lago do Curape,Saracura,22 Aug
1975 (st), D. Coelho et al. (INPA)52385 (INPA);Benjamin
Constant, Mapatjirana,16 Oct 1956 (9 fl-fr), Drees 5596
171
(INPA, U); Mun. Manaus,ReservaFlorestalDucke, 28 Sep
1962 (6), Duarte 7187 (F, G, INPA, RB); nr. mouth of Rio
Embira, 13 Jun 1933 (6), Krukoff4789 (A, BM, F, G, K,
MICH, MO, NY, S, U, US); Mun. Humaita,nr. Tres Casas,
14 Sep-11 Oct 1934 (st), Krukoff6237 (A, BM, F, G, K,
MICH, MO, NY, RB, S, U, US); Mun. Sao Paulo de Oli-
venqa,nr.Palmares,11 Sep-26 Oct 1936 (Y), Krukoff8083
(A, BM, F, G, K, MICH, MO, NY, P, S, U, US); Cano Tu-
cano, Rio Cauaburf,12 Nov 1965 (6), Maguireet al. 60161
(GH, K, MG, MO, NY, S, US); Distr. Agropecuario,ca.
2?24-25'S, 59?40-45'W, 15 Nov 1988 (6), Mori et al.
19821 (BG, INPA, NY); Mun. Manaus,rd. Manaus-Itacoa-
tiara,km 26, ReservaFlorestalDucke, 6 Dec 1976 (6), Nas-
cimentoet al. (INPA)66334; Mun. Manaus,ca. 80 km N of
Manaus,Distr. Agropecuarioda SUFRAMA, 19 May 1992
(Y fl-fr),Nee 42752 (INPA,K, NY); rd.Manaus-Itacoatiara,
km 45, 29 Feb 1968 (Y fl), Rodrigues et al. 8439 (INPA,
MO); Rio Jurua,Cachoeira, May 1901 (Y fl), Ule 5512
(MG). MARANHAO: Mun. Monqao,Rio Turiaqu,4 km NW
of Urutawy,5 Feb 1985 (st), Balee et al. 408 (NY); Mun.
Carutepara,Gurupiuna,4 Nov 1986 (st), Bale et al. 2801
(BG, NY). MATO GROSSO:Aripuana,29 May 1976 (Y fl-
fr), Monteiroet al. 1126 (INPA);Rio Peixoto de Azevedo,
15 Aug 1980 (st), Werner24/80 (MG). PARA: Ariramba,1
Jun 1957 (6), Egler 439 (MG); Peixe Boi, Jul 1907 (6),
Goeldi (MG) 8232 (BM, G, MG); Rio Tapaj6s,Boa Vista,
28 Jul 1933 (9 fl-fr), Kauffinann603 (F); Tucuruf,Cagan-
cho, Jan 1981 (st), Lisboa et al. 2271 (MG); rd. Belem-
Brasilia,km 93, 5 Sep 1959 (6), Kuhlmannet al. 202 (GUA,
MG, US); nr. Belem, Sep-Oct (st), Pires 51868 (NY, US);
nr.Cachoeira,rd. BR.22, km 96, 29 Oct 1965 (9 fl), Prance
et al. 1778 (GH, K, MO, NY, S, U, US); Mun. Almeirim,
Monte Dourado,rd. to Sul de Pacanari,20 Nov 1978 (6 fl),
M. R. Santos 352 (MG); Rio Jarn,Monte Dourado, 15 Oct
1968 (6), N. T Silva 1226 (NY, U, US). ROND6NIA: Mun.
Santa Barbara,rd. BR.364, km 120, 23 May 1982 (9 fr),
Teixeiraet al. 672 (INPA, NY). RORAIMA:Mun. Alto Ale-
gre, Upper Rio Uaikas, Maiongong village, Sep 1994 (st),
Milliken 2101 (BG); Serra dos Surucucus, 2042-47'N,
63?33-36'W, NW of Mission Station, 21 Feb 1969 (9 fl-
fr), Prance et al. 10152 (INPA, NY, U, US).
BOLIVIA. BENI: Prov. Moxos, 130 km S of San Ig-
nacio, 18 Oct 1991 (st), Del Aguila et al. 196 (BOLV,LPB);
Prov.Vaca Dfez, rd. Riberalta-Cobija,km 6.7, 28 Sep 1993
(9 fr), Michel et al. 2437 (BG); Prov. Ballivian, Serranfa
del Pil6n Lajas, 18 May 1989 (9 fl-fr), D. N. Smith 13218
(BG, BOLV,LPB, MO, USZ); 3 km E of Riberalta,rd. to
Guayaramirim,7 Jun 1982 (d), Solomon 7982 (LPB, MO,
NY, U); Tumapasa,11 Jan 1902 (st), Williams400 (BM, K,
NY, US). COCHABAMBA:Prov.Carrasco,Estaci6nValledel
SajtaUMSS, 29 Oct 1991 (st), Atahuachiet al. 24 (BOLV),
Galarza et al. 22 (BOLV,LPB); Prov. Carrasco,rd. Santa
Cruz-VillaTunari,km 240, 8-9 Jul 1989 (9 fr), D. N. Smith
et al. 13643 (BG, BOLV,LPB, MO, USZ). LA PAZ:Prov.
Abel Iturralde,Alto Madidi,21 May 1990 (st), Gentryet al.
70264 (MO). PANDO:Prov. Nicolas Suarez, Bella Flor, 10
Oct 1989 (9 fr), Becket al. 19266 (BG, COL);Prov.Nicolas
Suarez, Bella Flor, 10 Oct 1989 (9 fl-fr), Beck et al. 19266
(COL, LPB); Prov.Manupiri,12 km W of Conquista,4 Oct
172
1991 (st), Beck et al. 20092 (BG, LPB); Prov.Nicolas Sua-
rez, Cocamita,4 Sep 1987 (J), Buchanan-Smith3 (BG, K,
LPB); Prov. Nicolis Suarez, Mukden, Jun-Dec 1979 (6),
Izawa 25 (MO); Prov. Manuripe,35 km N of PuertoAmer-
icana, 3 Jun 1994 (Y fl-fr), Jardimet al. 871 (USZ); Prov.
Madre de Dios, Concesi6n de Mobil, 20-25 Aug 1992 (Y
fr), Killeen 4407 (BG, USZ); Prov. Nicolas Suarez,Cobija,
19 Oct 1978 (Y fr), Meneces 786 (BOLV,INPA);Rio Ma-
deira, 6 km above Abuna, 11 Jul 1968 (Y fr), Prance et al.
5857 (GH, INPA, K, MG, MO, NY, P, S, U, US).
This species is ratheruniform.The most apparent
variationis found in the length of the petiolate parts
of the segments of the lamina, varying from nearly
absent to 6 cm long. Cecropia sciadophylla is the
most common and widespreadforest species through-
out the Amazon basin and the Guianas.It usually out-
numbersthe otherforest species. However,in the area
in which this species co-occurs with C. herthae (Ec-
uador, Colombia), the two species often occur in
about equal proportions.These two species can be
easily confused. They have the same general habit,
but C. herthae does not have petiolulatebases of the
lamina segments, the upper surface of the lamina is
? scabrous, and it has trichilia, although these are
ratherinconspicuous.
Vernacular names. Colombia:yemeoba (Mirania,
Amazonas); zmuikaz (Amazonas); guarumo blanco
(Casanare). Venezuela: yagrumo morado (Ama-
zonas); sarasara-yek(Bolivar); yagrumo montaniero
(Delta Amacuro). Guyana: congo pump, wanasoro
(Arawak).Suriname:manbospapaja.FrenchGuiana:
diapapaYe,makamakalu(Wayana).Ecuador:su, yan-
tan (Morona-Santiago);dundu (Quichua,Napo); hu-
arumo,maguimehue(Huaorani,Napo); mangineowe
(Haorani, Napo); su or suu (Shuar, Napo); yantan
(Napo). Peru: yanat (Huambisa, Amazonas), setico
colorado (Amazonas);torog (Cuzco);cetico colorado
(Huanuco); Imuisai (Huitoto, Loreto); setico de al-
tura, suu (Jivaro, Loreto); torototza (Machiguenga,
Madre de Dios); setico colorado, setico roja, tacona,
tacona ojo de venado (Pasco); setico colorado (Uca-
yali). Brazil: imbauba gigante, imbauiba ver-
melha (Acre); torem do igapo (Amapa');tour6mda
folha grande (Amazonas); ama'y-ata, ama'y-puku
(Ka'apor,Maranhao);sakaka(Maiangong,Roraima).
Bolivia: ambaibo blanco (Cochabamba);ambaibo
colorado, toronoma(Pando).
52. Cecropia silvae C. C. Berg, Acta Bot. Neerl. 21:
655. 1972;Acta Amazonica8(2): 180. 1978. Type.
Brazil. Para:Rio Jarn,Monte Dourado,PlanaltoA,
11 Sep 1968 (9), N. T Silva 956 (holotype: U;
isotypes: MG, NY, US).
FLORA NEOTROPICA
Tree,to 30 m tall. Leafy twigs 2-4 cm thick, hir-
tellous to subhispid.Laminasubcoriaceous,ca. 30 X
30 cm to 70 X 70 cm, the segments (12-)15-17, ob-
lanceolate,the incisions down to nearthe petiole; api-
ces obtuse to subacuteto short-acuminate;uppersur-
face scabrousto scabridulous,hispidulous,often also
with sparse arachnoid indumentum;lower surface
minutelypuberulousto pubescenton the main veins,
with arachnoid indumentum in the areoles, the
smallerveins, or also on the main veins; lateralveins
in the midsegment ca. 40-45 pairs, submarginally
loop-connected, unbranched;petiole ca. 25-55 cm
long, puberulous to subhirsute;trichilia fused, the
brown indumentumintermixedwith ratherlong stiff
to soft white hairs;stipules 5-6 cm long, hirtellousto
subhirsuteoutside, sericeous inside. Staminateinflo-
rescences:peduncleca. 9 cm long, hirtellousandwith
sparse arachnoidindumentum;spathewhitish, hirtel-
lous and with dense arachnoidindumentumoutside,
glabrousinside; spikes ca. 40-50, 4-8 X ca. 0.3 cm,
with stipes 0.2-0.3 cm long and with dense brown
pluricellulartrichomes;rachis hairy.Staminateflow-
ers: perianth tubular,ca. 1-1.5 mm long, glabrous,
the apex convex to plane; filaments flat; anthersca.
0.5-0.6 mm long, appendiculate,detachedat anthesis
(?), reattachedto the margins of the apertureby the
appendages(?). Pistillate inflorescencesin pairs (?),
erect (?); peduncle3-7 cm long, hirtellous;spatheca.
6-7 cm long, the color and indumentumas in the
staminateinflorescence;spikes 4, ca. 5-6 x ca. 0.4
cm, to 9 X 1.5 cm in fruit, sessile; rachis hairy.Pis-
tillate flowers: perianthca. 2 mm long, with arach-
noid indumentumbelow the apex and on the apex to
nearthe apertureoutside, also below the style channel
inside, the apex convex, puberulouson the marginsof
the aperture;style rather long; stigma penicillate.
Fruit oblongoid, ca. 2.5 mm long, tuberculate,dark
brown.
Distribution (Fig. 11.2). Brazil,northernPara,ex-
tending to southernSurinameand FrenchGuiana.
Specimens examined. SURINAME.Sipaliwinisa-
vanne, 22 Jan 1970 (st), Oldenburgeret al. 1112 (U).
FRENCH GUIANA. Piste St. Laurent-Apatou,km 46,
25 Feb 1984(st,juv),Prevost1727(BM,INPA,NY,P,U).
BRAZIL. AMAPA: Mun. Mazagao, Morrodo Felipe V,
1 Oct 1986 (st, juv), Pires et al. 1394 (BG). PARA:Rio
Jacamacaru,1 Jun 1957 (st), Egler 445 (MG); Rio Jari re-
gion, rd. PlanaltoA.-Tinguelim,24 Aug 1970 (6), Silva
3284 (IAN).
This species is distinguishedby the narrowseg-
ments of the lamina with numerouslateral veins in
the free part of the midsegment.This species shows
morphologicalaffinitiesto Cecropia ulei. Due to the
scarcity of material, there are some gaps in the de-
SYSTEMATICTREATMENT 173

scription. The collections from Amapa'and French usually green to yellowish, sometimespartlyreddish,
Guiana are made from juvenile trees, and their iden- or occasionally to red-brown, often subpersistent,
tity is not certain. (rather) sparsely puberulous to hirtellous or some-
times densely brownish- (or whitish-)hirsute,occa-
sionallypartlysetose with irritatinghairsoutside,gla-
brous or sometimes (densely) hairy inside; terminal
53. Cecropia strigosa Trecul,Ann. Sci. Nat. Bot., Ser
bud ? inflated.Staminateinflorescencesin pairs (or
3, 8: 82. 1847 (Aug). Type. Peru.Withoutlocality,
solitary), often subtendedby white, caducous bracts,
(d + Y), Ruiz & Pav6n s.n. (FI-W, photograph
to 1 cm, sometimes to 8 cm long, the peduncle erect
seen, mixed collection, the sheet with the stami-
to spreadingand the spikes (sub)pendulous;peduncle
nate inflorescence here designated as the lecto-
4-8(-10) cm long, puberulousto hirtellous; spathe
type; isolectotype: G).
10-23 cm long, (pale) green to yellowish, (rather)
CecropiabicolorKlotzsch,Linnaea20:531. 1847(Oct). sparsely puberulous to hirtellous, or sometimes
Peru.Huanuco: Munia,(9), Ruiz& Pavons.n. (ho- densely brownishto whitish hirsuteoutside, glabrous
lotype: B, destroyed;isotype:FI-W,photograph or hairy inside; spikes ca. (15-)25-100, (3-)5-22 X
seen).
ca. 0.2-0.3 cm, with stipes (0.5-)1-2.5 cm long and
CecropiamultifloraSnethlage,Notizbl. Bot. Gart.
Berlin-Dahlem 8:367. 1923.Type.Peru.Junin:Prov. minutely puberulous;rachis (sub)glabrousor short-
Tarma,Chanchamayo Valley,San Arnoldo,Dec hairy. Staminateflowers: perianth short-tubularto
1900(6), Weberbauer1837 (holotype: B, destroyed, cup-shaped,0.8-1.2 mm long, 2-lobed, glabrous;fila-
photographs in F, G, GH,fragmentin F). ments slightly swollen; anthersca. 0.5-0.7 mm long,
CecropiarugosaCuatrecasas, RevistaAcad.Colomb. not appendiculate,at anthesisnot detached.Pistillate
Ci.Exact.9(36/37):335. 1956.Type.Peru.Huanuco: inflorescencesin pairs, erect, soon pendulous, often
Divisoria,1700 m, 18 Sep 1946 (6), Woytkowski subtendedby whitish, caducousbracts,to 8(-15) cm
34554(holotype:F-n.v.;isotypes:G, MO). long; peduncle 4-16(-21) cm long, (sparsely)puber-
Tree,to 10(-18) m tall. Leafy twigs (I.5-)3-1-0 cm ulous to hirtellousto subhirsuteor also with arachnoid
thick, green or to darkbrown,puberulousto hirtellous indumentum, sometimes densely brownish hirsute,
with straight hairs, sometimes densely hirtellous to occasionally partlysetose with irritatinghairs;spathe
brownish(or whitish) hirsute(or glabrousand bluish 8-18(-23) cm long, the color and indumentumas in
by a waxy layer);intemodes occasionallywith ample the staminateinflorescence;spikes 3-7(-9), occasion-
brownpith. Laminachartaceousto subcoriaceous,ca. ally tortuose,6-15(-33) X 0.3-0.6 cm, to 25(-40) X
30 X 30 cm to 90 X 90 cm (to 140 X 140 cm), the 1(-1.5) cm in fruit, (sub)sessile or with stipes to 0.9
segments (6-)8-10(-12), the free partsof upperseg- cm long and densely hairy;rachis(sub)glabrous.Pis-
ments elliptic to subobovateto oblong, the incisions tillate flowers: perianth,ca. (0.6-)l.2-1.5(-2.3) mm
down to 5/10-9/10; apices short-acuminate(to sub- long, with arachnoidindumentbelow the apex out-
acute or to obtuse);uppersurfacesmooth or ? scab- side, also in the lower partof the style channelinside,
rous, often + bullate,sparselyto densely hispidulous the apex convex with a collar-shapedrim aroundthe
(on the main veins); lower surface minutely puberu- aperture,punctateto muriculate;style short; stigma
lous (with uncinateto curvedhairs)on the veins, often comose. Fruitellipsoid to oblongoid, 1.5-2 mm long,
with sparse longer (uncinate to straight)hairs espe- almost smooth.
cially on the main veins, or sometimesthe main veins Distribution (Fig. 18.2). From northernPeru to
hirtellousto brownish(sub)hirsute,with dense arach-
Bolivia, in forest and secondarygrowth,at 400-1900
noid indumentum in the areoles and also on the
m.
smallerveins, sometimesextendingto the main veins,
occasionally almost absent; lateral veins in the free Representative specimensexamined.PERU.Cuzco:
part of the midsegment 8-17 pairs, submarginally Prov.Paucartambo, betweenAtalaya& Pilcopata,31 Jul
loop-connected, many of them branched,the lower 1988 (d), Berg et al. 1617(BG,COL,Centrode Medicina
Andina,Cuzco); Prov.La Convenci6n,Quillabamba,Salas-
ones usually not distinctly loop-connected; petiole
pampa,rd. to Kitem, Rio Urubamba,1100 m, 28 Oct 1986
(20-)35-70(-105) cm long, rather sparsely puberu-
(Y fl), Nuiiez et al. 6304 (MO); Prov.La Convenci6n,Quil-
lous to hirtellous with straight hairs or sometimes labamba,Mandor,1200 m, 16 Oct 1987 (Y fl), Nuihezet al.
densely brownishor whitish hirsute;trichiliafused or 8230 (BG, MO); Prov.Urubamba,rd. Cuzco-Machupicchu,
separate on the usually ? bulging (subscrotiform) km 112, 1050-1300 m, 18 Nov 1987 (d), Nuiiez 8616
base of the petiole, the brownindumentumintermixed (MO); Prov. Quispicanchis,Puente Inambari,600 m, 8 Dec
with long white hairs or also with sparse to dense 1966 (d), VargasC. 18459 (US). HUANCAVELICA:Prov.
arachnoidindumentum;stipules (8-)10-35 cm long, Tayacaja,nr. Marabamba,1700 m, 25 Apr 1964 (Y fl-fr),
174
Tovar4798 (US). HUANUCO: Rd. Huanuco-Tingo Maria,
below Chinchoa,ca. 1800 m, 23 Nov 1997 (d), Berg et al.
1734 (BG, COL, MOL); rd. Huanuco-Tingo Maria, nr.
Santa Catalina,ca. 1400 m, 24 Nov 1997 (Y fl-fr), Berg et
al. 1739 (BG, COL, MOL); rd. Tingo Maria-Aucayacu,nr.
Pumahuasi,25 Nov 1997 (6), Berg etal. 1745 (BG); 69 km
NE of Tingo Maria, on rd. to Tocache, 16 Jul 1982 (Y fl-
fr), Gentry et al. 37616 (BG); Prov. Puerto Inca, Dtto. Yu-
yapichis, DANTAS, 14 Apr 1989 (Y fl-fr), Kroell S. 251
(BG, G), 16-31 Mar 1991 (Y fl), Tello 1117 (G, NY), (6),
Tello 1118 (G, NY). JUNiN:Rd. Tarma-SanRam6n,ca. km
75, ca. 1800 m, 28 Nov 1997 (Y fl-fr),Berg et al. 1754 (BG,
COL, MOL), (6), Berg et al. 1755 (BG, COL, MOL); La
Merced,29 May-4 Jun 1929 (st), Killip et al. 24047 (F, NY,
US); Prov. Chanchamayo,rd. to Mina Pichita, 1880 m, 2
Feb 1999 (Y fl), Ntiunez 68 (BG), 3 Feb 1999 (d), Niniez 75
(BG); nr. San Ram6n, La Florencia, 1800 m, 13 Sep 1989
(Y), Reynel et al. 4692 (MO). MADREDE DIos: Prov.
Manu, Parque Nacional Manu, Rio Manu, Rio Sotileja, 2
Oct 1986 (d), Fosteret al. 11570 (BG), (Y fl), Foster 11876
(BG, LPB, MOL, US). PASCO:Rd. Puente Paucartambo-
Oxapampa,ca. 1000 m, 28 Nov 1998 (6), Berg 1756 (BG,
COL, MOL); 3.7 km N of Puente Paucartambo,900 m, 30
Jan 1983 (9 fl-fr), Gentry et al. 39840 (BG, MO, MOL,
TEX); Iscozacin, 1 Apr 1986 (st, juv), Salick 7390 (NY).
SAN MARTiN:Prov. Lamas, rd. Lamas-Pamashto,ca. 700
m, 4 Dec 1997 (9 fl-fr), Berg et al. 1764 (BG, COL, MOL);
Prov. San Martin,rd. Tarapoto-Yurimaguas, ca. km 12, ca.
700 m, 5 Dec 1997 (6), Berg etal. 1772 (BG, COL,MOL);
Prov.Moyobamba,rd. Moyobamba-Jepelacio,ca. km 8, ca.
1100 m, 8 Dec 1997 (9 fl-fr), Berg et al. 1790 (BG, COL,
MOL); San Roque, 1350-1500 m, Jan-Feb 1930 (9 fl-fr),
LI. Williams2373 (US). UCAYALI:Rd. Tingo Maria-Pu-
callpa, km 82, nr. Boquer6n, 26 Nov 1997 (9 fl), Berg et
al. 1746 (BG, COL,MOL), (6), Berg et al. 1747 (BG, COL,
MOL);rd. Tingo Maria-Pucallpa,ca. km 125, 26 Nov 1997
(9 fl-fr), Berg et al. 1748 (BG, COL, MOL);Prov.Coronel
Portillo, San Alejandro,between Pucallpa & Tingo Maria,
km 115, 8 Apr 1953 (9 fl), Ferreyra9036 (US); rd. Pu-
callpa-Tingo MarIa,km 90-130, Bosque Nacional Alexan-
der von Humboldt,(9 fl-fr), Gentryet al. 41409 (BG, NY).
BOLIVIA. BENI:Prov. Ballivian, rd. Yucumo-Caran-
avi, km 13, 900 m, 4 Mar 1993 (d), Berg 1712 (BG, COL,
LPB); Prov. Ballivian, SerraniaPil6n Lajas, rd. Caranavi-
San Borja, 25 km from Yucumo, 850-900 m, 16-26 Feb
1990 (9 fl-fr), D. N. Smithet al. 13989 (USZ, MO), 14013
(BG, MO). COCHABAMBA: Rd. Cochabamba-VillaTunari,
ca. km 95, 1800 m, 6 Dec 1995 (9 fl-fr), Berg et al. 1723
(BG, BOLV,COL, LPB), (6), Berg et al. 1725 (BG, BOLV,
COL, LPB). LA PAZ: Prov. Sud Yungas, rd. Caranavi-Yu-
cumo, nr. La Cascada, ca. 2.5 km SSW of Quiquibey,900
m, 4 Mar 1993 (9 fl-fr), Berg 1711 (BG, COL, LPB), (9
fl-fr), Berg 1714 (AAU, BG, COL, LPB, MO, NY).
The trichilia are either fused or separate, even on
the same tree. In the southern part of the species
range, the trichilia are more commonly separate than
in the northern part of the range. In the southern part
of the range, reddish-colored parts are lacking, but in
FLORANEOTROPICA
the northernpartthe presenceof a red dye in the "um-
bilicus" and the basal partsof the main veins, or also
in the stipules, is not uncommon.In the southernpart
of the species range, the incisions of the lamina are
mostly down to 7/10, but in the northernpart, the
incisions are often down to 8/10 or even 9/10, the
segments are often narrower(obovateto oblong), and
the lateralveins of the midsegmentare more numer-
ous. The base of the petiole is mostly ? scrotiform,
but in particularin the northernpart of the species
range sometimes or often hardly so. The terminal
buds are ? conspicuouslyinflated.The spatheof sta-
minate inflorescences often remains hanging on one
or two of the longest spikes; the tips of these spikes
are curvedinto the calyptrateapex of the spathe.The
spikes of the pistillateinflorescencesare usuallyto 15
cm long at anthesis,but in Ucayali (Peru)they can be
to 33 cm long. The number of lamina segments is
usually 9-10, but in Huanuco(Peru)individualswith
10-12 segments can be found. Materialwith densely
hirsuteleafy twigs, stipules,petioles, veins of the lam-
ina beneathand/orspathescan be found in Huanuco,
Ucayali and San Martfn(Peru).This species is prob-
ably related to Cecropia marginalis and to C. mem-
branacea.
Vernacular names. Peru: tacona blanco (San
Martin);shiari (Ucayali).
54. Cecropia subintegra Cuatrecasas,Rev. Acad.
Colomb.Ci. Exact.6(22/23): 279. 1945. Type.Co-
lombia. Valle: Rio Digua, Piedra de Moler, 1000
m, 25 Aug 1953 (9), Cuatrecasas 15111 (holo-
type: COL; isotypes: F, US).
Tree,to 20 m tall. Leafy twigs 1-3 cm thick,green,
puberulousto subhispidulouswith straightto curved
to uncinate hairs. Lamina chartaceousto subcoria-
ceous, ca. 20 X 20 cm to 50 X 50 cm, 3-lobed, the
main radiatingveins 7-10, the incisions down to ca.
3/10-4/10 in the upper part of the lamina and the
lower part shallowly lobate to subentire;apices acu-
minate to acute; upper surface smooth, sparsely pu-
berulous;lower surfaceminutelypuberulousandwith
sparselonger uncinatehairs on the veins, with arach-
noid indumentumabsent or confined to the margin;
lateralveins in the free part of the midsegment8-10
pairs, submarginally,unbranched or some (lower
ones) branched;petiole ca. 10-40 cm long, minutely
puberulousor also with sparseto ratherdense arach-
noid indumentum;trichilia fused, the brown indu-
mentum intermixed with short white hairs; stipules
4-12 cm long, (purplish-)redor greenwith red stripes,
SYSTEMATICTREATMENT
(sparsely) puberulous and with brown pluricellular
hairs or also with arachnoid indumentum outside,
hairy to subglabrousinside. Staminateinflorescences
solitary,erect;peduncle2.5-3.5 cm long, puberulous;
spatheca. 10-15 cm long, (purplish)red, sparselypu-
berulousand with dense brownpluricellularhairsout-
side, glabrousor sparsely hairy inside; spikes ca. 7-
12, ca. 9-12 X 0.3 cm, with stipes 0.3-0.5 cm long
andpuberulous;rachissparselyhairy.Staminateflow-
ers: perianth tubular, 1-1.5 mm long, with short
arachnoidindumentumbelow the apex, the apex con-
vex, smooth to slightly tuberculate,the aperturesur-
roundedby a rim; filamentsflat; anthers0.5-0.6 mm
long, appendiculate,detached at anthesis, reattached
to the marginsof the apertureby the appendages(?).
Pistillate inflorescencesin pairs or solitary, erect to
pendulous; peduncle 3-24 cm long, puberulous to
subhispidulous;spathe 12-20 cm long, the color and
indumentumas in the staminateinflorescence;spikes
4(-6), ca. 9-12 X 0.5-0.6 cm, to 32 X 0.8-1 cm in
fruit, sessile or with stipes to 1 cm long and puberu-
lous; rachis hairy. Pistillate flowers: perianth 1.5-2
mm long, with arachnoidindumentumbelow the apex
outside, also in the style channelinside, the apex con-
vex to conical, muriculate;style ratherlong, straight;
stigma comose. Fruit ellipsoid, 2-2.5 mm long,
smooth, darkbrown.
Distribution (Fig. 18.8). Pacific coastal region of
Colombia (Valle and Nariiio) and Ecuador(Carchi),
in forest, at 800-1300 m.
Specimensexamined.COLOMBIA.NARINO:Mun.
Barbacoas,Cgto. Altaquer,Vrda. El Barro, 1325 m, Franco
et al. 4909 (BG); Mun. Barbacoas,El Diviso, 850 m, 7 Oct
1993 (Y fl-fr), Franco et al. 4463 (BG); Mun. Barbacoas,
Cgto. Altaquer,rd. El Barro-Junin,El Mirador,1150 m, 12
Dec 1993 (6), Franco et al. 5174 (BG); Mun. Barbacoas,
rd. Altaquer-El Diviso, nr. Reserva Colibries, 1300 m, 10
Mar 1995 (9 fl-fr), Lozano C. et al. 6906 (COL); Mun.
Barbacoas,Cgto. Junin,rd. Junfn-Barbacoas,ca. 950-1100
m, 14 Mar 1995 (d), Lozano C. et al. 6987 (COL);Ricaurte,
ca. 1300 m, 30 Mar 1941 (9 fr), SneidernA.485 (A, MICH,
NY, S, TEX, U, US). VALLE: Mun. Trujillo, rd. Trujillo-
Naranjal,km 20.4, Rio Sanquinini, 1200 m, 11 Oct 1993
(d), Franco et al. 4482 (BG, TULV).
ECUADOR.CARACHI: N of Lita, N of Rio Mira & E
of Rio Baboso, 1 Jul 1994 (st), Boyle et al. 3304 (BG).
The shallowly incised leaves are found only on
lateral branches. Before branches are produced,the
leaves on the trunk are more or less deeply incised
and often larger.In the monocaul state, the leaves are
similarto the normalC. obtusifolia.Subjuveniletraits
seem to reappearin the leaves of the branches,not
only in the first leaves formed, but persistently.This
175
submontanespecies shows close affinitiesto Cecropia
obtusifolia, in particularto the type with relatively
few segmentsof the lamina.Cecropiasubintegramay
prove to be distinct only at the level of subspecies.
55. Cecropia tacuna C. C. Berg & P. Franco,Novon
6: 248. t. 13, 1996. Type. Peru. Pasco: Prov. Ox-
apampa,5 km SE of Oxapampa,1850 m, 25 May
1983 (Y fl-fr), D. N. Smith4179 (holotype: MO;
isotypes: BG, K). Fig. 44
Tree,to 20(-30) m tall. Leafy twigs 4-8 cm thick,
green to bluish gray,densely villous and with filiform
brown pluricellularhairs (or only with these pluri-
cellular hairs). Lamina (sub)coriaceous,ca. 50 X 50
cm to 100 X 100 cm, the segments 10-12 or 13-18
(but on new shoots down to 8), the free parts of the
segments oblanceolateto oblong, the incisions down
to ca. 6/10-9/10; apices subacuminateto acute;upper
surface scabrous to scabridulous,sparsely to rather
densely hispidulous to subhispid or to hirtellous or
largely hirsuteto subvillous and with sparseto dense
filiformbrownpluricellularhairs, "umbilicus"prom-
inent and villous; lower surface rathersparsely sub-
villous and with filiformbrown pluricellularhairs or
also sparse white arachnoidindument on the main
veins, varying to pilose (to subtomentose) on the
lesser veins, with arachnoidindumentumin the are-
oles and on the reticulum;lateralveins in the free part
of the midsegment 15-25 or 25-35 pairs, most of
them unbranchedor the lower ones branched,sub-
marginally( ? faintly)loop-connected;petiole 40-80
cm long, ? densely (sub)villous and with filiform
brown pluricellularhairs and sparse arachnoidindu-
ment or sparselyvillous in the upperpart,glabrescent
or (sub)glabrous;trichilia absent; stipules 15-35 cm
long, green (or red to grayish),caducous,densely vil-
lous outside, sparselyto densely hairy inside. Stami-
nate inforescences in pairs, erect to spreadingwith
the spikes curved upward,often subtendedby cadu-
cous bracts,to 13 cm long; peduncle 5-12 cm long,
? densely white-villous (at least in the upper part)
and also or only with filiform brown pluricellular
hairs, often also with sparse white arachnoidindu-
mentum or (sub)glabrous; spathe 14-20 cm long,
green (or purple),with dense filiformbrownpluricel-
lular hairs and sparselyvillous (glabrescent)outside,
hairy inside; spikes ca. 10-25, 12-20 X 0.3-0.5 cm,
sessile or with stipes to 1 cm long and with brown
pluricellularhairs or also sparsely hirtellous;rachis
hairy with stiff hairs and arachnoidindumentum.Sta-
minate flowers: perianth tubular, ca. 1.2-1.8 mm
long, with short arachnoid indumentumbelow the
176 FLORA NEOTROPICA

FIG. 44. Cecropia tacuna. 1. Lamina,reduced.2. Apex of lamina and venation. 3. Stipules and young leaf (Berg et
al. 1635). 4. Staminateinflorescence with spathe. 5. Staminateflower. 6. Stamen and filament after detachmentof anther
(Berg et al. 1634). 6. Pistillate inflorescence at anthesis. 7. Pistillate flower and fruit. 8. Pistillate flower. 9. Fruit (Berg et
al. 1635). (From Novon 6: 249. 1996, modified.)
SYSTEMATICTREATMENT
apex, the apex plane to slightly convex, sparsely to
densely muriculate; filaments swollen; anthers ca.
0.6-0.8 mm long, not appendiculate,remaining at-
tached at anthesis. Pistillate inflorescencesin pairs,
erect to spreadingand with the spikes curvedupward,
often subtendedby caducous bracts, to 13 cm long;
peduncle 5-12 cm long, with indumentumsimilarto
that of the staminateinflorescence;spathe 9-14 cm
long, the color and indumentumas in the staminate
inflorescence;spikes 3-7, 6-13 X ca. 0.5 cm, to 22
X 1.5 cm in fruit, (sub)sessile; rachis hairy with stiff
hairs and arachnoidindumentum.Pistillate flowers:
perianth, ca. 1.5-2 mm long, with arachnoidindu-
mentumbelow the apex outside, also in the lower part
of the style channel inside, the apex plane to slightly
convex, muriculate;style long, slightly curved, mur-
iculate; stigma comose. Fruit oblongoid, ca. 1.5 mm
long, finely tuberculate.
Distribution (Fig. 8.1). Peru, from Pasco to
Cuzco, in cloud forest, at (800-)1800-2500(-2700)
m.
56. Cecropia telealba Cuatrecasas, Revista Acad.
Specimensexamined.PERU. AYACUCHO:Between
Huanta & Rfo Apurimac, 750-1000 m, 7-17 Nov 1929
(juv),Killipet al. 23117 (NY,US). Cusco: Prov.Paucar-
tambo,rd.Pilcopata-Paucartambo,ca. 2000m, 2 Aug 1988
(d), Berg et al. 1634 (BG, Centro de Medicina Andina,
Cusco),2200-2300m,2 Aug1988(Y fl-fr),Bergetal. 1635
(BG,COL,CentrodeMedicinaAndina,Cusco);MachuPic-
chu, 19 Apr 1957 (st), Ellenberg880 (U); Prov.Paucar-
tambo,rd.Paucartambo-Pilcopata, km 132,2260-2290m,
27 Jun1978(st),Gentryetal. 23564(BG);Prov.Urubamba,
MachuPicchu, 2000 m, 4 Jul 1972 (5), Mueller 2861 (LZ);
Prov.Urubamba, &Cedrobamba,
betweenPampacahua Ma-
chu Picchu, 2800 m, 8 Jul 1989 (st), Nuhez et al. 11111
(MO); Prov. Urubamba,nr. Machu Picchu, Rio Mandor,
2055 m, 2 Jun 1982 (5), Peyton et al. 374 (MO); Prov.
Paucartambo,Kosfiipata,2700 m, 4 Jul 1972 (5), VargasC.
15490 (US). HUANUCO: Rd. Huanuco-Tingo Maria,Car-
pish,ca. 2500 m, 23 Nov 1997(Y fl-fr),Berget al. 1731
(BG,COL,MOL),2400m, 10Mar1982(5), Gentryet al.
36153(BG,MO,USM).PASCO: Prov.Oxapampa, RioBo-
queria,ca.26 kmfromOxapampa, viaRioYamaquizu, 2040
m, 3 Jun 1982 (d), D. N. Smithetal. 1832 (BG, MO, USM);
Prov. Oxapampa,5 km SE of Oxapampa, 1850 m, 9 Apr
1983 (5), D. N. Smith 3663 (BG, MO, MOL); Prov. Oxa-
pampa, Huancabamba,Rio Yanachaga,2280 m, 26 May
1983 (Y fl), D. N. Smith et al. 4196 (BG, MO); Prov.Oxa-
pampa, Oxapampa-VillaRica rd., 29 Sep 1983 (D fl-fr),
D. N. Smith et al. 5340 (BG, MO); Prov. Oxapampa,RIo
San Albert valley, E of Oxapampa,2300 m, 4 Jul 1984 (st),
D. N. Smithet al. 7608 (BG, MO, USM); Prov.Oxapampa,
Yanachagavia Rio San Daniel, 2500 m, 17 Jul 1984 (st),
D. N. Smithet al. 7848 (BG, MO, USM); Prov. Oxapampa,
Rio Alberto valley, E of Oxapampa,slopes of Cord.Yana-
chaga, 2400 m, 23 Jul 1984 (st), D. N. Smith et al. 7974
177
(BG, MO, USM); Prov. Oxapampa,Palmazu, 2100 m, 28
Sep 1984 (Y fl-fr), D. N. Smith 8555 (BG, MO). SAN
MARTiN: Prov.Rioja, rd. PedroRuiz-Moyobamba,2100 m,
7-9 Aug 1983 (d), D. N. Smithet al. 4686 (MO).
This species is probably related to the Bolivian
montane species, Cecropia elongata. It shows simi-
larities to the villous forms of C. angustifalia (see
page 62) in the indumentum. In the collections from
Huainuco (Peru), the leafy twigs are subglabrous,
bearing only brown pluricellular hairs, the peduncles
are glabrous, and the petioles are glabrous or sparsely
villous only in the upper part. Moreover, these col-
lections differ in the less deeply incised laminas
(down to 6/10-7/10), fewer incisions (10-12), and
fewer lateral veins in the free part of the midsegment
(15-25 pairs), features matching those of subjuvenile
leaves of material from Cusco and Pasco. Ants are
absent.
Vernacular name. Peru: tacuna (Pasco).
Colomb. Ci. Exact. 6(22/23): 293 (1945). Type.
Colombia. Valle: Rio Cali, Quebradahonda, above
La Glorieta, rd. to Miralindo, 2100-2250 m, 31
Oct-I Nov 1044 (d), Cuatrecasas 18406 (holo-
type: COL; isotypes: F, US).
Cecropia telealba Cuatrecasasvar. hirsuta Cuatrecasas,
Revista Acad. Colomb. Ci. Exact. 6(22/23): 294
(1945). Type.Colombia.Valle:Rio Calima,El Cairo,
1650-1750 m, 6-7 Jan 1943 (Y), Cuatrecasas13947
(holotype:COL; isotype: US).
Cecropia alborugosa Cuatrecasas,Revista Acad. Col-
omb. Ci. Exact. 9(36/37): 330. 1956. Type. Colom-
bia. Valle: Rio Digua, Quebradadel San Juan, be-
tween Paraguita& Queremal,1570-1740 m, 17 Mar
1947 (d), Cuatrecasas 23841 (holotype: US; iso-
types: F, VALLE).
Cecropia teleargenteaCuatrecasas,Revista Acad. Col-
omb. Ci. Exact. 9(36/37): 333. 1956. Type. Colom-
bia. Valle: Rio Bugalagrande,Jamaica, 1640-1700
m, 26 Apr 1946 (Y), Cuatrecasas21011 (holotype:
US; isotypes: F, VALLE).
Tree, to 20(-30) m tall. Leafy twigs 1.5-5 cm
thick, green, hirsute, sometimes only at the stipular
scars; intemodes partly to entirely (?) filled with
orange-brown pith. Lamina coriaceous, ca. 20 X 20
cm to 90 X 90 cm, the segments 8-10, the free parts
of the upper segments obovate or elliptic, the inci-
sions down to 7/10-8/10; apices acute to obtuse; up-
per surface smooth, often ? rugose, very sparsely
strigose and with dense to + sparse arachnoid indu-
mentum (or subglabrous), lower surface sparsely to
densely hirsute to hirtellous on the main veins, hir-
178
tellous to puberulouson the smallerveins, with arach-
noid indumentumin the areoles or extending to the
smallerveins or also to the main veins (not distinctly
in two layers as in C. telenitida);lateralveins in the
free part of the midsegment ca. 15-25(-30?) pairs,
submarginallyloop-connected,unbranchedor some-
times branched;petiole 15-65 cm long, sparselysub-
villous in the upperand lower partor also with arach-
noid indumentum;trichilia fused (or separated),the
brown indumentumintermixed with short (or also
long) white hairs;stipules(6-) 13-32 cm long, reddish
to purplishor pale green to yellowish, hirsuteto sub-
villous (to rathersparselystrigose to hirtellousto sub-
sericeous outside, sparselyhairy inside. Staminatein-
florescences in pairs or solitary, erect; peduncle 5-7
cm long, subvillous,often only in the upperandlower
part;spathe9-17 cm long, pink to reddishto purplish
or pale green to yellowish, subvillous outside, gla-
brous inside; spikes 8-12, 5-12 X 0.5-0.9 cm, with
stipes 0.5-1.2 cm long and sparselyhairyor glabrous;
rachis hairy. Staminateflowers with pedicels to 1.5
cm long and minutely puberulous;perianthtubular,
ca. 1.2-2 mm long, minutely puberulousbelow the
apex, the apex plane to slightly convex; filamentsflat;
anthers0.6-0.8 mm long, appendiculate,detachedat
anthesis (?), reattachedto the marginsof the aperture
by the appendages (?). Pistillate inflorescences in
pairs or solitary, erect, later on deflexed, subtended
by caducous bracts,to 4 cm long; peduncle 3-11 cm
long, subvillous, often only in the upper and lower
part or subglabrous;spathe 8-13 cm long, the color
and indumentumas in the staminate inflorescence;
spikes l-3(-4), (1.4-)5-9 X 0.4-0.6 cm, to 32 X 2.5
cm in fruit, sessile or with stipes to 1 cm long and
subvillous; rachis hairy. Pistillate flowers: perianth
ca. 2-3 mm long, with arachnoidindumentumbelow
the apex outside, also in the lower part of the style
channel inside, the apex convex, muriculateto min-
utely puberulous(only in the lower part);style long,
straight, minutely puberulous; stigma penicillate.
Fruit oblongoid, 3.5-4 mm long, finely tuberculate.
Distribution (Fig. 10.2). Colombia (Caldas to
Cauca), in (cloud) forest, in the westernCordilleraat
1300-2500 m.
Representativespecimensexamined.COLOMBIA.
CALDAS: Rio Boquia,Salento,1600-1900m, 22 Jul 1922
(st), Killip et al. 8827 (GH). CAUCA: Between Santanderde
Quilichao & Rfo Ovejas, 1500 m, 3 Feb 1965 (d), Cuatre-
casas et al. 26894 (VALLE- underthe same numbera col-
lection from Valle); Sevilla, Las Brisas, 2000 m, 10 Apr
1970 (Y fl-fr), Cuatrecasaset al. 28946 (US); between San
Antonio & Rio Ortega, 2000-2200 m, 1-2 Jul 1922 (st),
Pennell et al. 8042 (GH). QUINDIO: Withoutlocality, 1200
FLORANEOTROPICA
m, Mar 1876 (st), Andre2574 (K, NY); Mun.Circasia,Vrda.
BarcelonaAlta, Finca Buenos Aires, 1690 m, 23 Mar 1991
(6), Agudelo et al. 1043 (COL);Mun. Armenia,Vrda. Las
Colillas, Finca Las Mercedes, 1700 m, 12 Jun 1992 (9 fl-
fr), Wlez et al. 2926 (COL). RISARALDA: Nr. Termales
2060-2130 m, 13 Aug 1995 (9 fr), Wijninga541 (BG, U).
VALLE: Rio Cali, Rio Pichind6, Alto de las Brisas, 2050-
2100 m, 29 Oct 1944 (9 fl-fr), Cuatrecasas 18353 (COL,
F, US); Rio Pichinde, Los Carpatos,2250-2350 m, 24 Jul
1946 (9 fr), Cuatrecasas21691 (F, US, VALLE);El Tabor,
above Las Brisas, 1970-2100 m, 19 Oct 1946 (9 fl), Cua-
trecasas 22304 (F, US); above La Carbonera,between Las
Brisas & Alban, 2000 m, 24 Oct 1946 (st), Cuatrecasas
22504 (F); nr.Las Mesitas, aboveVilla Colombia,El Placer,
1800 m, 4 Feb 1965 (6), Cuatrecasaset al. 26894 (TULV,
US); Mun. Trujillo,rd. Trujillo-Naranjal,km 9, 1800 m, 11
Oct 1993 (9 fl-fr), Franco et al. 4479 (BG, COL); Mun.
San Pedro,rd. Buenos Aires-San Pedro,km 15, 1900 m, 13
Oct 1994 (6), Franco 4490 (BG); Mun. Yotoco, El Canay,
rd. Medio Canoa-Darien,1500 m, 15 Mar 1994 (6), Franco
et al. 4528 (BG); Mun. Loboguerrero,rd. to Buenaventura,
1350 m, 15 May 1994 (9 fl-fr), Franco et al. 4530 (BG,
TULV); Santa Helena, above Topacio, edge of Los Faral-
lones de Cali NationalPark, 1920 m, 12 Dec 1985 (9 fl-fr),
Gentry et al. 53202 (MO); E of Bitaco, 1500 m, 16 Nov
1963 (9 fr), Hutchinsonet al. 3039 (F, G, K, MICH, NY,
P); Mun.AnsermaNuevo, rd.Anserma-SanJos6del Palmar,
1860 m, 12 Nov 1985 (6), Lozano C. et al. 4978 (COL);
Mun. Darien, Alto Calima, 1970 (6), Mahechas.n. (UDBC
5789).
Cecropia telealba shows affinities to C. gabrielis,
from which it can be distinguished by the (usually)
dense arachnoid indumentum on the upper surface of
the lamina and the consistent presence of trichilia.
Cecropia gabrielis (normally) occurs at lower eleva-
tions than C. telealba. It is the conspicuous white-
leaved species on the mountain slopes bordering the
Cauca valley.
57. Cecropia telenitida Cuatrecasas, Revista Acad.
Colomb. Ci. Exact. 6(22/23): 295. 1945; Velas-
quez, Acta Bot. Venez. 6: 62, t. 15. 1971. Type.
Colombia.Norte de Santander:Mun. Toledo, Rio
Samaria, 2000-2100 m, 30 Oct 1941 (9), Cua-
trecasas, Schultes & E. Smith 12781 (holotype:
COL; isotype: F). Fig. 45
CecropiatelealbidaCuatrecasas,RevistaAcad. Colomb.
Ci. Exact. 6(22/23): 294. 1945. Type. Colombia.
Huila: Above Guadalupe,Resina, 1850-1900 m, 20
Mar 1940 (9), Perez Arbeldez & Cuatrecasas8371
(holotype:COL; isotype: F).
Cecropiatelenivea Cuatrecasas,Revista Acad. Colomb.
Ci. Exact. 6(22/23): 295. 1945. Type.Colombia.Pu-
tumayo: Rio Sibundoy, 2200 m, 1 Jan 1941 (6),
Cuatrecasas11582 (holotype:COL; isotype: F).
SYSTEMATICTREATMENT 179

FI.4.Ccoi keeiia .Lmn,rdcd .Ae flmn n eain(eg13) .Siue n on

4. Ape
leaf. StmiCecropiarescencwitidh Lamia,h
(Brgeduced. 265. oftaminaten floer.n.atamen (Berg 1232).
3. Stiulstillateun
inflorescence at anthesis. 8. Pistillate flower and style. 9. Fruit (Berg et al. 1666). (From Fl. Ecuador 48: 11l. 1993, modified.)
180
Cecropia teleincana Cuatrecasas, Revista Acad. Col-
omb. Ci. Exact. 6(22/23): 296. 1945. Type. Colom-
bia. Antioquia:Alto del Toyo, between Antioquia&
CaniasGordas,2200 m, 2 Mar 1942 (d), Cuatreca-
sas 13548A (holotype:COL; isotype: F).
Cecropia santanderensis Cuatrecasas, Revista Acad.
Colomb. Ci. Exact. 6(22/23): 296. 1945; Velasquez,
Acta Bot. Venez. 6: 58, t. 13. 1971. Type. Colombia.
Norte de Santander:Rfo Chitaga,Ventanas,Alto del
Loro, 1820 m, 17 Oct 1941 (Y), Cuatrecasaset al.
12346 (holotype:COL; isotype: F).
Cecropiaangelica C. C. Berg & P. Franco,Fl. Ecuador
48: 9, t. 1. 1993. Type. Ecuador.Morona-Santiago:
Angel Sabio, between Lim6n (Gral. Plaza) & Gual-
aco, ca. 2250 m, 28 Jan 1981 (Y), Berg 1229 (ho-
lotype: QCA; isotypes: AAU, BG, COL, NY, U).
Tree, to 15(-30) m tall. Leafy twigs 2-8 cm thick,
green to bluish by a waxy layer,glabrousor (initially)
with dense brown pluricellulartrichomes, hirtellous,
villous to hirsute, or puberulouswith straightwhite
(unicellular)hairs; intemodes with scarce pith. Lam-
ina (sub)coriaceous,ca. 30 X 30 cm to 90 X 90 cm
(to 100 X 100 cm), the segments 7-10(-1 1), the free
partsof the uppersegmentsovate to elliptic to oblong
to (sub)obovate,the incisions down to 4/10-9/10; api-
ces roundedto short-acuminate;uppersurfacesmooth
or scabridulous (to scabrous), very sparsely to
densely puberulous(to hirtellousor subvelutinous)or
hispidulous and with sparse to dense arachnoidin-
dumentum, often slightly bullate, lower surface
hirtellous to subtomentose (or puberulous), espe-
cially on the main veins hirsute to villous, or main
veins subglabrous,with very short arachnoidindu-
mentum in the areoles or also on the reticulumand
usually very sparse to ratherdense longer arachnoid
indumentumon the main veins and supportedby the
other indumentumon the smaller veins (the arach-
noid indumentumthus in two distinct layers); lateral
veins in the free part of the midsegment7-16 pairs,
submarginally loop-connected, unbranched or
branched,the smaller veins often prominent;petiole
ca. 25-90 cm long, glabrous,villous in the upperor
also the lower part, also with arachnoidindumentum
and/or dense brown pluricellulartrichomes (some-
times at the base dense, and suggesting the occur-
rence of trichilia, but Muillerianbodies lacking), or
puberulous; trichilia absent or present, weakly
developed without Mullerianbodies or with Muller-
ian bodies and separateor fused (andthen sometimes
lobate), the brown indumentum intermixed with
short or also long white hairs; stipules (12-)20-55
cm long, green to orange to pink to reddish to dark
maroon,with sparse to dense darkbrown pluricellu-
lar hairs and sparselyminutely puberulousto tomen-
FLORA NEOTROPICA
tellous or often also sparsely to densely hirsute to
villous to sericeous, subglabrous, subhirsute or
hirtellous outside, densely to sparsely subvillous to
subsericeous or subglabrousinside. Staminateinflo-
rescences solitary or in pairs, (usually/always?)sub-
tended by caducous bracts,to 8 cm long, the pedun-
cle erect and the spikes spreadingto subpendulous;
peduncle 2-11 cm long, glabrousor in the upperpart
sparselyto densely (sub)villous or puberulousto hir-
tellous to subhirsute; spathe 9-24 cm long, red-
brown to orange to pinkish or yellowish brown or
yellowish, rathersparsely to densely (sub)villous to
hirsute or also with sparse arachnoidindumentum
outside, sparsely hairy or glabrous inside; spikes 5-
12(-16), 6-25 X ca. 0.5-1 cm, sessile or with stipes
to 1 cm long and hairy;rachis hairy.Staminateflow-
ers: perianthtubular,1-2(-2.5) mm long, (sparsely)
puberulous or glabrous below the apex, the apex
slightly convex, glabrous;filamentsswollen; anthers
0.6-0.8 mm long, detachedat anthesis (?). Pistillate
inflorescencessolitary or in pairs, erect to deflexed,
(usually/always?)subtendedby caducousbracts,to 8
cm long; peduncle 4-12(-15) cm long, glabrous or
sparsely villous or ? densely hirtellous to puberu-
lous; spathe 6-13 cm long, the color and indumen-
tum as in the staminateinflorescence;spikes (I-)4-
5(-6), purplishor reddish,3-12 X 0.7-1.6 cm, to 25
X 2.5 cm in fruit, subsessile (or with to 2 cm long
stipes); rachis subglabrous.Pistillate flowers: peri-
anth2-2.5 mm long, with arachnoidindumentumbe-
low the apex outside, also below the style channel
inside (or the style channelminutelypuberulous),the
apex conical to convex, hispidulous near the slit-
shaped aperture; style rather long, puberulous;
stigma comose to subpeltate.Fruit oblongoid to sub-
obovoid, 2-2.5 mm long, tuberculateor smooth.
Distribution (Fig. 18.7). The Andean region of
Venezuela to southernColombia, in the central and
eastern cordilleras, and from southern Ecuador to
northernPeru, in cloud forest, as Podocarpus forest
in Ecuador,at (1200-)1400-2600 m.
Representativespecimensexamined.COLOMBIA.
ANTIOQUIA:Mun.Medellin,Cgto.SantaElena,12 km SE
of Medellin, 2200 m, 21 Jan 1994 (d), Callejas et al. 11110
(HUA, COL); CordilleraCentral,between Palmitas& Alto
(or Boquer6nde San Crist6bal),2200-2250 m, 5 Oct 1961
(d), Cuatrecasas 26247 (COL, P, US), (9 fl-fr), Cuatre-
casas 26248 (COL, P, US); El Retiro, Los Alpes, 15 Jan
1953 (st), Hno. Daniel 4492 (US); Mun. Medellfn, Santa
Elena, old rd. to Rionegro, 1700 m, 24 Mar 1994 (d),
Franco et al. 4579 (BG, HUA), (9 fl-fr), Franco et al. 4581
(BG, HUA);Yarumal,2150 m, 28 Mar 1994 (d), Franco et
al. 4602 (BG, HUA); Mun. Guarne,ParqueEcol6gico Pie-
SYSTEMATICTREATMENT 181

dras Blancas, 2350 m, 9 Feb 1994 (J), Rolddn et al. 2161 Dtto. Moran,Mun. Barbacoas,nr. Hato Arriba, 1870 m, 12
(COL, HUA). BOYACA: Mun. Pajarito,between Corinto& Aug 1983 (9 fl), Trujilloet al. 18553 (MO). MERIDA:Filo
Peniadel Gallo, 1800 m, 29 Jan 1995 (d fl-fr), Franco et al. del Sai-Sai-ElCarrizal, 1850 m, 24 Feb 1955 (9 fl-fr),Ber-
4640 (BG, COL). CAQUETA: Mun. Florencia,below Gabi- nardi 2072 (NY); above Las Cuadras,QuebradaMolino, N
nete, km 17, 2200 m, 15 Oct 1993 (6), Franco et al. 4494 of Torondoy,1820-2255 m, 27 Mar 1944 (st, juv), Steyer-
(BG, COL), (cd), Franco et al. 4498 (BG). CASANARE: mark55817a (F); Rio Capaz, above La Azulita, 2100-2400
Mun. Sacama,Vrda. Guivarin,1750 m, 27 Jan 1995 (Y fl- m, 1 Sep 1966 (d), Steyermarket al. 97107 (NY, US, VEN).
fr), Franco et al. 4639 (BG, COL, NY). CAUCA:Moscopan PORTUGUESA: Dtto. Sucre, La Divisoria de la Concepci6n,
region, Rfo San Jose, Aguabonita,2280 m, 30 Jan 1947 (Y 1700 m, 24 Oct 1985 (st), Werif7587 (BG, MO). TACHIRA:
fl-fr), Cuatrecasas 23543 (US, VALLE) & Cuatrecasas Dtto. Junin,betweenRubio& Delicias, 2200 m, 19 Oct 1984
23553 (VALLE). CESAR:Sierra de Perija, E of Manaure, (st), Bono 4317 (MO); Rio Copas, between Las Copas &
San Antonio, 1700 m, 15 Nov 1959 (i), Cuatrecasaset al. San Vincente de la Revancha,S of Alquitrana,SW of Santa
25346 (COL, US); Mun. Manaure,rd. Manaure-ElCinco, Ana, 2000-2400 m, 22 Jan 1988 (9 fl-fr), Steyermarket al.
1950 m, 14 Nov 1993 (Y fl-fr), Franco et al. 4520 (BG), 101275 (MO, NY, VEN); QuebradaAgua Azul, S of El Re-
(c), Franco et al. 4521 (BG). CUNDINAMARCA: Mun. poso, 14 km SE of Delicias, 2150-2300 m, 22-23 Jul 1979
Funza, Cerro de Rosales, ca. 2600 m, 26 Oct 1961 (6), (9 fl-fr), Steyermarket al. 118640 (MO, U, VEN). ZULIA:
Cuatrecasaset al. 26508 (COL, P, US), (Y fl), Cuatrecasas Sierrade Perijd,between Misi6n de Los Angeles de Tukuku
et al. 26510 (COL, P, US); Mun. Granada,2225 m, 12 Mar & Pishikakaro,1400-1850 m, 1-3 Apr 1972 (9 fl-fr), Stey-
1993 (? fl-fr), Franco et al. 4342 (BG, COL); Mun. Tena, ermarket al. 105784 (F, MO, NY, US, VEN).
rd. Bogota-La Mesa, Vrda. Laguneta, 1900 m, 6 Feb 1995 ECUADOR. LOJA:San Jose de Yanacocha,ca. 1 km W
(6), Franco et al. 4643 (BG, COL); rd. San Miguel-La of jct. of rd. Zamora-Loja& Rfo Jipiro, nr. border Loja-
Aguadita,2-3 km below QuebradaAguabonita,2100 m, 12 Zamora-Chinchipe,nr. Rio Jipiro,2300 m, 5 Jan 1991 (d),
Aug 1970 (9 fl-fr), Idrobo 6321 (COL);Mun. Pacho, 1975 Berg et al. 1665 (BG, GB, LOJA,QCA), ca. 2450 m, 5 Jan
(d), Mahecha 1764 (UDBC); Mun. Pasca, Vrda. El Pedre- 1991 (9 fl-fr),Berg etal. 1666 (BG, LOJA,QCA);rd.Loja-
gal, 2550-2600 m, 27 Jun 1987 (9 fl-fr), G. Morales et al. Zamora,7 km SE of Loja, 2300-2400 m, 3 Jul 1947 (9 fl-
1116 (COL). HUILA:Mun. La Plata, Vrda. Agua Bonita, fr), R. Espinoza 1575 (NY). LOJA/ZAMORA-CHINCHIPE:
Finca Merenberg,1200-1300 m, 15 Jul 1975 (6), Dfaz P Rd. Loja-Zamora,ca. km 14, 2500-2820 m, 29 Dec 1988
et al. 575 (COL); Finca Merenberg,2300 m, 23 Dec 1975 (9 fl-fr), Jorgensen et al. 65761 (AAU, BG), QCA).
(d), Gaulin 5 (COL);Finca Merenberg,Caucaborder,E of MORONA-SANTIAGO: Angel Sabio, between Lim6n &
Leticia, 2300 m, 8 Jul 1984 (9 fr), Gentry et al. 47757 Gualaceo,ca. 2250 m, 28 Jan 1981 (9 fl-fr),Berg 1232 (BG,
(JAUM,MO); 30 km NW of Palermo, ca. 2700 m, 9 Oct COL, QCA, U). ZAMORA-CHINCHIPE: 11 km E of Loja,
1944 (9 fr), Little 8780 (COL, US); Rfo Villalobos, nr. Rio 2600 m, 5 Sep 1975 (9 fl-fr), Little et al. 248 (COL,LOJA,
Suazita, 1400 m, Jan 1943 (9 fl-fr), Schulteset al. 5172 (F, Q, QAME, QCNE).
GH); confluence of Rio Villalobos & Rio Cachos, 1400 m, PERU. SAN MARTiN:Prov. Moyobamba,Rioja, Ven-
Jan 1943 (9 fl-fr), Schultes et al. 5205 (COL, F, GH). ceremos, 1985 (9 fl), Lao M. 21 (K); Prov. Mariscal Ca-
META:ParqueChingaza,rd. from San Juanito,17 Sep 1997 ceres, Rio Abiseo NationalPark,N-side of riveracross from
(st), E. Acero D. 43 (COL); CordilleraOriental, between La Playa base camp, 2600 m, 1 Sep 1985 (6), Young1544
Hda. El Pato & camp Los Cueros, ca. 2150 m, 12 Jan 1944
(F, MO).
(9 fr), Little 8088 (COL, US). NORTEDE SANTANDER:
Ocafiaregion, between La Maria& Jurisdicciones, 2200 m, The species is in cultivationin the BotanicalGar-
25 Sep 1969 (6), Cuatrecasas et al. 27940 (US); Ocaiia den "Jose Celestino Mutis" in Bogota (9 Jun 1993
Region, nr.Apasica, Quebradadel Guarumal,1900-2000 m, [d], Franco et al. 4461 [BG, COL]).
26 Sep 1969 (9 fl-fr), Cuatrecasaset al. 27949 (COL, NY, The type materialof Cecropia teleincana and C.
P, US); Mun. Toledo, 1850 m, 24 Jan 1995 (9 fl), Franco
santandarensis,or more generally, the materialcol-
et al. 4624 (BG, COL). QUINDIO:Mun. Pijao, Vrda. Car-
lected in Antioquiaand Santander(Colombia),differs
cineros, Finca La Esperanza, 1920 m, 14 Aug 1992 (9 fl-
fr), Velez et al. 2997 (COL). PUTUMAYO: 1.5 km E of Si- (somewhat)from the other collections of the species
bundoy, 2200 m, 3 Oct 1963 (9 fl-fr), Bristol 1441 (COL, in the relatively short stipules (to 25 cm), in the hir-
GH, US); rd. Mocoa-Pitalito, between Suacira& SantoDo- sute ratherthan villous indumentum,and in the rela-
mingo, 2000 m, 5 May 1994 ( 9 fr), Francoetal. 5540 (BG); tively long peduncle (to 15 cm long) and relatively
nr. Sibundoy,2250 m, 18 Feb 1942 (9 fr), Schultes 3275 short spikes of the pistillate inflorescence.Moreover,
(F, GH). SANTANDER: Rio Surata,above Surata,2000-2300 specimenswith this hirsuteindumentum(such as Cal-
m, 5-6 Jan 1927 (st), Killip et al. 16667 (GH, NY, US). lejas et al. 11110 and Franco et al. 4579) appearto
TOLIMA:Rio Claro, E of Nevada del Huila, ca. 2700 m, 30
be ratheroften (?) inhabitedby ants,in contrastto the
Sep 1944 (d), Little 8747 (COL, F, US).
VENEZUELA. LARA:Huymucaro,rd. to Barbacoas, type with villous indumentum
as common in other
1600 m, 1 Dec 1967 (st), R. E Smith V910 (VEN); Dtto. parts of the species range, for which the presence of
Jimenez, 10-15 km SSE of Sarare, 1750-1800 m, 7 Aug ants has never been recorded.The materialfrom Ec-
1970 (6), Steyermarket al. 103564 (K, NY, U, US, VEN); uadorand Peru shows some differencesin the length
182 FLORANEOTROPICA

of the hairs on the leafy twigs and longer spikes in minate inflorescence;spikes 1-3, often 2, 6-8 X ca.
the staminateinflorescences. The variationfound is 0.7 cm, to 15 X 1.1 cm in fruit, sessile or with stipes
not unusualin Cecropia species and does not justify to 0.3 cm long; rachis hairy. Pistillate flowers: peri-
recognition of infraspecifictaxa. The longest spikes anth ca. 2.5 mm long, with arachnoidindumentum
sometimes perforatethe apex of the spatheof the pis- below the apex outside, also below the style channel
tillate inflorescencesbefore anthesis. inside, the apex convex, hispidulous; style rather
long, slightly curved;stigma truncateand comose to
Vernacular name. Colombia: yarumo blanco
sublingulate.Fruit oblongoid to ellipsoid, 3-3.5 mm
(Antioquia). long, ? tuberculate.
Distribution (Fig. 11.2). CentralAmazonianBra-
zil, mostly in secondarygrowth.
58. Cecropia ulei Snethlage, Notizbl. Bot. Gart.
Berlin-Dahlem 8: 361. 1923; Acta Amazonica Specimens examined. BRAZIL. AMAZONAS: Mun.
Manaus,rd. BR. 174, km 45, Aug 1978 (Y fl), Benson8465A
8(2): 177. 1978. Syntypes. Brazil. Amazonas, nr.
(UEC, US), 7 Aug 1979 (Y fl), Benson 10341 (INPA,UEC);
Manaus,May 1910 (d), Ule 8838 (B, destroyed),
Manaus,INPA, 9 Sep 1973 (Y), Berg 275 (K, MO, U); Rio
(Y fl-fr), Ule 8838 (B, destroyed;lectotype, here Cuieras, 15 Sep 1977 (d), Berg 277 (U), (Y fr), Berg 278
designated:MG; isolectotypes: G, K, US). (F, U); Manaus-P6rtoVelho Hwy., km 160, 23 Mar 1974
(Y fl-fr), Campbellet al. P20862 (INPA,U); Rio Negro, nr.
Tree,to 4(-8) m tall, usuallyunbranched,the leafy
Rio Arara,SIDERAMA, 2 May 1973 (Y fl-fr), Loureiroet
partof the stem 3-5 cm thick, green,hispid to setose, al. (INPA) 37937 (INPA); Mun. Manaus, Distr. Agropecu-
sometimes also with sparse arachnoidindumentum. ario da SUFRAM, 27 km E of rd. BR.174, km 64, Fazenda
Lamina chartaceousto subcoriaceous,ca. (20 X 20 Esteio, 19 Mar 1981 ( fl-fr), Nascimento et al. 1103.371
cm to) 30 X 30 cm to 80 X 80 cm, the segments 12- (BG, INPA);Rio Madeira,Mun. Humaitd,rd. Humaita-La-
15, oblanceolate, the incisions down to 1.5-5 cm brea,km 50, 29 Nov 1966 (d), Prance et al. 3460 (F, INPA,
from the petiole; apices acute to acuminate (to ob- NY, U, US); Manaus-P6rtoVelho Hwy. BR.319, km 268,
tuse); upper surface ? scabrous,hispidulousto hir- 28 km S of Igap6 A,u, 17 Mar 1974 (? fl-fr), Prance et al.
20649 (INPA,K, MG, MO, NY, S, U, US). PARA:Santarem,
tellous, initially also with sparsearachnoidindumen-
Serra Diamantina, 14 Dec 1966 (? fr), Cavalcante et al.
tum; lower surface hirtellous on the veins, with 1757 (MG); Faro, Castanhalda B6a
Vista, 1 Feb 1910 (6),
arachnoidindumentumin the areoles, sometimes in- Ducke (MG) 10632 (MG); Serrado Humaitd,Rio Tapajoz,
tially also on the main veins; lateralveins in the free Boinu, 8 Apr 1924 (Y fl-fr), Kuhlmann1941 (RB 19881);
part of the midsegment ca. 20-25 pairs, submargin- Rio Trombetas,Monte Branco, 6 Oct 1982 (6), Revilla et
ally loop-connected, unbranchedor some branched; al. 6976 (INPA).
petiole 40-75 cm long, hispidulous to subsetose, This species is distinguished by its predominantly
sometimes also with sparse arachnoidindumentum; monocaul habit and the small size of the trees. It
trichilia fused, the brown indumentum intermixed shows some affinities to Cecropia silvae.
with ratherlong white hairs; stipules 8-13 cm long,
red-brownto grayish, hirtellous and with arachnoid Vernacular name. Brazil: imbauba roxa (Ama-
indumentumoutside, subsericeous inside. Staminate zonas).
inflorescencesmostly in pairs,the peduncleerect and
the spikes pendulous;peduncle 4-10 cm long, hispi-
dulous to subsetose;spathe 8-12 cm long, red-brown 59. Cecropia utcubambana Cuatrecasas, Phytologia
to grayish, hirtellous to subhispid outside, glabrous 52: 157. 1982; Berg & Franco Rosselli, Fl. Ec-
inside; spikes 2-5, often 4, 6-13 X 0.6 cm, with uador 48: 51, t. 9. 1993. Type. Peru, Amazonas:
stipes ca. 0.5 cm long and only with brown pluricel- Prov. Bongara, Rio Utcubamba, 4 km below Cam-
lular trichomes;rachis hairy. Staminateflowers: per- pamento Ingenio, 1250 m, 28 Jan 1964 (Y),
ianth tubular,2-2.5 mm long, glabrous,the apex al- Hutchinson & Wright 3844 (holotype: UC-n.v.;
most plane, the aperture surrounded by a rim; isotypes: G, K, LE, MICH, MO, NY, P, US).
filaments ? swollen; anthers0.5-0.7 mm long, ap- Fig. 46
pendiculate,detachedat anthesis(?),reattachedto the
Cecropiapuberula C. C. Berg & P. Franco, Novon 6:
marginsof the apertureby the appendages(?). Pistil- 248, t. 12. 1996. Type Peru. Ucayali: San Miguel de
late inflorescencessolitary or in pairs, erect; pedun- Semuya,S of CampVerde(at km 33 on rd. Pucallpa-
cle 7-10 cm long, hispidulousto subsetose;spathe7- Lima), 12 Aug 1988 (d), Berg et al. 1639 (holotype:
9 cm long, the color and indumentumas in the sta- MOL; isotypes: AAU, BG, COL, MO, NY, U).
SYSTEMATICTREATMENT 183

199'1

FIG. 46. Cecropia utcubambana.1. Lamina, reduced (Berg et al. 1640). 2. Apex of lamina and venation (Berg et
al. 1639). 3. Stipules and bases of petioles with trichilia (Berg et al. 1640). 4. Staminateinflorescences, one with spathe,
the other at anthesis, and bases of petioles with trichilia. 5. Staminateflower. 6. Stamen after detachmentof anther(Berg
et al. 1639). 7. Pistillate inflorescenceat anthesis and base of petiole with trichilium(Berg et al. 1640). 8. Pistillate flower.
9. Fruit(Berg et al. 1623). (From Novon 6: 247. 1996, modified.)
184
Tree, to 15(-25) m tall. Leafy twigs 1.5-3.5(-5)
cm thick, green (or partlybluish), puberulousto sub-
hispidulous with curved to uncinate hairs. Lamina
chartaceousto subcoriaceous,ca. (10 X 10 cm to) 25
x 25 cm to 60 X 60 cm, the segments 7-9(-1 1), the
free part of upper segments oblong to subobovateor
to obovate, the incisions down to 5/10-6(-7/1)0; api-
ces obtuseto short-acuminate;uppersurfacescabrous
to scabridulous(to almost smooth),puberulousto his-
pidulous (to hirtellous); lower surface minutely pu-
berulous,mostly also with longer uncinateto straight
hairs on the veins, with arachnoidindumentumin the
areoles or (almost) confined to the margin; lateral
veins in the free part of the midsegment9-16 pairs,
submarginallyloop-connected,some or most of them
branched;petiole 15-50 cm long, greenor red-brown,
minutely puberulous;trichilia fused, the brown in-
dumentum intermixed with sparse, (rather) short,
white hairs; stipules 4-12 cm long, red-brown to
brown,darkred to purplish,or greenish,minutelypu-
berulous or on the ribs sparsely hirtellous outside,
sparsely to densely sericeous with white (or brown-
ish) hairs inside. Staminate inflorescences in pairs,
pendulous;peduncle (6-)10-21 cm long, puberulous
and with very sparse arachnoidhairs; spathe 7-26
(-32) cm long, yellowish brownto greenish,darkred
to orange, or grayish, puberulousto sparsely strigil-
lose to subhirtellous or also with sparse to dense
arachnoidindumentumoutside, (almost) glabrousin-
side; spikes 4-7(-15), (4-)10-22(-3 1) X 0.2-0.5
(-0.8) cm, sessile (or with stipes 0.8-1.8 cm long and
hairy); rachis glabrous or sparsely hairy. Staminate
flowers: perianthtubular,0.8-1.5 mm long, with short
hairsjust below or also on the marginof the apex, the
apex slightly convex to plane; filaments flat; anthers
0.4-0.7 mm long, not appendiculate,detachedandre-
maining attachedby 2 filiform connections between
the connective and the uppermarginof the filament.
Pistillate inflorescencesin pairs,pendulous;peduncle
15-50 cm long, puberulousto subhispidulous;spathe
9-20 cm long, the color and indumentumas in the
staminate inflorescence; spikes 4-5(-6), 10-20 X
0.8-1 cm, to 35 X 1.2 cm in fruit,(sub)sessile;rachis
hairy to subglabrous.Pistillate flowers: perianthca.
2 mm long, with arachnoidindumentumbelow the
apex outside, also below the style channelinside, the
apex sparsely to densely muriculate; style rather
short; stigma penicillate. Fruit ellipsoid, 2-2.5 mm
long, smooth, the upper and lower part dark brown,
the middle partpale brown.
Distribution (Fig. 11.3). Southern Ecuador to Cecropia utcubambana and C. puberula, consid-
southern Peru (Madre de Dios), in forest (margins) ered to be morphologically, ecologically, and geo-
and secondarygrowth, at elevations to 2000 m. graphically distinct (Berg & Franco Rosselli, 1996),
FLORANEOTROPICA
Representative specimens examined. ECUADOR.
MORONA-SANTIAGO:Between Plan del Milagro & Lim6n
(Gral. Plaza), 1500 m, 30 Jan 1981 (i), Berg 1235 (AAU,
BG, COL, MO, QCA, U). ZAMORA-CHINCHIPE:Rd. Loja-
Zamora, between Sabanilla & Zamora, ca. 1250 m, 3 Jan
1991 (? fl-fr), Berg et al. 1654 (BG, LOJA,QCA); rd. Val-
ladolid-Rio Palanda,ca. km 6-8, ca. 1500 m, 31 Jan 1985
(Y fl-fr), Harling et al. 21363 (BG, GB, QCA); El Padmi,
5 Jun 1995 (Y fl-fr), Merino et al. 4514 (BG, LOJA).
PERU. AMAZONAS: Prov. Bongara, Dtto. Sipabamba,
above QuebradaFortuna,on trail to Subida Alba, ca. 1350
m, 5 May 1981 (st), Younget al. 341 (BG, NY). Cuzco:
Prov. Paucartambo,between Pilcopata & Atalaya, 29 Jul
1988 (Y), Berg et al. 1604 (BG, COL, Centrode Medicina
Andina, Cuzco); Prov. Paucartambo,between San Jorge &
Salazar,Rio Tono, ca. 600 m, 1 Aug 1988 (Y), Berg et al.
1623 (BG, Centrode MedicinaAndina,Cuzco). HUANUCO:
Rd. Huhnuco-TingoMaria,nr. TambilloGrande(km 515),
24 Nov 1997 (Y fl-fr), Berg et al. 1741 (BG, COL, MOL);
nr. Tingo Maria,Cueva de las Lechuzas, 25 Nov 1997 (c),
Berg et al. 1744 (BG, COL, MOL);Prov.PuertoInca, Dtto.
Yuyapichis,DANTAS, 18 Jul 1989 (Y fl-fr), Kroell S. 508
(BG, G); Prov.Pachitea,ca. 26 km S of PuertoInca, 21 Sep
1988 (? fr), Morawetzet al. 15-21988 (BG); Prov.Leoncio
Prado,Dtto. RupaRupa,E of Tingo Maria,CerroQuemado,
10 Sep 1978 (9 fl-fr), SchunkeV 10593 (BG); Prov.Puerto
Inca, Dtto. Yuyapichis,DANTAS, 1-15 Oct 1990 (9 fl-fr),
Tello 242 (G), 16-30 Jun 1991 (i), Tello 2639 (G). LOR-
ETO:Purancanchim,Rio Sinchiuyacu,22-27 Nov 1986 (9
fr), Lewis et al. 12181 (U). MADREDE Dios: Prov. Tam-
bopata, Cuzco Amaz6nico, 13 Dec 1989 (st), Gentry et al.
68643 (BG, MO), 14 Jul 1991 (9 fl-fr), Fisher 210, 211
(BG), 14 Jun 1989 (9 fl), Niiuez et al. 10727 (BG). PANDO:
Prov.Manupiri,Conquista,23 Aug 2000 (st), Balcdzar2177
(USZ). PASCO:Rd. Oxapampa-Llaupi-Cerrode Pasco, ca.
17 km, nr.MariaTeresa,ca. 1650 m, 29 Nov 1997 (d), Berg
et al. 1760 (BG, COL, MOL); Prov. Oxapampa,Panjil, 12
km from PuertoInca, 27 Sep 1982 (9 fl-fr), D. N. Smithet
al. 2422 (BG). SANMARTiN:Rio Huallaga,Cerros de Es-
toraqui,W of Shapaja,trail to Tarapoto,2-8 km, 4-7 Aug
1937 (9 fl-fr), Belshaw 3224 (GH, MICH, NY, US); Prov.
San Martfn, rd. Tarapoto-Yurimaguas,ca. km 27, 5 Dec
1997 (9 fr), Berg et al. 1777 (BG, COL, MOL), ca. km 39,
5 Dec 1997 (6), Berg et al. 1780 (BG, COL, MOL); Prov.
San Martin,rd. Tarapoto-Chazuta,km 24, 6 Dec 1997 (c),
Berg et al. 1783 (BG); Prov. Lamas, rd. Tarapoto-Moyob-
amba,ca. km 60, 8 Dec 1997 (9 fr), Berg et al. 1787 (BG);
Prov. Rioja, rd. Rioja-Pomacocha,ca. 50 km, 8 Dec 1997
(9 fl-fr),Berg et al. 1795 (BG, COL, MOL).UCAYALI: San
Miguel de Semuya, S of Campo Verde (at km 33 on rd.
Pucallpa-Lima), 12 Aug 1988 (9 fl-fr), Berg et al. 1640
(BG, COL, K, MO, MOL); rd. Pucallpa-Tingo Maria, ca.
km 149, 26 Nov 1997 (d), Berg et al. 1751 (BG, COL,
MOL);rd. Pucallpa-TingoMaria,ca. km 179, nr.Boquer6n,
26 Nov 1997 (9 fl-fr), Berg et al. 1752 (BG, COL, MOL).
SYSTEMATICTREATMENT 185

had to be mergedas recentcollecting reducedthe dif- arachnoidindumentum,intermixedwith brown plur-

ferences considerably.In the northernpartof the dis-icellularhairs outside, darkred, with very shorthairs
tribution range, the species is submontaneand the on the ribs or glabrous inside. Staminate inflores-
arachnoidindumentumis confined to the margin of cences in pairs, the peduncle erect and the spikes
the lamina. In the southernpartof the range,the spe-pendulous (?); peduncle 7-14 cm long, brown-
puberulousto-subhispidulous, towardthe apex hir-
cies is a lowland element, and it often has arachnoid
indumentumin the areolesof the laminabeneath.The tellous to setulose, also with arachnoidindumentum
species has a green morphand a red one (with red to and dense brown pluricellularhairs;spathenot seen;
red-brownstipules, spathes,petioles, and peduncles).spikesca. 8-10, 12-20 X ca. 0.3 cm, with stipes 0.7-1
This species shows strong affinities to C. latiloba. It
cm long and sparselyhirtellousand with dense arach-
differs from the latterin the indumentumof the stip-noid indumentum;rachis glabrous. Staminateflow-
ers: perianth tubular, 1-1.5 mm long, with dense
ules, the indumentumon the veins at the lower surface
of the lamina, and the lateralveins being submargin-arachnoidindumentumbelow the apex, the apex mur-
ally loop-connectedinsteadof marginally.It is clearly
iculate; filamentsflat; anthersca. 0.6 mm long, at an-
distinct from C. latiloba in its ecology, as it is a true
thesis detached,reattachedto the marginsof the ap-
ertureby the appendages(?). Pistillate inflorescences
upland species. In its habit, the species resembles C.
putumayonis,as brieflydiscussedunderthe latter.The solitary, erect to deflexed; peduncle 7-14 cm long,
pistillate inflorescences with long peduncles (and dark green, with sparse arachnoidindumentumand
toward the base and apex brownish-subhispidulous;
spikes) of one of the Ruiz & Pavon s.n., on sheets at
B and MA, indicatedas C. peltata and in G without spathes not seen; spikes 4-5, ca. 25-45 X 1-1.5 cm
name, might belong to C. puberula. Matchingleaves in fruit, + curved;rachissubglabrous.Pistillateflow-
have not been found among the Ruiz & Pavon col- ers: perianthca. 1.5-2 mm long, with arachnoidin-
lections. dumentumbelow the apex outside, also the lowerpart
of the style channel inside, the apex almost plane,
Vernacular name. Peru: kabeari (Machiguenga,
muriculate; style rather long, straight to slightly
Madrede Dios).
curved,muriculate;stigma comose. Fruitellipsoid to
subovoid, ca. 2 mm long, slightly tuberculate,dark
brown.
60. Cecropia velutinella Diels, Notizbl. Bot. Gart.
Distribution (Fig. 18.4). Ecuador,fromPastazato
Berlin-Dalhem15: 367. 1941. Type.Ecuador.Pas-
Morona-Santiago, and in Peru (San Martin), in
taza:Mera,ca. 1100 m, 2 Dec 1938 ( Y), Schultze-
(sub)montaneforest, at 1200-1700 m.
Rhonhof3068 (holotype:B, destroyed,duplicates
not traced), herewith replaced by: Ecuador. Specimens examined. ECUADOR. MORONA-
Morona-Santiago:Rd. Gualaquiza-Don Bosco, SANTIAGO: Rd. Lim6n(Gral.Plaza)-Cuenca, ca. km 15,
km 19, ca. 1500 m, 16 Feb 1994 (Y), Berg et al. Plan del Milagro, ca. 1700 m, 30 Jan 1981 (juv), Berg 1234
BG). Fig. 47 (AAU, BG, GB, MO, QCA, U); rd. Gualaquiza-DonBosco,
1695 (neotype:QCNE; isoneotype:
km 19, ca. 1500 m, 16 Feb 1994 (juv), Berg et al. 1695A
Tree,to 25 m tall. Leafy twigs 5-13 cm thick,dark (BG, COL); Cord. de Cutucu, 5-10 km from Logronio,
brownto black, white-hirtellousand with darkbrown 1200-1500 m, 7-9 Oct 1975 (d), Little et al. 651 (COL,
to blackpluricellularhairs.Laminasubcoriaceous,ca. LOJA,QAME,QCNE).
50 X 50 cm to 130 X 130 cm, the segments 14-20 PERU.SAN MARTiN: Prov. Rioja, rd. Rioja-Pomaco-
cha, km 402. ca. 1350 m, 9 Dec 1997 (Y fr), Berg et al.
(-24), lanceolate to linear, the incisions down to the
1804(BG,COL,MOL).
petiole; apices acute;uppersurfacesmooth or scabri-
dulous, minutelypuberulousand initially with sparse This spectacularsubmontanespecies can be easily
brown pluricellular hairs; lower surface with very recognized by the numerous narrow leaf segments
dense arachnoidindumentum,on the main veins (in- with more lateral veins than in any other Cecropia
itially) also with brown pluricellular hairs; lateral species. Mullerian bodies are abundantlyproduced,
veins in the midsegment50-80 pairs, submarginally but the trees are probably rarely inhabitedby ants.
loop-connected, mostly unbranched;petiole 30-100 The absence of ants may be relatedto the fact thatthe
cm long, with dense arachnoidindumentumand with stems of young trees are covered by densely brown-
brownpluricellularhairs, the upperpart(pale brown) hirsuteindumentumand (sub)persistentstipules cov-
hirsute to subvillous; trichilia fused, the dark brown ered with very dense and matted,white-arachnoidin-
indumentumintermixedwith shortgrayishhairs;stip- dumentum mixed with filiform brown-pluricellular
ules 25-60 cm long, white to brownish, with dense hairs.
186 FLORA NEOTROPICA

i:

FIG. 47. Cecropia velutinella. 1. Lamina, reduced. 2. Apex of lamina and venation (Berg 1234). 3. Stipules and
young leaf. 4. "Umbilicus"of lamina. 5. Interiorof leafy twig (Berg et al. J695A). 6. Pistillate inflorescenceafter anthesis.
7. Pistillate flower. 8. Fruitwith remnantof style (Berg et al. 1695).
SYSTEMATICTREATMENT 187

Vernacular names. Ecuador:su (Shuar,Morona- convex, muriculate,the aperturewide andround,sur-

Santiago);guarumoblanco (Pastaza). roundedby a rim; style rathershortthe apex slightly
convex, muriculate,with a wide round aperturesur-
roundedby a rim; stigma comose. Fruit ellipsoid to
61. Cecropia virgusa Cuatrecasas, Revista Acad. ovoid, ca. 2 mm long, tuberculate.
Colomb. Ci. Exact. 6(22/23): 286. 1945; 7(27): t.
Distribution (Fig. 8.1). From northwesternEc-
5.2. 1947. Type. Colombia. Valle: Between El
uador to eastern Panama,in wet forest or secondary
Aguacate & Quebradade La Yuca, 8 Feb 1944
growth, at low elevations.
(d), Cuatrecasas16093 (holotype:COL;isotype:
F). Specimensexamined.PANAMA:DARIEN: Cruce de
Mono, ParqueNacional Darien, 5 Nov 1989 (st), Fisher 55
Tree,to 15 m tall. Leafy twigs 2-4 cm thick, green (BG).
to purplishbrownwith conspicuouslenticels, sparsely COLOMBIA. CAUCA:Rio Dagua, Juntas,21 Sep 1882
puberulousto hispidulous.Laminasubcoriaceous,ca. (i), Lehmann1969 (G). CHOC6:Ca. 20 km SE of Quibd6,
30 X 30 cm to 80 X 80 cm, the segments (6-)7-8, 8 Jul 1986 (Y fl), Berg 1542 (BG, COL);nr.Tutunendo,ca.
the free parts of upper segments ovate, the incisions 25 km ENE of Quibd6, 9 Jul 1986 (i), Berg 1545 (BG,
down to 3/10-6/10; apices subacuminateto rounded; COL); Serraniade Baud6, rd. Las Animas-Pato, Rio Pato,
uppersurfacealmostsmoothto scabridulous,sparsely 18 Apr 1983 (6), Croat56100 (BG, JAUM,MO); between
hispidulous, on the main veins and the "umbilicus" Quibd6 & Tutunendo,4 Apr 1958 (Y fl-fr), Cuatrecasaset
sparsely hirtellous to subhirsute; lower surface al. 24221 (COL, US); rd. Quibd6-Yuto, nr. Yuto, 12 Apr
1990 (Y fl), Franco et al. 3057 (COL); rd. Yuto-Llor6, 18
sparselypuberulouswith brownpluricellularhairson
Jan 1979 (f), Gentryet al 24376 (BG, COL, MO);between
the veins, with arachnoidindumentumin the areoles
Andagoya & Condoto, 9 Sep 1949 (Y fl-fr), Idrobo 1949
and extending to the main veins; lateralveins in the
(COL); Bahia Solano, QuebradaJellita, 22 Feb 1939 (st),
free partof the midsegmentca. 10-15 pairs, submar- Killip et al. 33551 (US); Novita, Mar 1853 (st), Triana1865
ginally loop-connected, the lower ones usually (K). NARINO: Mun. Barbacoas,nr.Guayacana,12 Dec 1993
branched;petiole 15-40 cm long, sparselypuberulous (d), Franco et al. 5180 (BG, COL);rd. Junin-Barbacoas,7
to subhispidulousand with ratherdense arachnoidin- Jan 1989 (Y fl), Gentry et al. 64565 (PSO); Mun. Tumaco,
dumentum;trichilia separate,sunken into 2 pockets ResguardoIndigenaAlto Albi, 30 May 1992 (st), M. S. Gon-
with slit-shaped apertures; stipules (7-)10-26 cm za'lez115 (COL); La Guayacana,3 Jun 1986 (6), Leon et
long, reddish to brownish or to purplish, ? curved, al. 1245 (U). VALLE: Rio Calima, La Trojita, 19 Feb-10
sparsely hirtellous and with dense arachnoidindu- Mar 1944 (Y fl), Cuatrecasas 16716 (MO, VALLE);Rio
mentumand with brown pluricellulartrichomesout- Cajambre,Barco, 21-30 Apr 1944 (6), Cuatrecasas17266
side, densely pale yellow-sericeous inside. Staminate (F, US, VALLE);Rio Digua, C6rdoba,14 Nov 1945 (Y fl),
Cuatrecasas19859 (F, US); RIo Calima,between Qubradas
inflorescencesin pairs,erect or + deflexed;peduncle
de Aguaclara & La Brea, 21 May 1946 (Y fl-fr), Cuatre-
7-11 cm long, hispidulousor also with sparsearach-
casas 21170 (F, US, VALLE);Rio Calima, nr. La Brea, 25
noid hairs;spathe7-12 cm long, reddishto brownish May 1946 (st), Cuatrecasas 21314 (F, US); Rio Calima,
or to purplish,sparselyhirtellousto strigose and with Quebradade L6pez, 23 Sep 1961 (Y fl-fr), Cuatrecasaset
dense arachnoidindumentumandbrownpluricellular al. 26043 (COL, P, US); 16 km NW of Buenaventura,25
trichomes outside, densely yellowish-sericeous in- Apr 1987 (st), Faber-Langendoenet al. 256 (MO); nr. Bue-
side; spikes 2-4, 4-13 X 0.3-0.7 cm, sessile; rachis naventura,15 Mar 1994 (? fl-fr), Franco et al. 4534 (COL,
hairy. Staminateflowers:perianthtubular,1-1.5 mm TULV); Rio Calima, ca. 6 km N of Buenaventura,13 Dec
long, with stiff hairs on the ribs below the apex, the 1981 (Y fl), Gentry35641 (BG, COL, JAUM,MO, TULV);
apex plane to slightly convex, glabrousor muriculate; Bajo Calima, Canalete, 15 Jun 1968 (Y fl), Gentry et al.
anthersca. 0.4-0.5 mm long, appendiculate,detached 62936 (BG); Rio Dagua, 10 May 1922 (juv), Killip 5352
at anthesis (?). Pistillate inflorescencesin pairs, pen- (NY, US); Mun. Buanaventura,1970 (st), Mahecha s.n.
dulous (?), subtendedby caducous bracts, 0.5-5 cm (UDBC 11162); Bajo Calima, 1970 (Y fl-fr), Mahecha s.n.
(UDBC 11673); Bajo Calima, Concesi6n Pulpapel/Buena-
long; peduncle 5.5-15 cm long, hispidulous or also
ventura,2 Jun 1987 (9 fl), Monsalve B. 1501 (BG, MO);
with sparse arachnoidhairs;spathe 6-8 cm long, the
rd. San Isidro-Juanchaco,15 Nov-6 Dec 1979 (9 fl-fr),
color and indumentumas in the staminateinflores- Rooden et al. 515 (BG, COL, U).
cence; spikes 2-4, 3.5-6 X 0.4-0.5 cm, to 11 X 1.5 ECUADOR.ESMERALDAS:Between Mataje& Molina,
cm in fruit, sessile; rachis hairy with straighthairs or 5 Sep 1991 (9 fl), Jaramillo et al. 13742 (COL, GB, NY,
also arachnoidindumentum.Pistillate flowers: peri- QCA);ReservaEcol6gicaCotacachi-Cayapas,
LuisVargas
anth2-2.5 mm long, with arachnoidindumentumbe- Torres,Rio Santiago, 8-14 Dec 1993 (9 fl-fr), Tiradoet al.
low the apex outside, absent inside, the apex slightly 735 (QCNE); Reserva Ecol6gica Cotacachi-Cayapas,Rio
188
17-26Jul1994(st),Tiradoet al. 1109
Santiago,Angostura,
(QCNE).
Cecropia virgusa has an exceptional type of tri-
chilium. The two trichilia are hidden in lateral (de-
pressions) pockets with a slit-shaped opening at the
base of the petiole. The pockets develop as grooves
departingfrom the base of the petiole and become
longer and deeper; this occurs before trichilia are
formed. Tanninis not found in the developing peri-
carps, as usual in the genus. Ants are usually present.
Vernacular names. Colombia: llarumo (Choco);
cosedera teu, tud teu (Awapit, Nariiio); virgusa
(Valle).
EXCLUDED NAME
Cecropia scabra Martius, Flora 24(Beibl. 2): 95.
1841. Type. Brazil. Para':Rio Amazonas,Martius
630(2) (M), a mixed collection consisting of a leaf
of Pourouma,almost certainlya juvenile one of P.
guianensis Aublet subsp. guianensis, and a pistil-
late inflorescence,possibly of Cecropia ulei. The
leaf is here designatedas the lectotype.
DOUBTFUL NAMES
Cecropia ambaci Rojas, Bull. Geogr. Bot. 28: 161.
1918. Type not designatednor traced.Possibly C.
pachystachya.
Cecropia argentea Visiani, Ort. Bot. Padov. 135.
1842. Type not designatednor traced.
Cecropia dentata Klotzsch, Linnaea 20: 533. 1847.
Type. Peru. Huanuco:Mufia, (st), Ruiz & Pav6n
s.n. (holotype:B, not found, destroyed?),probably
not Cecropia.
Cecropiaglauca Rojas, Cat. Hist. Nat. Corrientes80.
1897. Included in Gunneraceae.Type not desig-
nated nor traced.
NOMINA NUDA
Cecropia arenaria Warburg,in Karsten& Schenck,
Vegetationsbilder4(1): t. 2. 1906.
Cecropia bella Pittier, in Pittier, Luces de Febres,
Badillo & Lasser,Cat. Fl. Venez. 1: 258. 1945.
Cecropiafloccosa Pittier,in Pittier,Luces de Febres,
Badillo & Lasser,Cat. Fl. Venez. 1: 258. 1945.
Cecropiaguanipensis Pittier,in Pittier,Luces de Fe-
bres, Badillo & Lasser, Cat. Fl. Venez. 1: 258.
1945.
CecropiajavitensisPittier,in Pittier,Luces de Febres,
Badillo & Lasser,Cat. Fl. Venez. 1: 258. 1945.
FLORANEOTROPICA
Cecropia nigra Pittier, in Pittier, Luces de Febres,
Badillo & Lasser,Cat. Fl. Venez. 1: 258. 1945.
Cecropia solanoensis Pittier,in Pittier,Luces de Fe-
bres, Badillo & Lasser, Cat. Fl. Venez. 1: 258.
1945.
Cecropia paludosa Warburgex Glaziou, Bull. Soc.
Bot. France59(Mem. 3): 645. 1913.
Cecropia stenostachya Warburg, in Karsten &
Schenck,Vegetationsbilder4(1): t. 1-2. 1906.
ACKNOWLEDGMENTS
This study has been supportedby The Research
Council of Norway (NFR) throughits programCon-
servation of Biological Diversity, The Norwegian
Councilfor Science andHumanities(NAVF),andThe
MeltzerFoundationof the Universityof Bergen,mak-
ing (joint) fieldworkand visits to herbariapossible.
The Olaf Grolle Olsen Foundationof the University
of Bergen financedmost of the illustrationsprepared
for the present contribution.Studies by the second
authorhave been supportedby the Universityof Ber-
gen for a stay of some months (1987) to study Ce-
cropia,to Conciencias(Colombia)to carryout further
researchin Bergen and Utrecht(Aug-Dec 1994), Red
Latinoamericanade Botainica(RLB) for a stay at
Universidadde la Plata (Argentina),with special rec-
ognition to J. V. Crisci and J. J. Morone.Universidad
Nacional de Colombia,particularyInstitutode Cien-
cias Naturales (ICN) supportedand provided facili-
ties. Studentsof UniversidadNacional de Colombia
(Bogota'and Medellin) assisted in fieldwork. D. W.
Davidson is thanked for her willingness to write a
chapter on the Cecropia-ant relations and for her
comments on the introductorypart of this contribu-
tion, and D. W. Yu (HarvardUniversity) for infor-
mation about names and use of species by the Ma-
chiguenga people in Madre de Dios (Peru). The
University of Bergen and the NorwegianArboretum
(at Milde) areacknowledgedfor fundingandfacilities
providedto work togetherduringan early and a later
phase of the present study. Fieldwork was essential
for the preparationof the present monograph. It
startedin 1973 and finishedin 1998 andin thatperiod
numerouscolleagues and institutionshave provided
logistical and administrativesupportto one or both of
us, which is gratefullyacknowledged.Some of them
have to be named: C. D. Adams (then at Port of
Spain), S. G. Beck (La Paz), F. Borchsenius(then at
Quito), R. Callejas (Medellin), J. P. P. Carauta(Rio
de Janeiro),W. Devia (Tulua'),R. L. Dressler(thenin
Panama),G. Hatschbach(Curitiba),L. Holm-Nielsen
(then at Quito), J. Jaramillo (Quito), B. Klitgaard
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Ilavicencio),W. Steiner (then at Sapecho), J. Rios T.
(Lima), Centro de Medicina Andina (Cuzco), Her-
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NUMERICAL LIST OF TAXA

1. CecropiaalbicansTrecul 33. C. montana Snethlage
2. C. andina Cuatrecasas 34. C. multisecta P. Franco & C. C. Berg
3. C. angulata I. W. Bailey 35. C. mutisiana Mildbraed
4. C. angustifoliaTrecul 36. C. obtusa Tr6cul
5. C. annulata C. C. Berg & P. Franco 37. C. obtusifolia Bertoloni
6. C. bullata C. C. Berg & P. Franco 38. C. pachystachya Trecul
7. C. chlorostachyaC. C. Berg & P. Franco 39. C. palmata Willdenow
8. C. concolor Willdenow 40. C. pastasana Diels
9. C. distachyaHuber 41. C. peltata Linnaeus
10. C. elongata Rusby 42. C. pittieri Robinson
11. C. engleriana Snethlage 43. C. plicata Cuatrecasas
12. C. ficifolia Snethlage 44. C. polystachya Trecul
13. C. gabrielis Cuatrecasas 45. C. purpurascens C. C. Berg
14. C. garciae Standley 46. C. putumayonis Cuatrecasas
15. C. glaziovii Snethlage 47. C. reticulata Cuatrecasas
16. C. goudotiana Trecul 48. C. sararensis Cuatrecasas
17. C. granvilleanaC. C. Berg 49. C. saxatilis Snethiage
18. C. herthae Diels 50. C. schreberiana Miquel
C. C. Berg & P. Franco
19. C. heterochroma a. subsp. schreberiana
20. C. hispidissima Cuatrecasas b. subsp. antillarum (Snethlage) C. C. Berg & P.
21. C. hololeuca Miquel Franco
22. C. idroboi Cuatrecasas 51. C. sciadophylla Martius
23. C. insignis Liebmann 52. C. silvae C. C. Berg
24. C. kavanayensis Cuatrecasas 53. C. strigosa Trecul
25. C. latiloba Miquel 54. C. subintegra Cuatrecasas
26. C. litoralis Snethlage 55. C. tacuna C. C. Berg & P. Franco
27. C. longipes Pittier 56. C. telealba Cuatrecasas
28. C. marginalis Cuatrecasas 57. C. telenitida Cuatrecasas
29. C. maxima Snethlage 58. C. ulei Snethlage
30. C. megastachya Cuatrecasas 59. C. utcubambana Cuatrecasas
31. C. membranacea Trecul 60. C. velutinella Diels
32. C. metensis Cuatrecasas 61. C. virgusa Cuatrecasas

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Acevedo R., P. [et al.], 2845 (SOb), 2956 (50a), 7490 3114 (35).
(31). Aguilar, 0. T., 508 (38).
Acosta C., S., 1217 (37). Ake Assi, L. et al., 3960 (41).
Acosta Solis, M., 5848 (6), 12777 (4), 16221 (23). Alarc6n, R., 88 (11).
Adalardo-Oliveira, A., 2771 (12). Albert de Escobar, L., 984 (26).
LIST OF EXSICCATAE 197

Albuquerque,B. W. [et al.], 5 (45), 8 (51), 13 (8), 311 18867 (28), 18877 (46), 18892 (12), 18944 (28),
(25). 19007 (26), 19039 (29), 19204 (2), 19205 (2), 19215
Alexiades,M. [et al.], 290 (11), 353 (31), 603 (31), 807 (4), 19228 (2), 19233 (28), 19349 (4), 19400 (28),
(11), 997 (31), 1097 (31). 19432 (46).
Allen, P. H., 4311 (37), 4422 (37), 5302 (37), 5937 (37), Assungao, P.A.C.L. et al., 3204.2267.2 (45), 3209.3028
6543 (41), 6545 (37), 6688 (23). (9), 3304.6610 (58).
Almeidade Jesus,J., 2017 (15). Atahuachi, M. et al., 23 (31), 24 (51).
Alonso, F. et al., 10193 (47). Atwood, J. T. et al., 70 (41).
Alston,A.H.G.et al., 140 (38). Aubreville, A., 15 (36), 23 (41), 354 (25).
"Aluisio,"72 (25). Aulestia, C. et al., 1385 (37).
Alvarez,A., 740 (26), 877 (2), 881 (4), 2065 (4), 2269 Aulestia, M. et al., 2826 (51).
(18). Aviles, S., 61a (37).
Alves, E. S., 7 (15), 8 (15), 9 (15), 22 (15), 23 (15), 25 Axelrod, F. et al., 2062 (SOa), 2427 (SOb), 2787 (SOb),
(15). 3772 (SOa),4921 (SOb).
Amaral,I. L., 2082 (51). Ayala, F. (et al.), 689 (12), 2901 (31), 3310 (25), 5791
Amorin,A.M.A.et al., 1475 (38). (25), 5930 (25).
Amorozo,M. C., 240 (39). Aymard C., G. et al., 811 (41), 3655 (41), 4146 (3).
Ancuash,E., 50 (31), 1020 (46), 1020 (46), 1094 (46).
Andeer,M., s.n. (38). Bacariqa, ? et al., 15 (38).
Anderson,A. B. et al. 2073 (39), 2181 (39). BAFOG (Bureau Agriculture Forestier Guyanais), 39M
Anderson,A. I. et al., 1450 (8). (51), 7561 (51).
Anderson,W.R. [etal.], 2184 (38), 3000 (41), 3001 (41), Bailey, I. W., 3 (3), 4 (3), 5 (3), 6 (3), 8 (3), 9 (3), 12
3010 (41), 3020 (41), 6552 (49), 7495 (49), 9258 (51), 13 (51), 14 (3), 16 (51), 17 (3).
(49), 10370 (38), 10556 (38), 36458 (49). Bailey, L. H., 864 (41).
Andrade,J. Cardosode [et al.], 14 (38), 15 (38), 16 (38), Baird, J. N., 1000 (37), 1004 (37), 1013 (41), 1017 (41),
17 (38), 19 (38), 21 (38), 22 (38), 23 (38), 24 (15), 1018 (41), 1020 (41), 1021 (41), 1023 (41), 1024
25 (15), 27 (38), 29 (38), 31 (38), 32 (38), 48 (15), (41), 1025 (41), 1027 (41), 1029 (37), 1038 (37),
61 (38), 76 (15), 77 (15), 101 (38), 102 (38), 103 1041 (37), 1042 (37).
(38), 104 (38), 105 (38), 106 (38), 107(38), 108(38), Baker, C. F., 2007 (41).
110 (38), 111 (38), 114 (38), 118 (15), 122 (38), 129 Baker, M. A. [et al.], 6317 (28), 6724 (33).
(15). Baksh, Y. S. et al., 321 (41).
Andrade,R., 33149 (31). Balansa, B., 2575 (38).
Andre,E., 2574 (56). Balcazar, J. [et al.], 1033 (44), 2177 (59).
Andreata,R.H.P.et al., 937 (15). Balee, W. L. [et al.], 408 (51), 809 (8), 1830 (36), 2670
Angeli, C., 20 (15), 239 (38), 326 (15), 344 (15), 345 (8), 2783 (36), 2801 (51).
(15), 347 (15), 48 (15), 50 (21). Balick, M. et al., 1811 (41), 2183 (41), 2630 (37).
Antonio,T., 1309 (20). Balslev, H. (et al.), 1984 (13), 2280 (31), 84740 (26),
Appun,C. F., 1710 (25). 97160 (26), 97161 (26), 97162 (25).
Arana, A., 7 (11), 74 (51). Bamps, P., 5018 (38).
AraqueM., J. et al., 159 (37). Bang, M., 2260 (10).
Araquistain,M. [et al.], 1718 (37), 3033 (37), 3244 (37). Barbosa, C., 1191 (23), 1194 (23), 1334 (31).
Araujo,D. S. Dunnde [et al.], 224 (15), 608 (38), 4918 Barbour, P. J., 5820 (11).
(38), 5016 (38), 5246 (38), 9055 (38). Barclay, A. S. [et al.], 623 (51), 2730 (41), 3197 (48),
Arbo,M. M. et al., 1463 (38), 5346 (49). 3506 (35), 3557 (41).
Arbocz,G. et al., 33417 (15). Barclay, G. W., 2064 (41).
Archer,W.A., 588 (41), 7645 (39). Barfod, A. et al., 48433 (37), 48615 (13).
Argent,G. [et al.], 6426 (38), 6705 (38). Barkley, F. A. [et al.], 17C675 (13), 38C212 (41), 39574
Aristeguieta,L. et al., 1482 (41), 2308 (41), 7435 (25), (41).
7531 (41). Barlow, F. D., 30 (41).
Amason,T. et al., 17016 (41). Barr, R. J. et al., 62-305 (37).
Arroyo,L. et al., 326 (9). Barrera, E., 228 (4).
AscencioR., J., 140 (41). Barringer, K. et al., 1607 (4).
Asplund,E., 7849 (2), 9140 (28), 9198 (12), 9261 (31), Barros, F. de [et al.], 1929 (38), 2382 (15), 2385 (21),
9296 (51), 9301 (46), 9419 (51), 10150 (2), 10290 2469 (38), 2585 (38), 2694 (38).
(11), 12446(44), 12928(4), 13311(51), 14118(12), Barroso, G. M., 282 (38).
14166 (12), 14177 (12), 14350 (25), 14413 (9), Bartolomeu, J. [et al.], 12999 (38), 13000 (38), 15193
14499 (25), 14534 (25), 14731 (25), 14735 (25), (15).
14810 (26), 15377 (26), 15548 (37), 16167 (29), Bastos, M., 23 (36).
16402 (47), 17579 (26), 18745 (4), 18829 (12), Bausen, E., s.n. (21).
198
BBS (boschbeheerSuriname),4 (25), 223 (25).
Beard,J. S., 180 (SOa).
Beck, H. T. et al., 32 (36), 2237 (37).
Beck, S. G. et al., 2504 (8), 3329 (8), 3793 (44), 5292
(25), 5933 (8), 5934 (8), 9931 (11), 12082(4), 13549
(10), 13913 (44), 16603(31), 16662 (11), 16663(8),
19266 (51), 19266 (51), 19665 (8), 19723 (10),
20085 (51), 20092 (51), 20168 (12), 20424 (51),
21207 (44).
Begazo,N., 48 (51).
Be1anger,G. P., s.n. (50a).
Bel6m,R. P. et al., 2074 (38).
Belshaw,C. M., 3224 (59).
Beltran,W., 13 (4).
Benavides,0. de, 10102 (14), 11138 (6).
Bennett,B. C. et al., 3801 (11), 4474 (46).
Benoist,R., 279 (36), 442 (51).
Benson, W., 7954 (38), 8304 (12), 8465A (58), 10341
(58), 10345 (45), 10349 (9), 10350 (8), 10352 (25),
10836 (38), 10837 (15), 10838 (15), 10877 (38),
10878 (21).
Bequaert,J., 2 (8), 3 (45), 14 (37).
Berendson,?, 311 (37).
Berg, C. C. [et al., p.p. et P. Francol,244 (38), 245 (8),
246 (8), 249(45), 252 (39), 253 (39), 264 (45), 275
(58), 277 (58), 278 (58), 279 (8), 280 (8), 281 (45),
282 (9), 284 (39), 298 (20), 299 (19), 391 (37), 393A
(19), 393 (19), 394 (19), 397 (41), 398 (41), 401 (19),
402 (20), 410A (23), 410 (27), 416 (13), 417 (4), 418
(4), 422 (47), 430 (37), 432 (37), 439 (2), 441 (4),
585 (36), 586 (39), 773 (9), 784 (39), 949 (37), 978
(41), 979 (41), 984 (41), 1032 (25), 1043 (26), 1046
(26), 1056 (25), 1057 (25), 1066 (12), 1067 (51),
1071 (12), 1072 (28), 1073 (28), 1074 (28), 1075
(11), 1089 (12), 1090 (31), 1091 (28), 1092 (31),
1093(46), 1104(11), 1117(4), 1134(15), 1135 (38),
1135 (38), 1136 (15), 1142 (21), 1146 (15), 1149
(38), 1149(38), 1201(40), 1204(4), 1215(46), 1217
(28), 1224(12), 1229(57), 1232(57), 1233(4), 1234
(60), 1235 (59), 1236 (40), 1237 (26), 1238 (26),
1239(20), 1240(23), 1242(37), 1244(47), 1248(4),
1254 (37), 1260 (23), 1263 (4), 1279 (4), 1280 (47),
1281 (4), 1291 (13), 1526 (SOa),1527 (50a), 1528
(SOa),1533 (50a), 1534 (SOa),1540 (37), 1541 (37),
1542(61), 1545(61), 1549(37), 1550(12), 1553(9),
1562 (25), 1563 (31), 1570 (25), 1571 (25), 1573
(12), 1575(25), 1586(26), 1597(9), 1598(9), 1600A
(31), 1600 (31), 1600B(31), 1600C(31), 1604 (59),
1605(8), 1608A(44), 1608(44), 1609(8), 1610(11),
1610A (4), 1612 (44), 1615 (53), 1617 (53), 1621
(51), 1623 (59), 1626 (9), 1627 (9), 1628 (8), 1629
(11), 1630 (4), 1632 (4), 1633 (4), 1634 (55), 1635
(55), 1636 (11), 1637 (12), 1638 (44), 1639 (59),
1640(59), 1641(44), 1643(25), 1644(18),1645(29),
1646 (4), 1647 (47), 1648 (6), 1650 (4), 1651 (4),
1652 (2), 1653 (2), 1654 (59), 1655 (40), 1656 (33),
1657 (11), 1658 (33), 1659 (28), 1660 (46), 1661
(33), 1662 (33), 1664 (40), 1665 (57), 1666 (57),
1682 (20), 1687 (13), 1688 (4), 1691 (33), 1692 (4),
FLORA NEOTROPICA
1693 (40), 1694 (46), 1695 (60), 1695A (60), 1696
(44), 1697 (8), 1698A(5), 1698 (31), 1699 (5), 1700
(11), 1701 (5), 1703A(8), 1704 (5), 1707A(8), 1708
(8), 1710 (11), 1711 (53), 1712 (53), 1713 (4), 1714
(53), 1715 (4), 1717 (5), 1720 (39), 1721 (8), 1722
(10), 1723 (53), 1724(4), 1725(53), 1726(10), 1728
(8), 1729 (31), 1730 (1), 1731 (55), 1732 (1), 1733
(40), 1734(53), 1735(44), 1736(40), 1737(4), 1738
(40), 1739 (53), 1740 (44), 1741 (59), 1742 (33),
1743 (33), 1744 (59), 1745 (53), 1746 (53), 1747
(53), 1748(53), 1749(33), 1750(4), 1751(59), 1752
(59), 1753 (4), 1754(53), 1755(53), 1756(53), 1757
(1), 1758 (1), 1759 (4), 1760 (59), 1761 (40), 1762
(4), 1763(40), 1764(53), 1765(11), 1766(11), 1768
(33), 1769 (44), 1770 (44), 1771 (31), 1772 (53),
1773 (33), 1774 (46), 1776 (12), 1778 (12), 1780
(59), 1781(9), 1782(31), 1783(59), 1787(59), 1788
(33), 1789 (12), 1790 (53), 1792 (11), 1793 (11),
1794 (40), 1795 (59), 1796 (2), 1797 (2), 1798 (4),
1799 (7), 1800 (1), 1801 (4), 1802 (7), 1804 (60),
1805(4), 1808(23), 1809(41), P.17584(31), P.17585
(31), P.17586 (25), P.17590 (8), P.18419 (12),
P.18526 (12), P.18808 (45), P.19769 (25), P.19817
(8), P.19818(8), P.19839(9), s.n. (8, 13).
Bergeron,S. et al., 753 (12).
Berhaut,R. P.,984 (41).
Berlin,B., 229 (12), 592 (11), 736 (9), 2041 (51), 2057
(1 1).
Bermudez,?, 34923 (35).
Bernacci,L. C. [et al.], 62 (38), 193 (38), 194 (15), 823
(38).
Bernal,R. et al., 359 (14), 952 (37), 997 (43), 2039 (25).
Bernardi,A. L., 416 (41), 826 (51), 1295 (4), 1313 (48),
1612 (9), 1966 (4), 2031 (48), 2072 (57), 2943 (57),
3043 (3), 3298 (41), 5776 (41), 6878 (4), 7109 (41),
7629 (4), 16330 (9).
Bertero,C. G., s.n. (SOa).
Betancur,J. et al., 1255 (27), 1256 (37).
Biesmeijer,J. C. et al., 188 (41).
Bigay, ? et al., DI.805 (31).
Black,G. A., 51-14103 (36).
Blackmore,S. et al., 1655 (37), 3797 (41).
Blanchet,J. S., 464 (38), s.n. (15).
Blanco,C. A., 616 (3), 790 (41).
Blum, K. E. [et al.], 1169 (41), 1383 (37), 1861B (37),
1862B (41), 2054 (37), 2097 (19), 2098 (37), 2099
(20), 2153 (23), 2154 (41), 2157 (41), 2158 (37),
2176 (23), 2176 (41), 2205 (19), 2206 (19), 2216
(37), 2217 (37), 2218 (23), 2225 (4), 2230 (37), 2241
(27), 2443 (41), 2496 (37), 2515 (41), 2529 (41),
2530 (20), 2531 (37), 2544A (37), 2600 (4), 2607
(37).
Bohlin,J.-E.et al., 1468 (12).
Boldingh,I., 254 (50a),904 (SOa),955 (SOa),2158 (SOa).
Bono, J., 4317 (57).
Bonpland,A.J.A, 1295 (SOb),s.n. (38).
Boom, B. M. [et al.], 2086 (36), 2161 (36), 2162 (36),
4257 (51), 4264 (12), 4846 (12), 5252 (9), 6493 (41),
LIST OF EXSICCATAE 199

6894 (SOa),7190 (41), 7943 (SOa),8533 (45), 8718 11161 (25), 11183 (25), 15832 (41), P.20862 (58),
(51). P.22441(25).
Borges,J. R. et al., 101 (38). Carauta,J.P.P.[et al.], 175 (38), 201 (38), 208 (15), 21 lB
Bosse, G., 7633 (31), 8707 (31), 8797 (41). (15), 225 (38), 237 (15), 287 (15), 296 (38), 301 (38),
Boswaldo,I., 554 (38), 688 (38), 1363 (38). 303 (38), 349 (38), 353 (38), 404 (21), 413 (21),
Bottia,J., 1 (32). 414A (15), 414B (15), 491 (38), 536 (15), 538 (39),
BoudetFernandes,H. 0. [et al.], 8 (38), 1251 (38), 1620 544 (38), 545 (38), 551 (38), 553 (15, 39), 554 (39),
(38), 2623 (15), 2624 (15). 564 (38), 581 (21), 596 (21), 601 (21), 620 (38), 621
Bourgeau,E., 1869 (37), s.n. (37). (38), 622 (38), 623 (15), 624 (15), 644 (15), 645 (15),
Boyle, B. et al., 1574 (47), 1730 (47), 1916 (4), 3304 665 (15), 666 (15), 667 (38), 723 (49), 724 (38), 725
(54). (38), 810 (38), 869 (38), 965 (39), 969A (39), 969B
Brade,A. C., 14130 (21). (39), 1061 (38), 1062 (38), 1286 (15), 1287 (15),
Brand,J., 1083 (37), 1201 (27). 1303 (38), 1467 (38), 1468 (38), 1471 (39), 1474
Bravo,E. et al., 291 (28). (38), 1475 (38), 1502 (38), 1503 (38), 1557 (38),
Breedlove,D. E. [etal.], 9628 (41), 9822 (41), 9912 (37), 1652 (38), 1899 (38), 2000 (38), 2171 (38), 2172
20725 (41), 23943 (37), 28781 (41), 30939 (37). (38), 2328 (21), 2358 (38), 2976 (15), 3016 (8), 3077
Bristan,N., 121 (14). (38), 3297 (38), 3298 (38), 3300 (38), 3302 (38),
Bristol,M. L., 1441 (57). 3303 (38), 3304 (38), 3362 (38), 3363 (38), 3425
Brito, H. S. [et al.], 228 (38), 333 (49), 335 (38), 336 (15), 3425 (38), 3436 (15), 3437 (15), 3438 (38),
(38, 49). 3448 (38), 3450 (38), 3452 (38), 3471 (38), 3472
Britton,N. L. [et al.], 258 (SOb),508 (41), 572 (SOa), (38), 3486 (15), 3487 (15), 3504 (38), 3508 (38),
715 (SOb),1654 (41), 2048 (41), 2790 (41), 7490 3509 (38), 3510 (38), 3511 (38), 3512 (38), 3514
(SOa). (38), 3515 (38), 3575 (38), 3829 (38), 3830 (38),
Broadway,W. E., 259 (41), 582 (36). 3830 (38), 3879 (38), 3893 (38), 4423 (21), 4444
Brown,S. et al., 1683 (41). (38), 4462 (38), 4463 (38), 4463 (38), 4868 (15),
Buchanan-Smith, 3 (51). 4918 (38), 5418 (38), 5419 (38), 5481 (15), 5683
Buchtien,O., 2118 (4). (21), 5772 (38), 5776 (49), 5777 (49), 5800 (38),
Bulhoes,M. et al., 226 (38). 5816 (21), 5943 (38), 6485 (39), 6907 (21).
Bullock,2022 (37). Cardenas,D. [et al.], 679 (41), 897 (23), 1115(37), 1934
Bunting,G. S. [et al.], 1052 (41), 3790 (12), 6341 (41), (41), 4394 (12), 4397 (12), 4411 (12), 4412 (31),
6554 (41), 7323 (41), 9404 (41), 9853 (41), 10029 4509 (12), 4513 (12), 4599 (31), 4600 (25).
(41), 11458 (41), 12087 (4). Cardenas,H., 186 (41).
Burchell,W.J., 1452 (49), 1452 (38), 5146 (38), 5474 Cardenas,M., 1165 (10).
(38), 4553 (21), 6444 (49), 8023 (49). Cardona,F., 1690 (51), 1720 (51), 1951 (24).
Burger,W.C. et al., 4549 (37), 5201 (37), 6934 (37), Carmona,R., 132 (37).
7611 (41), 8428 (4). Carrasquilla,L., 362 (42), 2177 (19).
Burkart,A. E., 27588 (38). Carri6n, A. et al., 133 (9).
Busey,P., 822 (19). Carvalho,A.M.V.de [et al.], 347 (38), 6139 (38).
Buttura,?, 227 (38). Castanio,L. et al., 200 (31).
BW (Boschwezen,Suriname),1533 (36), 1593 (51), Castellanos,A. [et al.], 23456 (15), 23556 (38), 23558
1613 (36), 1621 (36), 1711 (36), 1892 (36), 2241 (38), 23671 (8), 23702 (25), 23712 (31), 23756 (12),
(36), 2967 (36), 3031 (36), 3818 (36), 3836 (51), 25695 (21), 27517 (25), 27525 (8), 27609 (12).
3953 (36), 4208 (36), 4708 (36), 5077 (36), s.n. (51). Castillo,A., 711 (41), 1337 (32), 2911 (12).
Byer,M. S., 66-960 (SOa). Castro,M. et al., 103 (8).
Castroviejo,? et al., 11978 (22).
Cabrera,E. [etal.], 1116(41), 3483 (41), 5099 (41), 5099 Cavalcante,P. [et al.], 207 (25), 1391 (39), 1392 (39),
(41). 1394 (39), 1395 (39), 1396 (39), 1757 (58), 2403
Cabrera,I., 3753 (31). (41), 2475 (41), 3097 (12), 3098 (12).
"CAC,"2482 (38). Cavalcanti,T. B. et al., 1294 (49).
Cadena,A. et al., 3008 (41). Cer6n,C. E. [et al.], 712 (31.note),1659(51), 2149 (28),
Calder6n,S., 104 (37), 1264 (37), 1840 (37). 3332 (51), 3408 (9), 3847 (11), 3860 (51), 4021 (18),
Callejas,R. et al., 4448 (41), 8744 (37), 8753 (20), 8757 4776 (29), 5429 (46), 5615 (12), 6081A (31), 7633
(47), 8793 (37), 10879 (41), 11110 (57), 11111 (4), (46), 8052 (46), 8414 (12), 8593 (13), 8865 (31.note),
11112 (4). 9407 (12), 9417 (51), 9418 (4), 13470 (26).
Calzada,J. I., 927 (37). Chac6n,I. A. et al., 1563 (4), 1816 (4).
Camp,W.H., E.3634 (26). Chagas,J., 138 (8), 1288 (45), 7280 (8).
Campbell,D. G. et al., 6806 (9), 7148 (51), 8098 (12), Chaguaro,?, 36 (51).
8144 (12), 8162 (51), 8197 (31), 9720 (44), 10334 Chalot,C., 48 (41).
(51), 10756(25), 10858(12), 11052(25), 11053(25), Chan,C. et al., 1432 (41).
200
Chavelas,J. et al., 2463 (41).
ChavezA., R., 389 (44), 409 (53), 463 (44), 1721 (44),
3389 (44).
Chickering,A. M., 43 (41).
Christensen,?, 82750 (31), 82794 (31).
Churchill,H. W., 5720 (41).
Clark,H. L., 6879 (12), 6880 (31).
Clark,J. L. et al., 140 (47), 1004 (51), 1121 (28), 1530
(26), 4089 (20), 4751 (20), 4792 (20), 4806 (47),
4885 (20), 4889 (37).
Clarke,D., 3937 (25).
Clarke,O., s.n. (41).
Claussen,P., 21 (38), 158 (38).
Coelho,?, 90393 (39).
Coelho,D. et al., 52385 (51).
Coelho,L. et al., 1795 (44), 1908 (25).
Cogollo,A. C. [et al.], 1405 (41), 3624 (37), 6153 (43),
6158 (43), 6161 (43), 6167 (43), 6194 (43), 6201
(43), 6383 (43), 6611 (43), 6692 (43), 7242 (43),
7612 (43).
Coimbra,C., 71B (11).
Colella,M. et al., 1444 (3).
Combs,R., 111 (50b).
Conant,D. S., 1053 (8).
Contreras,E., 1007 (41), 1390 (41), 5832 (41), 5845
(41).
Converse,O., s.n. (4).
Conzatti,C. [et al.], 3242 (37), 4518 (37).
Cook, 0. F., et al., 3 (41), 336 (41), 813 (SOa).
CorbusierBoland,?, 2061 (38).
Comejo,X. et al., 3578 (47).
CorreaA., M. D. et al., 27 (27), 575 (37), 850 (37), 1806
(19).
CorreaL., G. et al., 384 (9), 392 (9), 407 (9).
Cortes,L. et al., 111 (37).
CortesB., R. et al., 1545 (41), 1546 (23).
Costa,J. C. da et al., 1202.2406(51), 1202.2862(58).
Cowan,C. P., 38280 (36).
Cox, D. K., 561 (37).
Crane,C. L., 54 (41), 499 (41).
Cremers,G. [et al.], 12329(51), 12330(36), 12490(36),
13269 (25), 13270 (25), 15193 (51).
Croat,T. B. [et al.], 1884 (27), 4796 (37), 5030 (41),
5033 (23), 5131 (41), 5282 (37), 6000A (23), 6452
(41), 6712 (37), 6781 (27), 6816 (37), 7023 (23),
7127 (37), 7254 (27), 7421 (37), 7554 (25), 7827
(37), 8143 (37), 8449 (37), 8795 (23), 10113 (41),
10113 (27), 10114 (41), 10115 (27), 10117 (23),
10150 (37), 10386 (37), .11286 (37), 11323 (23),
11496 (41), 11587 (19), 11588 (20), 11716 (37),
11749 (41), 11800 (37), 11831 (41), 12623 (41),
13211 (23), 13887 (19), 15083 (27), 15246 (27),
15248A (27), 17338 (19), 17656 (51), 18128 (12),
18177 (12), 19018 (51), 20818 (31), 21791 (41),
22273 (23), 22562 (37), 23990 (41), 24563 (37),
24887 (37), 24965 (41), 25880 (19), 27627 (19),
34288 (19), 38234 (41), 56100 (61), 63278 (37),
65447 (37), 65546 (37), 70406 (19), s.n. (27).
FLORANEOTROPICA
Croizat,L., 196 (32), 501 (51), 511 (25), 957 (31), s.n.
(25).
Cruxent,J. M., 158 (51).
Cruz,J. S. de la, 3752 (3).
CuadrosV., H. [et al.], 711 (4), 891 (31), 892 (31), 937
(37), 1352 (41), 1372 (41), 1374 (27), 1383 (41),
3402 (41), 4173 (37).
Cuatrecasas,J. [et al.], 3082 (35), 4490 (31), 6503 (16),
6818 (12), 7328 (12), 7479 (12), 7617 (31), 8135 (4),
8334 (35), 8697 (4), 8782 (4), 8816 (12), 8953 (31),
9092 (12), 9106 (51), 9204 (40), 9397 (35), 9410
(41), 9843 (41), 10140(41), 10178(41), 10437(41),
10438 (41), 10551 (51), 10702 (12), 10787 (12),
10867 (31), 11053 (46), 11063 (31), 11069 (18),
11141 (12), 11203 (51), 11243 (28), 11244 (11),
11263 (46), 11308 (33), 11582 (57), 11662 (2),
12346 (57), 12781 (57), 13098 (48), 13197 (48),
13355 (48), 13548A (57), 13689 (37), 13855 (4),
13872 (4), 13947 (56), 13969 (37), 13979 (37),
14266 (37), 14845 (37), 14910 (30), 15111 (54),
15117 (47), 15260 (37), 15469 (43), 15540 (4),
15550 (4), 15988 (37), 16000 (37), 16093 (61),
16096 (23), 16347 (31), 16349 (31), 16467 (14),
16505 (37), 16716 (61), 16830 (23), 16960 (37),
17266 (61), 17268 (37), 17272 (14), 17675 (37),
17681 (37), 18353 (56), 18406 (56), 18785 (4),
18786(4), 18787(4), 19804(14), 19805(37), 19859
(61), 21011(56), 21170 (61), 21242 (31), 21262 (23),
21314 (61), 21315 (23), 21510 (37), 21511 (31),
21577 (31), 21619 (56), 21691 (56), 22015 (4),
22304 (56), 22504 (56), 22511 (4), 22682 (4), 22763
(47), 22891 (4), 22902 (4), 23543 (57), 23553 (57),
23841 (56), 23883 (47), 23884 (13), 23967 (43),
23968 (56), 23986 (20), 23987 (20), 24008 (47),
24008A (47), 24009 (23), 24010 (23), 24050 (37),
24164 (37), 24189 (37), 24192 (37), 24194 (37),
24203 (31), 24204 (37), 24221 (61), 24419 (41),
25020 (41), 25346 (57), 25356 (41), 25393 (4),
25420 (4), 25522 (41), 25535 (41), 26042 (37),
26043 (61), 26107 (31), 26112 (37), 26113 (31),
26114 (37), 26186 (23), 26187 (23), 26213 (41),
26214 (41), 26247 (57), 26248 (57), 26487 (4),
26491 (4), 26508 (57), 26510 (57), 26626 (15),
26627 (15), 26644 (38), 26646 (38), 26653 (39),
26654 (36), 26655 (39), 26658 (39), 26894 (56),
26894 (56), 27125 (12), 27126 (12), 27236 (12),
27240 (31), 27244 (12), 27250 (25), 27940 (57),
27949 (57), 27950 (4), 27953 (4), 27965 (41), 27968
(41), 28946 (56), 30079 (41), 30096 (41).
Curran,H. M. [et al.], 65 (41), 72 (38), 328 (38), 339
(15), 408 (38), 714 (38), 1009 (41).
Curtiss,A. H., s.n. (50b).
CustodioFilho,A., 654 (21).
Dahlgren,?, 51/102 (SOb).
Daly,D. C. et al., 6722 (51), 6972 (12), 7143 (51), 7418,
(12), 7529 (12), 7726 (11), 7727 (51), 8471 (12).
Daniel,Hno., 1230 (41), 2843 (4), 3312 (4), 4492 (57).
Darfo,M., 355 (4).
LIST OF EXSICCATAE
Davidse, G. [et al.], 1221IA (25), 12305 (32), 13258
(25), 14037(25), 14302(32), 14500(25), 14637(32),
14998 (32), 14998 (32), 15555 (32), 15831 (32),
15968 (32), 16969 (9), 20921 (48), 26532 (12),
27161 (12), 27450 (12), 35776 (37).
Davidson,C. [et al.], 7165 (37), 9793 (25), 862 (37),
8990A (37), 10290 (25).
Davis, D. H., 79 (41), 239 (3), 240 (3), 247 (51).
Davis, E. W. et al., 921 (51).
Del Aguila,M. et al., 90 (11), 168 (31), 196 (51).
Delascio C., F. et al., 2578 (41), 2782 (51), 3446 (41),
7845 (41), 8010 (3), 9304 (31), 9318 (12), 10959
(25), 11050 (51), 11720 (32), 12225 (41).
Del Carpio,[et al.], 1634 (25), 2087 (25).
Delgado,L. [ et al.], 20 (41), 241 (41), 720 (25), 961
(12).
Devia A., W., 301 (31), 313 (4), 396 (47), 406 (4), 1267
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Dias, A.T.G.et al., 219 (38).
Diaz, C. et al., 1470 (9), 4567 (7).
Diaz, J., 536 (12).
Diaz P., S. [et al.], 575 (57), 685 (57), 3263 (35).
"Dioscoreabrigade,"3778 (37), 3924 (37), 6577 (37),
6658 (37).
Dodson,C. H. [et al.], 1263 (26), 5579 (23), 5601 (37),
5880 (26), 6233 (26), 6235 (31), 6235 (37), 6237
(37), 6451 (47), 6662 (26), 6668 (47), 6834 (23),
7303 (20), 7304 (47), 7305 (4), 7334 (26), 7335 (4),
7413 (20), 8869 (26), 8941 (4), 8953 (4), 8959 (47),
8960 (47), 9005 (20), 9922 (26), 10541 (4), 11287
(26), 12724 (47), 12769(20), 14841(2), 15011(28).
D6mingez,V., 698 (41).
Dorr,L. J., et al. 5367 (48).
"DPM,"36 (41).
Drees,E. M., 5596 (51).
Dressler,R. L., 4326 (19), 4460 (42).
Duarte,A. Pereira,5485 (38), 6078 (38),7187 (51),7188
(45), 7189 (39), 7190 (8).
Dubs,B., 371 (38), 464 (38).
Duchassaing,D. P., s.n. (41, Sla).
Ducke,A. [et al.], 1690(39), 1910(15), 2615 (39), 3664
(21), 9912 (39), 10199(25), 10564(39), 10632(58),
11711 (25), 11803 (25), 12355 (12), 13071 (38),
19160 (38), 19456 (25), 19457 (25), 19469 (51).
DugandG., A. [et al.], 407 (41), 612 (41), 3357 (41).
Duke,J. A. [etal.],4904 (41), 5355 (31), 5846 (41), 6645
(41), 9883 (37), 11728 (41), 12204(37), 12500(41),
14274 (37).
Dulmen,A. van [et al.], 58 (12), 147 (31), 148 (12).
Dumont,M. et al., 41 (35).
Duno, R. et al., 174 (25).
Duque-Jaramillo, J. M., 2030 (25), 2206 (25), 2590 (35),
2590 (41), 4048 (4).
Dusen,P.K.H.,3542 (15), 11217(15),14043 (15),16456
(38), 17438 (38).
Duss, A., 1405 (50a), 2860 (50a), 3621 (50a), 4207
(SOa).
Dwyer,J. D. [et al.], 147 (41), 553 (37), 3447 (19), 7110
(20), 9885 (37), 11017 (41).
201
Echavarria, J. A. et al., 149 (27), 249 (41).
Eden, M. J., 12A (12).
Edwall, G., 6403 (21).
Eggers, H.F.A., 6829 (50a), s.n. (SOb).
Eggers, L. et al., 3402.5108 (51).
Egler, W. A., 413 (39), 437 (36),439 (51),445 (52),1018
(38), 45948 (25), 47658 (36).
Eiten, G. et al., 6196 (38).
Ekman, E. L., 983 (SOb), 2013 (38), 6548 (SOb), 9376
(SOb), 12324 (SOb).
Elcoro, S., 81 (3).
Elias, Bro., 578 (41), 1230 (41).
Elias, T. S. et al., 1606 (41).
Ellenberg H., 880 (55), 2242 (51), 2296 (51), 2385 (31),
2430 (31), 2820 (31).
Encarnaci6n, F., 990 (25), 25003 (12), 25004 (12), 25005
(12), 25006 (12), 25007 (9), 25008 (12), 25009 (12),
25011 (12), 25012 (9).
Erlanson, C. O., 233 (41).
Ernani Diaz, B., 446 (15), 452 (38), 453 (38).
Ernst, A., 1756 (4).
Ervik, F., 36871 (29).
Escobar, L. et al., 9935 (SOb).
Escobar M., E., s.n. (4).
Espadas, M. et al., 96 (37).
Espina, J. [et al.], 695 (37), 1920 (41), 2745 (41).
Espinal T., S., 2325 (56), 2746 (31).
Espinoza, R., 1575 (57).
Espinoza, S. et al., 379 (51), 585 (51).
Evans, ?, 1551 (37).
Ewan, J. A., 16731 (12), 16732 (11).
Faber-Langendoen, D. et al., 256 (61), 1106 (14).
Falcao, J. I. Almeida de et al., 1155 (38).
Fanshawe, D. B., 731 (51), 732 (3), 2990 (3), 3033 (41).
Farinias,M. [et al.], 349 (31), 378 (51), 501 (11), 502
(25), 506 (11), 689 (51).
FD (Forest Department, British Guiana), 3467 (51), 3468
(3), 6320 (3), 6363 (41), 7769 (25).
Fearnside, P. M., 1019 (44).
Feddema, C., 2112 (41).
Fendler, A., 1239 (41), s.n. (4).
Fernandes, A. et al., s.n (38).
Ferrnndez, A. [et al.], 1965 (12), 2993 (32), 3527 (25),
3964 (3), 4399 (25), 4819 (25), 5007 (3), 5031 (35),
5505 (3), 5505 (24), 6533 (25), 6533 (41), 6775 (25),
9798 (37).
FermnndezC., J. [et al.], 3709 (38), 4133 (38), 4138 (38),
4139 (38), 4171 (38), 5645 (38), 7907 (44), 7913 (8),
8052 (44), 8152 (12), 8163 (8), 8203 (31), 8238 (8),
8241 (44), 8410 (25), 10690 (SOb).
Fermnndez-Perez,A., 20106 (31).
Ferreira, A.J.C. et al., 1101.481 (9), 3209.873 (51),
3209.2016 (51), 3209.2631 (45), 3209.2634 (45).
Ferreira, A. R., s.n. (31).
Ferreira, C. A. Cid et al., 2542A (44), 9836 (9).
Ferreira, E., 58/239 (12), 5929 (8), 6442 (12).
Ferreira, L. V., 109 (25), 180 (25), 322 (25), 1341 (25).
Ferreira,V. F. [et al.], 67 (38), 4219 (38).
202
Ferreyra,R. [et al.], 398 (44), 9030 (31), 9031 (31), 9032
(12), 9033 (31), 9035 (51), 9036 (53), 9037 (33),
9038 (31), 9039 (31), 9040 (31), 9041 (25), 9042
(51), 9043 (12), 9044 (11 and31), 9045 (11), 10186
(44), 13047(12), 13075(12), 15939(41), 16102(12),
17201 (12), 17204 (12), 17205 (12), 17423 (44),
17609 (12), 17621a (12), 17621 (12), 17625 (25),
17629 (11), 17633 (12), 17637 (12), 17639 (12),
17649 (51), 17878 (33), 18132 (12), 17205 (12).
Ferrucci,M. M. et al., 6195 (38).
Feuillet,C., 10312 (9), 10313 (36).
Fevereiro,V.P.B.et al., M.169 (39).
Fisher,B. L., 15 (37), 19 (19), 20 (19), 21 (37), 23 (19),
25 (37), 26 (41), 27 (41), 29 (19), 30 (19), 31 (19),
33 (23), 34 (37), 41 (27), 44 (27), 45 (27), 46 (41),
47 (27), 52 (19), 53 (27), 54 (37), 55 (61), 55 (31),
56 (31), 57 (47), 177 (31), 178 (44), 179 (25), 210
(59), 211 (59).
Fleury,M., 1026 (36), 1063 (36), 1065 (51).
"FloraFalc6n(HW,ES),"460 (4).
FloresN., C., 312 (31).
Focke,H. C., s.n. (41).
Folli, D. A., 310 (21), 1051 (38), 3567 (15).
Folsom,J. P. [et al.], 5226 (41), 6161 (19), 6351 (47),
7766 (41), 8979 (37).
Fonnegra,R. et al., 2070 (37), 4221 (37).
Fonseca,M. L. et al., 1570 (38).
Forero,E. [et al.], 116 (57), 1554 (37), 1871 (41), 1890
(27), 4665 (8), 4969 (37), 6415 (51), 6480 (47), 8682
(38), 10239 (32).
Foresta,H. de, 495 (25).
Forzza,R. C. et al., 291A (45), 1242 (49).
Foster,R. B. [et al.], 1484 (37), 1896 (19), 2293 (41),
8035 (51), 8948 (31), 11350 (25), 11435 (8), 11444
(8), 11526 (44), 11528 (8), 11533 (31), 11534 (11),
11570 (53), 11777 (12), 11875 (53), 11876A (53),
11960 (25), 12052 (23), 12084 (31), 12169 (31),
12189 (51), 12202 (31), 12202 (31), 12410 (8),
12426 (8), 12535 (11), 12825 (31), 12827 (31),
13042 (31), 13284 (31), 14728 (19).
Foumier,L. A., 311 (42).
Franco,P. [et al., p.p. et C. C. Berg], 1192 (37), 1982
(37), 2409 (35), 2441 (35), 2443 (35), 2483 (35),
2485 (35), 2963 (41), 2965 (23), 2968 (41), 2970
(35), 2973 (35), 2973 (35), 2974 (35), 3001 (41),
3006 (23), 3011 (37), 3055 (37), 3057 (61), 3058
(37), 3061 (37), 3062 (37), 3349 (31), 3442 (37),
3508 (37), 3605 (37), 3606 (37), 3607 (14), 4342
(57), 4343 (4), 4344 (35), 4345 (35), 4346 (41), 4347
(41), 4456 (4), 4457 (4), 4457 (4), 4459 (48), 4461
(57), 4462 (47), 4463 (54), 4464 (14), 4465 (32),
4465 (4), 4466 (13), 4467 (6), 4468 (4), 4469 (13),
4471 (13), 4474 (47), 4475 (4), 4477 (4), 4479 (56),
4481 (4), 4482 (54), 4485 (30), 4486 (30), 4487 (31),
4488 (4), 4489 (37), 4490 (56), 4493 (4), 4494 (57),
4495 (4), 4496 (4), 4498 (57), 4502 (4), 4503 (2),
4505 (35), 4506 (35), 4507 (51), 4508 (28), 4509
(46), 4512 (40), 4513 (4), 4514 (46), 4515 (4), 4516
(11), 4517 (11), 4519 (12), 4520 (57), 4521 (57),
FLORANEOTROPICA
4522 (4), 4523 (4), 4525 (4), 4526 (4), 4527 (4), 4528
(56), 4529 (43), 4530 (56), 4532 (23), 4534 (61),
4535 (14), 4535 (20), 4536 (41), 4537 (37), 4538
(37), 4539 (37), 4541 (37), 4542 (41), 4543 (41),
4544 (37), 4551 (14), 4552 (20), 4554 (20), 4556
(13), 4559 (6), 4560 (47), 4561 (6), 4562 (4), 4563
(43), 4564 (30), 4565 (30), 4567 (23), 4568 (23),
4569 (20), 4570 (43), 4573 (37), 4574 (34), 4575
(34), 4576 (20), 4577 (4), 4578 (4), 4579 (57), 4580
(4), 4582 (4), 4582 (4), 4583 (41), 4584 (41), 4585
(23), 4586 (20), 4589 (13), 4590 (13), 4591 (4), 4592
(43), 4593 (43), 4594 (13), 4595 (47), 4596 (37),
4597 (37), 4598 (20), 4599 (37), 4600 (23), 4601
(31), 4602 (57), 4605 (4), 4606 (4), 4623 (48), 4624
(57), 4625 (4), 4626 (48), 4627 (48), 4628 (48), 4629
(48), 4632 (48), 4633 (48), 4634 (48), 4635 (48),
4636 (48), 4637 (4), 4637 (48), 4638 (4), 4639 (57),
4640 (57), 4641 (4), 4642 (48), 4643 (57), 4644 (35),
4646 (41), 4649 (41), 4650 (16), 4651 (16), 4652
(16), 4653 (16), 4654 (23), 4655 (32), 4655 (48),
4655 (48), 4656 (32), 4658 (12), 4659 (51), 4660
(32), 4661 (32), 4662 (32), 4663 (48), 4664 (12),
4666 (51), 4667 (9), 4669 (31), 4670 (32), 4671 (32),
4672 (48), 4673 (48), 4674 (48), 4675 (48), 4676
(28), 4677 (4), 4678 (46), 4679 (31.note), 4680 (28),
4681 (26), 4682 (12), 4683 (31), 4684 (33), 4685
(33), 4686 (28), 4687 (11), 4688 (11), 4689 (46),
4690 (11), 4691 (40), 4692 (40), 4693 (14), 4694
(29), 4696 (47), 4697 (20), 4698 (6), 4909 (54), 4959
(13), 5107 (13), 5142 (13), 5174 (54), 5175 (29),
5176 (14), 5177 (14), 5178 (37), 5180 (61), 5181
(37), 5186 (4), 5206 (4), 5206 (4), 5343 (40), 5344
(40), 5458 (4), 5464 (46), 5497 (4), 5498 (4), 5511
(12), 5512 (33), 5513 (46), 5515 (4), 5516 (33), 5517
(11), 5518 (28), 5519 (28), 5520 (11), 5528 (4), 5529
(2), 5536 (4), 5536 (4), 5540 (57), 5541 (4), 5541 (4),
5542 (13), 5543 (13), 5544 (26), 5545 (26), 5546
(26), 5547 (4), 5549 (43), 5550 (6), 5551 (47), 5552
(47), 5553 (6), 5554 (43), 5555 (30), 5556 (30), 5557
(47), 5558 (37), 5559 (14), 5560 (23), 5561 (41),
5564 (4), 5565 (20), 5566 (41), 5567 (37), 5568 (4),
5569 (34), 5570 (4), 5571 (37), 5572 (34), 5573 (34),
5574 (18).
Fredholm, A., 3266 (41).
Freeman, J., 35 (41).
Freire, E. et al., 2784 (9), 3260 (51).
Freire, F.M.T., s.n. (49).
Freire-Fierro, A. et al., 1797 (49).
Freitas, ?, 129 (31).
Fr6es, R. de Lemos, 11634 (38), 19911 (39).
Fromm, E. et al., 471 (8).
Fuchs, H. P. et al., 21921 (37), 22112 (37).
Fuertes, M., 78 (SOb).
Furtado, C. X., s.n. (41).
Furtado, P. Pires, 154 (38).
Gabriel, A., s.n. (39).
Gabrielli, A. C. et al., 8751 (38), 8752 (38), 8758 (44),
8759 (44), 8759 (15).
LIST OF EXSICCATAE
Galarza,M. I. et al., 2 (8), 22 (51).
Galeano,G. et al., 512 (35), 513 (35), 552 (35), 562 (35),
642 (4), 645a (35), 645 (4), 675 (4), 1093 (25), 1093
(25), 2204 (9).
GaleanoS., W., 3 (8), 9 (9).
Galo, M., 3 (41).
Garber,P. A., 57 (41), 154 (41).
Garcia,H. et al., 60 (8).
Garcia-Barriga, H. et al., 4692 (35), 12286 (41), 14012
(51),14442 (12), 14722(12), 17053(22),19686 (41),
19686 (41).
Gardner,G., 1845 (38), 3981 (38 and 15 or 39), 3982
(38 and49).
Gamier,Bro. A., 1061B(41), 492 (41), 1712 (41).
Garwood,N. [et al.], 49 (37), 1130(37), 1144(37), 1146
(37), 1147 (37), 1162 (41), 2208A (41).
Gasche,J. et al., 222 (9).
Gaulin,?, 4 (57), 5 (57).
Gaumer,G. F. [et al.], 1459 (41), 24321 (41).
Gay,C., s.n. (44).
Gehrt,A., 45875 (38).
Gentle,P.H., 455 (41), 1566 (41), 1664 (41), 5490 (37).
Gentry,A. L. [et al.], 2618 (41), 3447 (19), 4895 (19),
6109 (19), 6149 (20), 7022 (47), 7451 (23), 8482
(12), 8958 (41), 9673 (26), 11791 (37), 12442 (11),
12759 (12), 13422 (19), 15501 (12), 15633 (12),
16504 (12), 17664 (41), 18194 (41), 18307 (25),
18558 (25), 18958 (51), 18998 (51), 20223 (37),
20289 (41), 20290 (41), 20514 (25), 20941 (25),
21300 (12), 21333 (12), 21817 (31), 23564 (55),
23820 (37), 24088 (20), 24195 (4), 24357 (37),
24357 (41), 24376 (61), 24521 (37), 25410 (12),
25762 (31), 25870 (25), 26527 (13), 26662 (29),
27376 (44), 27757 (51), 28103 (31), 28521 (37),
28801 (41), 28977 (12), 29516 (51), 29531 (25),
29982 (12), 30051 (20), 30086 (37), 31998 (51),
32123 (25), 32257 (37), 34941 (14), 35584 (37),
35641 (61), 35671 (20), 35680 (30), 35775 (44),
36153 (55), 37251 (9.note),37566 (33), 37566 (33),
37616 (44), 37616 (53), 37903 (46), 39501 (12),
39534 (12), 39840 (53), 39956 (1), 39956 (4), 40079
(44), 40290 (37), 41409 (53), 42136 (31), 42327 (31),
42783 (11), 43032 (31), 44666 (4), 44925 (33),
44934 (44), 44934 (44), 45194 (40), 45194 (40),
46069 (9), 47163 (12), 47757 (57), 50369 (50a),
50717 (SOb),51929 (38), 52282 (12), 53202 (56),
53288 (37), 53726 (37), 53945 (4), 55117 (13),
55425 (41), 55865 (51), 56424 (51), 57097 (31),
57488 (41), 57660 (51), 58001 (12), 58001 (9),
59325 (38), 60519 (6), 60723 (41), 60920 (12),
61217 (4), 61909 (31), 62936 (61), 63245 (4), 64399
(13), 64550 (14), 64565 (61), 64579 (4), 64746 (4),
64821 (31), 65484 (6), 65672 (12), 68643 (59),
68643 (25), 68813 (31), 69558 (12), 69785 (31),
69901 (13), 70264 (51), 72460 (47), 72522 (4),
72939 (47), 72939B (47), 72939A (23), 73169 (13),
75902 (47), 77156 (51), 80664 (9).
Gereau,R. E. et al., 2223 (37).
Gibbs,P.E. et al., 6638 (15).
203
Gibson,A. C. et al., 2450 (37).
Gillespie,L. J. [et al.], 803 (3), 995 (41), 1729(41), 2195
(25), 2344 (25).
GimateL., J., 642 (37).
Gines,H., 5156 (3).
Giordano,L. C. et al., 545 (15).
Giraldo-Gensini, J. et al., 411 (6).
Glassman,S. F., 1694 (41).
Glaziou,A.F.M.,6684 (38), 6684 (38), 7924 (15), 10060
(38), 11559 (15 and 38), 11599 (15), 12173 (15),
15426 (38), 17224 (38), 18497 (21), 18498 (21),
18499 (15), 18500 (15 and 38), 18501 (15 and 38),
18502 (38), 18503 (38), 18504 (38), 18505 (15),
18506 (15), 18507 (38), 18906 (38), 19863 (38),
22144 (38), 22144 (38), 27224 (38), s.n. (21, 38).
Gleason,H. A., 198 (3), 663 (51), 864 (25).
Glenboski,L. L., C.101 (12).
Godoi, J. V. et al., 1 (15), 2 (15), 24 (15), 25 (15), 56
(15), 174 (38), 269 (44), 270 (44), 330 (38).
Goeldi,A., 3957 (25), 3957 (12), 7726 (39), 7727 (36),
7728 (9), 7729 (51), 7730 (51), 7776 (39), 7777 (8),
7778 (36), 8232 (51), 19466 (39).
Gomes,J. C., 1 (38), 2 (38), 3 (21), 4 (15).
G6mez,C., s.n. (41).
G6mez,R. et al., 216 (32).
G6mez-Pompa, A. [et al.], 167 (37), 913 (37), 2568 (41),
3413 (37).
Gonzales,A. [et al.], 1012 (41), 1405 (41).
Gonzalez,G., 12 (44).
Gonzdles,H. et al., 42 (9).
Gonzdlez,M. S. [et al.], 115 (61), 1619 (47), 8200 (47).
Gonzdlez-Espinosa, M. et al., 1486 (41).
GonzdlezM., F. [et al.], 2419 (37), 5180 (37).
GonzalezQ., L., 575 (37).
Gooding,E.G.B.,331 (41).
Goodland,R., 3188 (38).
Gorts-vanRijn,A.R.A.et al., 317 (25).
Gottsberger,G. et al., 35-2575 (31), 127-21183 (25),
156-13375(38), 321-15183(36).
Goudot,J., 1 (16), s.n. (41).
Goulding,M., 7 (25), Ila (25), 1252 (25), 2117 (25).
Graham,?, 1395 (37).
Grande,D. A. de et al., 162 (38).
Grandez,C. et al., 3880 (11), 4091 (11), 4107 (18), 4156
(18).
Granville,J.-J. de [et al.], 2871 (25), 4397 (51), 6000
(17), 6611 (17), 7788 (36), 9319 (17), 9392 (17),
12995 (51), 13104 (9).
Gregory,L. E., 67 (50a).
Grenand,P., 1542 (36), 1690 (39), 2428 (8).
Grewal,M. S., 215 (3).
Griffis,J. D. et al., s.n. (SOb).
Grijalva,A. et al., 16 (41), 68 (51), 91 (12), 1355 (41).
Groll-Meyers,J. van, 105 (SOa).
Grubb,P.J. et al., 1627 (28), 1331 (2).
Guanchez,F., 786 (9), 787 (12).
Gudiiio,E. [etal.], 164(12), 179(18), 519 (46), 586 (46),
2021 (26), 2241 (51), 9720 (6).
Guedes,M. L., 1581 (38).
204
Guillaumet, J.-L. et al., 5850 (32), 5851 (32).
Guillen, R. et al., 1314 (39), 2686 (49), 3170 (25), 4430
(8).
Guti6rrez, G. et al., 17C675 (13), s.n. (13).
Gutierrez, V. et al., 542 (12), 769 (51).
Gutte, P. et al., 1566 (11), 1578 (12), 8161 (44).
Guzman, M. et al., 378 (41).
Haber, W. A.[et al.], 455 (4), 1658 (4), 6117 (37), 9970
(41), 10486 (37).
Hage, J. L. [et al.], 662 (15 and 38), 2255 (38).
Hagen, C. von et al., 1063 (41), 1264 (37).
Hahn, L., 210 (SOa), 342 (37), s.n. (50a).
Hahn, R., 6 (31).
Haines, ?, 467 (37).
Hall, E., 53 (41).
Hall6, F., 992 (36).
Hamilton, C. et al., 616 (19), 1108 (37).
Hansen, B. et al., 9158 (50a).
Harley, R. M. [et al.], 10838 (39), 10968 (38), 15223
(38), 18099 (38), 27357 (38), 50510 (38).
Harling, G. [et al.], 3809 (12), 21363 (59).
Harmon, W. E., 1828 (37), 4932 (41), 5198 (37).
Harriman, ?, 14663 (37).
Harris, C. G., 1612 (41).
Harris, S. A., Y.29 (51), TP.578 (3), TP.583 (3).
Hartshorn, G. S. [et al.], 2490 (38), 2694 (33).
Harvey, D. R., 5282 (37).
Hassler, E., 617a (38), 617b (38), 2278 (38), 7924 (38),
7924a (38), 12442 (38), 12442a (38).
Hatschbach, G. [et al.], 5103 (38), 7333 (38), 9920 (38),
19138 (38), 32454 (38), 34702 (38), 37457 (49),
41973 (38), 42244 (38), 43138 (38), 44846 (38),
44931 (38), 44969 (15), 45015 (38), 46710 (38),
48598 (21), 48643 (15), 48892 (15), 49743 (38),
52158 (15), 52159 (15), 52160 (38), 52174 (21),
53946 (49), 60929 (38).
Hayes, S., 671 (37), 863 (37).
Heinrich, ? et al., 7800 (37).
Helme, ? et al., 898 (9).
Helstone, M., 4 (25).
Henkel, T. W. et al., 2584 (25), 3519 (25), 4664 (25),
5318 (25), 5542 (41).
Heringer, E. P. [et al.], 10 (38), 491 (38), 493 (38), 509
(38), 513 (38), 515 (38), 516 (38), 552A (38), 997
(49), 1096 (49), 1685 (21), 1738 (38), 1826 (38),
2902 (38), 4495 (38), 5420 (38), 5746 (38), 7112
(38), 7311 (38), 7650 (38), 10171 (38), 10961 (49),
13024 (38), 15037 (38), 15054 (21), 15057 (21),
17350 (38), 17384 (38), 17630 (38), 17634 (38).
Hermnndez,R. et al., 488 (41).
HermnndezX., E. H., 606 (37), 635 (37).
Herrera, H. et al., 644 (41), 1763 (19).
Herzog, T., 1508 (8).
Hespenheide, H. A. et al., 874 (41).
Heyde, E. J. [et al.], 6238 (37).
Heyde, N. M., 710 (51).
Hill, M., 16 (37).
Hill, R. J., 1722 (8).
FLORANEOTROPICA
Hind,D.J.N.et al., 40 (38).
Hinojosa,I. et al., 1104 (44).
Hinton,G. B., 3568 (37).
Hioram,Bro., s.n. (SOa).
Hladik,A., 497 (41).
Hodge,W.A. [et al.], 632 (SOa),3168 (SOa),3663 (SOa),
4403 (50b).
Hoehne,F. C. et al., s.n. (38).
Hoehne,W. [et al.], 2004 (21), 2005 (21), 3325 (15),
12783 (38), 12787 (15), 13234 (38), 13237 (38),
17467 (38), 28268 (15).
Hoff, M., 6244 (36), 6866 (51).
Hohenkerk,L. S., l1SA (51).
Holdridge,L. R., 1061 (SOb).
Holm,R. W. et al., 234 (41), 950 (41).
Holm-Nielsen,L. et al., 20205 (31), 20211 (31), 20426
(12), 21754 (28), 24409 (4), 24410 (47), 24518 (4),
24705 (47), 24706 (29), 26201 (4), 26350 (4).
Holt, E. G. et al., 248 (25), 657 (25).
Holton,I. F., 251 (31), s.n. (31).
Hopkins,M.J.G.et al., 872 (25), 1422 (45).
Homer,C. et al., 73 (25), 360 (25).
Hostmann,F.W.R.et al., s.n (36).
Howard,R. A. et al., 18982 (50a).
Hoyos,J. et al., 822 (41).
Huashikat,V., 253 (12), 793 (12), 982 (12), 1189 (9),
1358 (51), 1828 (9), 1887 (51).
Huber,J., 494 (39), 3025 (39), 3026 (39), 3417 (39),
3869 (32), 4169 (31), 4175 (8), 4179 (25), 4184 (31),
4195 (25), 4237 (31), 4245 (12), 4474 (51), 4522
(31), 6877 (39), 7285 (8 and 39).
Huber,O., 6348 (41), 9235 (24).
Hunt,D. R., 5778 (38).
Hunteret al., ?, 462 (37).
Hunziker,J. H. et al., 11026 (38).
Hurtado,F. et al., 2403 (31.note),2457 (4), 2581 (4).
Hutchinson,P.C. et al., 3035 (4), 3009 (4), 3028 (4),
3039 (13 and57), 3844 (59).
IbarraM., G. et al., 1467 (37), 2002 (37), 2468 (37).
Ibarrola,T. S., 4060 (38).
Idrobo,J. M. [et al.], 604 (32), 662 (22), 792 (51), 1092
(4), 1093 (4), 1216 (12), 1949 (61), 4846 (12), 6298
(31), 6299 (31), 6300 (31), 6301 (31), 6310 (4), 6311
(4), 6312 (4), 6313 (4), 6314 (41), 6315 (41), 6316
(37), 6318 (31), 6319 (35), 6320 (35), 6321 (57),
6489 (32), 6518 (31), 6520 (12), 8086 (51), 8315
(31), 8316 (31), 8394 (12), 8753 (4), 9016 (12), 9975
(4), 10126 (37), 10680 (4), 10988 (16).
Irusta& FortoulLtd.s.n. (31).
Irvine,D. I., 103 (51), 114 (12), 210 (28), 211 (12), 728
(46).
Irwin,H. S. [et al.], 2199 (15), 5234 (38), 5234 (38),
8497 (38), 11799 (49), 14415 (38), 15807 (38),
16554 (38), 17688 (38), 18150 (38), 18307 (38),
21539 (38), 21539 (38), 24223 (49), 24910 (38),
24910 (38), 31509 (49), 32099 (49), 47159 (36),
47747 (36), 48703 (36), 55577 (51), 57681 (41).
LIST OF EXSICCATAE
Isern,J., 5PB (28).
Izawa,K., 25 (51).
Jack, J. G., 4355 (38), 4356 (SOb),4485 (SOb),5271
(SOb),5558 (50b), 7481 (SOb).
Jangoux,J. [etal.], 256 (38), 257 (38), 10146(25), 10147
(25), s.n. (25).
Jansen-Jacobs, M. J. et al., 971 (41), 1260 (41), 1542
(51), 2658 (41), 2997 (36), 4231 (25).
Jaramillo,J. [et al.], 3001 (51), 5763 (4), 5764 (2), 6714
(37), 7436 (20), 8145 (4), 10914 (12), 11475 (26),
12091(4), 12580(4), 12711(46), 13520(12), 13740
(14), 13742 (61), 14521 (6), 14651 (6), 14657 (47),
15100 (28), 16518 (25), 16525 (25), 16878 (12),
16961 (46), 17110 (18), 17182 (51), 31352 (31).
Jaramillo,N. et al., 517 (46), 579 (11), 851 (46), 994
(31).
JaramilloM., R. [et al.], 378 (32), 1017 (48), 1146 (32),
1146 (32), 1237 (51), 1261 (51), 2153 (31).
Jardim,J. G. et al., 396 (12), 871 (51), 1913 (15), 3067
(39).
Jativa,C. [et al.], 802 (37), 803 (14), 976 (26), 1099(37),
1151 (14), 2105 (14), 2160 (26), 2217 (37), 2223
(26), 2224 (26), 2233 (23).
Jenman,G. S., 4975 (36).
Jimenez,S. et al., 358 (10).
JimenezM., A., 2682 (37), 3184 (42), 3473 (41).
Johann,M. [et al.], 1/3-31188(51), JO-2/35(11), 2/32-
151188 (44), JO-2/26(11), 4/21-20888 (44), 4/5-
18888 (44).
Johnson,G. E. et al., 18C240(41), 22J181(41).
Johnson,W. et al., 127 (47).
Johnston,I. M., 78 (41), 154 (41), 155 (41), 471 (41),
1503 (27), 1758 (23).
Joly,A. B., 7349 (15).
Jones,G. C. et al., 3034, 3465 (41).
Jones,G. N. 11024 (SOa).
Jong,W. de, 163 (12).
JordyFilho,S., 53 (39).
J0rgensen,P. [et al.], 2158 (38), 3836 (38).
J0rgensen,P.M. et al., 65761 (57).
Joseph,?, 13100 (41).
Josse,C. et al., 785 (37).
Juncosa,A. [et al.], 695 (37), 1134 (37).
Juzepczuk,S., 4863 (41), 6411 (12), 6710 (41), 6928
(41).
Kalliola,R. et al., P5-4 (25), P5-9 (31), P5-13 (25), PS-
16 (25), P5-34 (25), P5-54 (31), P5-64 (31), P5-65
(25), P5-67 (31).
Kapelle,M. et al., MK.519(4).
Kauffmann,E., 603 (51).
Kayap,R., 213 (51), 368 (46), 442 (12), 680 (28), 805
(31), 832 (11), 1052 (28), 1099 (11).
Kellerman,W.A., 6039 (37), 6416 (41).
Kenoyer,L. A., 310 (41).
Kernan,C., 1064 (23).
Khan,R. et al., 1155 (41).
Killeen,T. [et al.],, 1083 (8), 3072 (4), 4407 (51).
205
Killip,E. P. [et al.], 588 (41), 5312 (31), 5352 (61), 7873
(47), 7874 (4), 8827 (56), 13571 (SOb),14243 (41),
14892 (41), 15010 (41), 16667 (57), 20852 (48),
23117 (55), 24047 (53), 24737 (4), 26499 (51),
26614 (12), 27053 (12), 28787 (12), 33221 (14),
33551 (61), 33712 (4), 38712 (37), 38713 (14).
King,R. M., 715 (41), 4426 (37).
Kirizawa,M., 2119 (38), 2120 (38).
Kirkbride,J. H. [et al.], 21 (19), 79 (41), 2381 (4).
Klein,R. M., 113 (15).
Klug,G., 399 (12).
Knab,C., 179 (41), 196 (51).
Knapp,S. [et al.], 5844 (19), 8482 (12).
Knees,S. G.[et al.], 2812 (41), 2836 (41).
Korning,J. et al., 47692 (11).
Kramer,K. U., et al., 2317 (41).
Krapovickas,A. [et al.], 23862 (38), 35177 (51), 35179
(12), 40699 (38), 40700 (38),42964 (38), 43065 (38).
Kress,W.J. et al., 94-422 (23).
Krieger,L. [et al.], 9054 (15), 10538 (21), 12284 (31),
12579 (12).
KroellS., B., 190 (31), 251 (53), 439 (59), 454 (59), 508
(59).
Krukoff,B. A., 1438 (25), 4517 (12), 4789 (51), 5307
(44), 5389 (51), 6237 (51), 6420 (12), 6718 (25),
7214 (9), 8061 (11), 8083 (51), 8333 (12), 8336 (11),
8407 (25), 8571 (31), 8921 (9), 10304 (4), 10305
(44), 10465 (44), 10811 (4).
Kuhlmann,J. G., 407 (11), 591 (12), 592 (51), 952 (8),
1047 (25), 1085 (39), 1139 (25), 1193 (25), 1205
(12), 1486 (31), 1582 (31), 1613 (31), 1617 (25),
1941 (58), 2328 (38), 3664 (21), 19881 (58), 19882
(51), 19884(31), 19886(25), 19889(25), 19890(12),
19891 (25), 19892 (25), 19893 (12), 19896 (31),
19903 (31), 20199 (39), 20200 (8), 111679 (21),
136990(21), 141261(21), 141262(21).
Kuhlmann,M. [et al.], 227 (15), 3326 (15), 202 (51),
12788 (38), 3988 (38), 2006 (38), 284 (36), 2003
(15), 3770 (15), 2674 (15), 2673 (15), 2567 (21).
Kujikat,A., 132 (11).
Kuntze,O., s.n. (37).
Kvist,L. P. et al., 48433 (37).
Ladeira,J. et al., 264 (15).
Laessoe,T. et al., H.52562(38).
Laguna,A., 399 (41), 400 (41).
Lallathin,B. R., 5065 (23).
Lamotte,S., 123 (9), 126 (12), 166 (25), 231 (25), 248
(9), 325 (11), 326 (11), 344 (9), 375 (26), 395 (12).
Lanjouw,J. [et al.], 453 (36), 720 (25), 824 (51), 922
(41), 2485 (51), s.n. (25).
LannaSobrinho,J. de Paula[et al.], 257 (15), 290 (8),
403, (25), 429 (31), 451 (12).
Lao,E. A. [et al.], 120 (27), 121 (27), 122(41), 169(37),
181 (20), 203 (41), 211 (23).
Lao M., R., 21 (57), s.n. (59).
La Rotta,C. et al., 543 (51), 723 (37).
LaSalle,? et al., 810704-8 (37).
Laughlin,R. M., 1282 (41).
206
Lauri,?, 187 (36).
Lawesson,J. E. et al., 39398 (12).
Lazor,R. L. [et al.], 1710a(37), 1710d(37), 2653 (37),
2824 (41), 2825 (41), 3066 (41), 3439 (41), 3440
(27), 3440 (27).
LBB (LandsBosbeheerSuriname),7616 (25), 8628 (36),
10621 (25), 12739 (41), 12742 (25), 12743 (25),
12744 (51), 12745 (51), 12746 (51), 12747 (51),
12750 (39), 12791 (51), 15274 (25).
Lechler,W., 2436 (4 and53).
LeDoux,D. G., et al., 2122 (37).
Leemans,J., 83 (38).
Leeuwenberg,A.J.M.,4504 (41).
Leguizamo,I., 493 (41), 494 (41).
Lehmann,F. C., 1969 (61), 2341 (35).
Leija,G. et al., 1112 (37), 2720 (37).
LeitaoFilho, H. F., 20345 (38).
Leite, J. E., 2300 (38).
Lemonnier,L. G. (herb.),s.n. (36).
Lent, R. W., 1031 (23), 1614 (4), 2052 (4), 2119 (23),
2120 (4), 2948 (37), 3154 (4), 4016 (37), 4017 (37),
4101 (4). Le6n,H. et al., 1492 (14), 1244 (37), 1245
(61).
Leonard,E. C. [et al.], 4875 (SOb),8532 (SOb),8655
(SOb),9990 (SOb),10026b(SOb),11496(SOb),11496
(SOa),13998 (SOb),14084 (SOb).
Leonard,J., 76 (38).
Leoni,L. [ et al.], 1218 (21), 1219 (21), 1699 (15), 2237
(21).
Leprieur,F.R.M.(herb.),195 (36), s.n. (36).
Letouzey,R., 12474 (41), 14828 (41).
Levy,P., 52 (41), 473 (37).
Lewis,W.M [et al.], 868 (37), 10346 (31), 11038 (31),
11206 (31), 12181 (59), 12601 (31), 13287 (12),
13413 (51).
Liebmann,F. M., 5889 (37), 5890 (37), 5891 (37), 14303
(37), 14306 (37).
Liesner,R. L. [etal.], 517 (19), 2066 (41), 3481 (9), 4076
(12), 5815 (41), 6573 (31), 6617 (12), 6965 (51),
8664 (51), 8825 (12), 9987 (4), 10437 (48), 10737
(41), 16194 (12), 16945 (12, 24), 16951 (12, 24).
Lima,E. et al., 64 (36).
Lima,L. et al., 529 (11).
Lindeman,J, C. et al., 3537 (38), 3617 (51), 5304a (38),
5364 (38).
Lindeman,J. C. & A.R.A. Goerts-vanRijn et al., 288
(25).
Lindeman,J. C. & E. Mennegaet al., 150 (41).
Lindeman,J. C. & A. L. Stofferset al., 367 (36), 400
(36), 516 (36), 549 (51).
Linden,J. J., s.n. (37).
Lindman,C.A.M.,A.3761 (38).
Liogier, A. M. (Bro. Alain) [et al.], 6432 (SOb),6433
(SOb),20542 (SOb),28274 (50a), 35988 (SOa).
Lira,C.M.S. [et al.], 137 (38), 409 (38).
Lisboa,P. et al., 950 (8), 1446 (25), 1897 (9), 2271 (51).
Lissot,J., 75/35 (51).
Little,E. L. [et al.], 6 (46), 19 (51), 35 (18), 61 (46), iSO
(2), 167 (12), 169 (46), 178 (46), 213 (2), 227 (4),
FLORA NEOTROPICA
234A (18), 234 (4), 248 (57), 347 (33), 386 (12), 430
(51), 572 (51), 651 (60), 654 (51), 669 (28), 1681
(33), 1764 (51), 6185 (37), 6220 (23), 6361 (37),
6386 (26), 6610 (37), 7343 (35), 7562 (35), 7595
(41), 7749 (51), 7750B (51), 7763 (4), 7765 (46),
7766 (46), 7771 (12), 8088 (57), 8088 (4), 8242 (32),
8337 (31), 8352 (51), 8478 (35), 8747 (57), 8780
(57), 8788 (2), 9498 (12), 9684 (26), 9710 (51), 9785
(51), 9787 (31), 13014 (50a), 13103 (50a), 15119
(41), 15120(41), 15437 (4), 15451(41), 15452(41),
15854 (41), 16215 (41), 16215 (41), 16775 (41),
16954 (51), 17668 (41), 17669 (41), 17708 (25),
21052 (37), 21082 (14), 21097 (14), 21175 (26),
21176 (26), 21224 (23), 25025 (41), 25073 (37),
25077 (37), 25201 (23), 25340 (23), 25356 (23),
25382 (23), 40030 (38), 40313 (41).
Lizot,J., s.n. (11).
LlanosH., F. et al., 1372 (35).
Lleras,E. [et al.], P.17033(12), P.17161(12), P.17161
(12), P.17553(8).
Locke,R. D. et al., 73 (38).
Loefgren,A. et al., 1657 (38).
Loiz, A. et al., 37 (50a).
Lojtnant,B. et al., 13297 (28), 14569 (28).
Lombardi,J. A. [et al.], 331 (38), 637 (38), 1757 (49),
2214 (49).
Lopes,J. M. et al., 2206.2257 (9).
Lopes,D., s.n. (45).
L6pez,F., 537 (41), 788 (37).
L6pez,G. et al., 262 (37), 351 (37).
Lorence,D. H. et al., 3436 (37).
Loroca,S., 25 (38).
Lott,E. J. et al., 1154 (37), 2213 (37).
Loubry,D., 1570 (36), 1570 (39), 1766 (51), 2351 (36).
Loureiro,A. et al., 37937 (58), 38041 (8), 48349 (8).
Lowe, J., 3999 (8).
Lozano,P. et al., 901 (4), 902 (26), 903 (26), 912 (26).
LozanoC., G. [et al.], 314 (12), 901 (4), 1078 (4), 2877
(41), 4900 (37), 4907 (20), 4975 (37), 4978 (56),
5082 (14), 5572 (14), 5652 (37), 6906 (54), 6940
(14), 6987 (54).
Lugo S., H., 118 (40), 1012 (28), 1065 (28), 1450 (28),
1461 (28), 1480 (28), 1685 (28), 1705 (28), 2003
(31), 2104 (31), 2614 (28), 2650 (28), 3302 (28),
3542 (31), 4825 (46), 4825 (46).
Lundell,C. L. [et al.], 2891 (37), 2902 (41), 3006 (41),
3016 (41), 3295 (41), 3296 (41), 4813 (41), 4814
(41), 6373 (37), 6374 (37), 7548 (41), 15972 (41),
16355 (37), 18191 (37).
Luteyn,J. L. et al. 4155 (41), 8637 (31), 12128 (47).
Maas,P.J.M.[et al.], 245 (8), 2411 (3), 2589 (3), 2594
(36), 3146 (38), 3525 (36), 3848 (36), 4551 (12),
4603 (25), 5105 (12), 5384 (24), 5571 (41), 5859
(51), 5981 (44), 5983 (11), 6035 (33), 6160 (8), 6161
(44), 6285 (25), 6414 (SOb)6420 (SOb),6934 (12),
6935 (9), 7339 (25), 7501 (36), P.12674(31), P.12771
(9) P.13281(31), P.13319(12).
Macbride,J. F., 5741 (4).
LIST OF EXSICCATAE
Macedo,A., 1503 (38).
Macia,M. J. et al., 325 (12), 2491 (51).
Maciel,U. N. et al., 2113 (31).
Mackenzie,C. A. et al., 2206.3034 (45).
Madriiain,S. et al., 406 (41), 407 (41), 458 (41), 458A
(41), 982 (12), 1000 (12).
Maestro,A. L. et al., 6 (38).
Maguire,B. et al., 34787 (12), 41691 (12), 54150 (25),
60161 (51).
Mahecha,G. [et al.], 281 (4), 1764(57), 3572 (51), 5588 Mills, A. P. 22 (44).
(32), 5589 (32), 5779 (51), 5824 (41), 6244 (31),
6389 (41), 7235 (51), 7853 (31), 8093 (51), 11162
(61), 11168 (41), 11673 (61), s.n. (4, 31, 37, 52, 57,
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Malme,G.O.A.,11.869(38), 11.1811(38), s.n. (38).
Manara, B., s.n. (4).
MarcanoBerti,L. [et al.], 89 (51), 641 (51), 2963 (41), Moore,et al., 3642 (37).
94-980 (32), 109-980 (41), 113-980 (41), 114-980 Mora0., L. E., 2310 (37), 2311 (14), 2310 (37), 3185
(41), 115-980(41), 144-981(3), 146-981(41), 189-
981 (41), 277-981 (48), 280-979 (4), 456-979 (41), Moraes,M.C.C.et al., 279 (49).
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Marquete,N., 1590 (15).
Marquez,W., 717 (37).
Mars,L. et al., 3304.2552 (9).
Marshall,N. [et al.], 86 (37), 188 (36), 394 (41).
Martin,B.-A. (herb),s.n. (36).
Martinelli,G. et al., 5011 (38), 6965 (25), 9908 (21).
MartinezC., R. et al., 10849 (38).
Martinez-Calderon, G., 3106 (37).
MartinezS., E. M., 7004 (37).
Martins,H. J. et al., 1301.523(45).
Martins,L.H.P.et al., 17 (45).
Martius,C.F.P.von, 630(1) (51), 630(2) (excl. name),
2834 (38), s.n. (25, 37).
Marulanda,0. et al., 1204 (9).
MatAsri, FRI25549 (41).
Mathes,L.A.F.,31C (38), 10084 (38).
Mathias,M. E. et al., 3446 (53).
Matthes,L., 1 (15).
MattosSilva, L. A. et al., 1155 (38), 2674 (21).
Matuda,E., 3240 (41), 16411 (37).
Maxon,W.R. [et al.], 5132 (37), 7193 (41).
Maxwell,I., s.n. (41).
Maxwell,J. F., 79-59 (44).
McDaniel,S. et al., 29832 (12).
McDonagh,J. F. et al., 79 (20), 405 (37).
McDowell,T., 2455 (25).
McVaugh,R., 717 (37), 11957 (37).
Meave,J. et al., B.265 (37).
MeijerDrees,E., 20 (41).
Mejia,M. et al., 6732 (SOb),6733 (SOb).
Melendez,E. J. et al., 175 (SOa).
M6linon,M., 18 (36), s.n. (39).
Mello, M.M.R.F.et al., 565 (38).
Mello-Silva,R. et al., 547 (21).
Mendonca,R. C., 235 (38).
MendoncaFilho,C. V., 28 (38).
Meneces,E. [et al.], 45 (11), 786 (51).
Merino,B. et al., 4514 (59), 4779 (2).
207
Merklen,E., s.n. (41).
Metcalf,R. et al., 30168 (37).
Mexia,Y, 1275 (37), 4349 (15), 4389 (21), 9225 (37).
Michel,R. de et al., 2437 (51).
Miege, E. et al., 3960 (41).
Miller,G. S. [et al.], 1251 (SOb),1350 (41), 6000 (SOa).
Miller,J. S. et al., 953 (41), 1604 (SOb).
Milliken,W. [et al.], 159 (41), 388 (25), 408 (41), 2101
(51).
Miranda,?, s.n. (39).
MolinaR., A. [et al.], 427 (41), 3455 (37), 13571 (37),
17928 (37), 18102 (37), 18220 (37).
MonsalveB., M., 627 (37), 1501 (61), 1843 (37).
Monteiro,L. et al., 122 (38), 123 (38).
Monteiro,0. P. [et al.], 1126 (51), 1278 (45).
(4), 4474 (4).
MoraesR., M. et al., 263 (31), 800 (8), 982 (31), 1239
(25), 2091 (8), 2286 (8).
Morais,H. C., 9201 (21), s.n. (49).
Morales,G. et al., 1116 (57).
Morales,M. E. et al., 647 (46), 648 (46).
Morawetz,W. et al., 11-11988 (9), 11-18988 (51), 15-
21988 (59), 11-211085(44), 18-131085(44).
Moreno,P. P., 1 (31), 1707 (41), 2796 (41), 3990 (41),
10791 (41), 15989 (41), 24261 (41).
Mori, S. [et al.], 4044 (37), 4989 (37), 5122 (19), 6998
(27), 6999 (41), 7511 (12), 11060 (38), 11357 (15),
11521 (38), 17671 (51), 18706 (36), 18749 (51),
18864 (36), 19817 (45), 19821 (51), 20907 (36),
21555 (51), 21959 (25).
Morillo,G. [et al.], 113 (32), 4193 (31), 5167 (9), 7548
(12), 9154 (51).
Morong,T., 717 (38).
Morton,C. V. [et al.], 2468 (37), 5419 (SOa),5571 (SOa).
Mosen,H., 1020 (38), 3214 (15), 3677 (15), 3926 (38).
Mosquera,E., 10855 (51).
Mostacedo,B. [et al.], 800 (9), 2261 (8), 2305 (44), 2391
(44), 2530 (44), 2610 (44), 2916 (8).
Mueller, ?, 2861 (55).
Muiioz,E. et al., 534 (37).
Murphy,H. et al., 531 (37).
Murray,N. J., 394 (41).
Mutis,J., 2887 (56).
Narvaez,L. H. et al., 280 (12).
Nascimento,J. R. et al., 66159 (52), 66244 (52), 66246
(52), 66261 (52), 66291 (52), 66294 (52),66295 (52),
66297 (52), 66334 (52), 66363 (52), 66364 (52),
66394 (52), 1201.1815(9), 1207.1589(9).
Nascimento,J.R.C.et al., 1103.371(59), 1105.404(52),
1302.1524 (52), 1302.3271 (52), 1302.3275 (52),
1302.3276 (52), 2206.3191 (52).
Nascimento,J.R.M. et al., 1109.374 (59), 1201.1319
(59), 1302.2957 (46), 1302.3270 (46), 1302.3273
(46), 1302.3312(46).
208
Nash, G. V., 889 (50a).
Navarette, H., 17 (31), 38 (31).
Nee, M. [et al.], 6643 (20), 6659 (41), 7533 (41), 7652
(41), 7949 (20), 9084 (37), 11004 (19), 22573 (37),
28120 (41), 28174 (41), 28268 (41), 28468 (41),
29824 (37), 35924 (44), 37513 (44), 37514 (44),
37515 (44), 39708 (44), 39842 (44), 40049 (8),
41051 (44), 41225 (49), 41226 (49), 42151 (8),
42752 (51), 42810 (8), 42895 (45), 43005 (8), 43272
(8), 44158 (SOb),44967 (44).
Negrelle, R. et al., A.743 (15).
Neill, D. A. [et al.], 3094 (41), 3392 (23), 5994 (4), 6003
(46), 6176 (28), 6234 (28), 6335 (51), 6336 (12),
6337 (28), 8252 (51), 9345 (8), 9345 (11), 9504 (12),
9703 (28), 9704 (33), 9742 (11), 9743 (18), 9743A
(18), 9744 (11), 10107 (31), 10130 (31).
Nelson, B., 671 (12).
Nelson, C. et al., 8737 (41).
Nelson, E. W., 337 (37).
Nevers, G. de [et al.], 3731 (19), 3752 (19), 3753 (41),
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Nicholson, D., 2037 (SOa).
Noberto, F. et al., 215 (38).
Noblick, L. R. et al., 2106 (38).
Norby, ?, 133 (37).
Noriega, R., s.n. (9).
Nuiiez I., 85 (44), 86 (44), 87 (33).
Nufiez, P. [et al.], 65 (40), 5461 (51), 5748 (51), 6044
(11), 6304 (44 and 53), 6876 (44), 6896 (25), 8230
(53), 8578 (44), 8616 (53), 10121 (44), 10727 (59),
11111 (55), 11276 (11), 11944 (4), 12793 (44),
14382 (11), 14627 (31).
Nuiiez, T. et al., 259 (26).
Oberwinkler, B. et al., 13409 (41), 15607 (41).
Oersted, A. S., 5861 (41).
Oldeman, R.A.A. [et al.], M.58 (SOa), 90 (31), T.737
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Oldenburger, F.H.F. et al., 1112 (52), 1297 (36), 1362
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Oliveira, E., 6139 (39).
Oliveira, A. A. de, 2771 (45).
Oliver, R. L. et al., 1884 (27), 1901 (41).
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O'Neill, H., 8819 (41).
Orcutt, C, R., 3030 (41).
Orozco, C. I. et al., 2181 (20).
Ortega, J. G., 4761 (37), 5086 (37), 5194 (37).
Ortega, R., 128 (SOa).
Ortega U., A., 148 (28), 205 (51), 361 (41).
Ortiz, L., 203 (47).
Ortiz, M., 805 (38).
Ortiz S., G. A. [et al.], 96 (37), 110 (41), 1979 (41).
Outer, R. W. den, 881 (41).
Pacheco, M. et al., 174 (8).
Paixao, J. L. et al., 74 (38).
Palacios, E., 66 (41).
Palacios, P. et al., 2655 (9)
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Palacios,W. [et al.], 1447 (46), 1750 (12), 1774 (18),
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(46), 1975 (51), 4460 (46), 4462 (51), 4465 (12),
4494 (18), 4684 (18), 5363 (4), 5673 (12), 5741 (46),
8178 (33), 8293 (28), 8884 (12), 9150 (26), 9150
(25), 9312 (9), 9627 (2), 9720 (6), 10036(29), 11164
(23).
Palmer,W. et al., 768 (SOb).
Pardo,M. de et al., 75 (25).
Paredes,R., 680 (19), 681 (19), 924 (19).
Pastore,U. et al., 84 (38).
Pedersen,T. M., 945 (38), 5923 (38), 13463 (38).
Pedralli,G. et al., s.n. (38).
Peixoto,A. Lunaet al., 359 (38).
Pennell,F.W.,3635 (16), 8042 (56).
Pennington,R. T. et al., 31 (11).
Pennington,T. D. et al., 9073 (37), 9273 (4), 14054(23),
14176 (23), 14180 (37), 14530 (37), 15230 (20),
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Pereira,B.A.S. et al., 3448 (38), 3514 (38), 3542 (38).
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Pereira,E. [et al.], 271 (38), 2838 (38), 3247 (39), 3715
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Peterson,P.M. et al., 6858 (37).
Peyton,B. et al., 374 (55).
Pfeifer,W., 1726 (41).
Pfeiffer,H. W. et al., 2619 (SOa).
Philipson,W.R. et al., 1623 (12), 1982 (4), 1983 (4),
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Phillips,0. et al., 172 (31), 181 (51), 554 (31).
Pic6n N., G. [et al.], 1108 (24), 1484 (41).
Pimentel,J. et al., 575 (50b).
Pinheiro,P.R., 175 (38).
PintoE., P. et al., 941 (12), 1165 (32), 6268 (31), 6290
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Pipoly,J. et al.,8217 (41), 11288(41), 11316(41), 13115
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Pires,J. Murqa[et al.], 910 (36), 1392 (36), 1393 (51),
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(38), 16911(25), 16931(25), 16963a-b(25), 16963a-
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Pittier,H. [et al.], 385 (41), 1985 (37), 2673 (37), 3823
LIST OF EXSICCATAE
(27), 3825 (27), 3826 (37), 3999 (37), 4056 (41),
4057 (37), 4060 (41), 6656 (37), 6666 (37), 6829
(41), 8626 (41), 9996 (37), 10266 (41), 10418 (41),
10518 (41), 11077 (37), 11077 (37), 11511 (41),
12143 (41), 14842 (41), 14842 (32), 14845 (32),
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Pl6e, A., 179 (50a).
Plowman,T. [et al.], 2272 (12), 5100 (31), 6433 (25),
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Poeppig,E., 1593 (44), s.n (44).
Pohl,J. E., s.n. (21 and 38).
Poiteau,P. A., s.n. (39).
Polak,M. et al., 333 (3).
Porter,D. M. [et al.], 4068 (19), 4194 (37).
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Prance,G. T. et al., 1700 (36), 1778 (51), 3460 (58),
4361 (39), 4665 (8), 5857 (51), 6574 (39), 7226 (8),
7226 (8), 8910 (12), 10152 (51), 10289 (25), 10473
(24), 11207(25), 12171(51), 12603(51), 15375(12),
15780 (9), 17827 (9), 17840 (9), 18008 (9), 18294
(12), 18294A (12), 20649 (58), 20925 (25), 20926
(25), 20927 (25), 20929 (25), 25548 (36), 25549 (36),
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Prescott,G. W. et al. 34 (51), 70 (51).
Prevost,M. F. [et al.], 292 (39), 706 (36), 708 (39), 731
(39), 732 (36), 733 (36), 801 (51), 811 (36), 812 (36),
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Proctor,G. R.[et al.], 8692 (41), 17949 (SOa),17949
(SOa),19153(SOa),19153(SOa),21135 (SOa),26152
(SOa),27245 (23), 32563 (41), 41467 (50a).
Pulle, A. A., 94 (25).
Purpus,C. A., 15387 (37).
Pursell,? et al., 8241 (41).
Quadros,J. et al., 1677 (38), 1678 (38), 1679 (15), 1680
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Quarfn,C. et al., 2446 (38).
Quarlesvan Ufford,L. H., 94 (37).
Quelal,C. [et al.], 65 (13), 273 (47).
Quevedo,R. et al., 2572 (39).
Quisumbing,E., 7933 (38).
Rabelo,B. V., 62 (8), 3112 (51).
Ramage,G. A., s.n. (SOa).
Ramalho,R. S. [et al.], 316 (38), 1337 (21), 1388 (15),
1825 (15), 3398 (38).
Rambo,B., 29298 (38), 53542 (38), 63578 (38).
Ramfrez,J. G. et al., 691 (23), 909 (37), 1212(37), 4151
(43), 4184 (43), 4242 (43), 4327 (37).
RamfrezP.,B. R. [et al.], 4167 (4), 6061 (29).
Ramos,J. E. et al., 3071 (31).
Rangel,0. et al., 13234 (4).
Ratter,J. A. et al., 202 (38), 5081 (38), 5503 (39), 6300V
(25).
Raunkiaer,C., 1450 (SOb),s.n. (SOb).
Raven,P. H., 21644 (37).
209
Reark,J. B., 5687 (37).
Reekmans,M., 9622 (38).
Regnell,A. F., 194 (15), 1.415(38), 111.1100(21).
Reinders,M. A., 36 (51).
Reitz,R. [et al.], 177 (38), 319 (15), 521 (38), 853 (38),
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Renner,S., 52 (8).
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Renterfa,E. et al., 1688 (41), 1913 (41), 1929(31), 4713
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Restrepo,C. [et al.], 378 (6), 440 (13), 441 (6).
Revilla,J. [et al.], 33 (25), 3577 (12), 6976 (58), 8642
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Reyes, G. et al. 1266 (41).
ReynaR., N., 20 (51), 65 (51).
ReynelR., C. [et al.],480 (44), 480 (44), 4016 (44), 4417
(44), 4692 (53), 4706 (44), 5131 (51), 5144 (12),
5396 (44).
Ribeiro,J.E.L.S.[et al.], 822 (45), 1293 (8), 1424A (9),
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Ribeiro,R., 21 (21).
Richardson,W.D., 683 (41).
Ricksecker,A. E., 449 (SOb).
Ridley,H. N. et al., s.n. (38).
Ridley,?, s.n. (SOa).
Riedel,L. [et al.], 74 (15, 78).
Riera,B. J. J.-L. [et al.], 649 (36), 650 (39), 905 (51),
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RimachiY., M., 593 (12), 8352 (25).
Rios, M. et al., 154 (20).
Rios T., J., 100 (4), 1801 (44).
Rivera,G., 1933 (23), 1934 (4).
Rivero,E., 135 (11).
Rizzini,C. M., 205206 (49).
Rizzo, A. et al., 4642 (49).
Roberts,L., 12791 (51).
Robinson,J.W.L.,364 (31).
Robles,R., 1446 (37).
Robleto,W., 401 (41).
Rocha,D.M.S., 10370 (38).
Rodrigues,R. S., 7841 (39), 7842 (39).
Rodrigues,W. [et al.], 403 (12), 1793 (51), 2090 (45),
2129 (9), 5009 (8), 5282 (12), 8421 (8), 8439 (51),
8891 (12), 8892 (31), 8893 (8), 8906 (8), 8907 (45),
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Rodriguez,?, 240 (38).
Rodriguezdel A., L. T., 23 (51).
Rolddn,F. J. et al., 2161 (57).
Romaniuc,S. [et al.], 760 (15), 778 (38), 787 (15), 791
(38), 1015 (15), 1116 (15), 1199 (38), 1200 (38),
1207 (38), 1218 (38), 1237 (38), 1272 (38), 1273
(15), 1274 (38), 1275 (15), 1276 (15), 1277 (38),
1278 (38), 1279 (15), 1280 (38), 1281 (38), 1282
(38), 1283 (38), 1300 (21), 1364 (38), 1366 (21),
1374 (38), 1377 (15), 1396 (15).
Rombouts,H. E., 64 (25).
Romero-Castafieda, R., 1137 (27), 3324 (14), 4847 (41),
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210 FLORANEOTROPICA

Rooden,J. van [et al.], 251 (4), 257 (37), 341 (31), 344
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Roque,N. et al., 92 (38).
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RosasR., M., 317 (37).
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Rovirosa,J. N., 383 (41).
Rubio,D. et al., 978 (47), 1263 (14), 1535 (14), 1946
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Rudas,A. et al., 3336 (51), 3352 (51), 3353 (12), 3489
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Ruiz, H. & J. Pavon,s.n. (1, 4, 44, 53).
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RufzZ., T. et al., 2806 (48).
Rusby,H. H. [et al.], 99 (41), 273 (25), 620 (44).
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Rzedowski,J., 10595 (37), 25511 (37).
Sabino,B., s.n. (37).
Sagot,P., 515 (36), 861 (51), s.n. (51).
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Saint-Hilaire,A.F.C.P.,89 (15), 90 (38), 91 (38).
Saldias,M. [et al.], 1038 (31), 1236 (8), 1429 (8), 1440
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Salino,A. 3574 (38).
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SanchezS., D. [et al.], 56 (41), 74 (41), 198 (37), 990
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Santos,J. U. et al., 659 (39).
Santos,M. R., 352 (51).
Santos,R. R. de et al., 1503 (38).
Santos,T. S., 1950 (21), 2063 (38).
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Sastre,C. [et al.], 2404 (12), 3110 (12), 3116 (12), 6909
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Schinini,A. [et al.], 10866(38), 17724(38), 20584 (38),
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Schreber,J.C.D.von (herb.),s.n; (SOa).
Schultes,R. E. [et al.], 684 (50b), 2003 (12), 3275 (57),
3697 (12), 3778 (31), 3821 (11), 3964 (51), 3996
(25), 5128 (35), 5172 (57), 5205 (57), 6583A (25),
6720 (25), 6781 (51), 6912 (25), 7156 (25), 7180A
(25), 8446 (51), 13028(12), 13679(51), 13904(12),
13990 (51), 14240 (22), 14370 (22), 14555 (12),
15876 (12), 16343 (12), 16811 (25), 17191 (9),
18500 (22), 22508 (4), 24021 (25), 24272 (22).
Schulz,J. P. [et al.], 7616 (25), 8628 (36), 10109 (51),
10621 (25).
Schulze-Rhonhof, H., 2996 (40), 3068 (60), 3072 (18).
Schunke,V., J., 1417 (12), 2313 (12), 7694 (44), 10579
(9.note),10593 (59), 10595 (33).
Schupp,E., 2 (38), 7 (38).
Schwacke,W., 11128 (21), 11129 (38).
Schwarz,G. J., 11407 (38).
Schwindt,E., 2983 (38).
Seemann,B., 150 (37), 499 (37).
SEF (Studieson EcuadorianForest),8566 (51), 8604
(51), 8644 (51), 8849 (11), 9227 (51), 10099 (31),
10108 (28), 10183 (51), 10223 (46), 10245 (46),
10249 (28), 10366 (51).
Segadas-Vianna, F. [et al.], 838 (38), 3729 (38), 3730
(38).
Seibert,R. J., 565 (41).
Seidel, R. [et al.], 994 (10), 6224 (44), 6628 (25), 6674
(25), 7316 (11), 7394 (44), 7395 (8).
Sellow,F., s.n. 2157/2146(38).
Semir,J. et al., 5026 (38).
Sermefno, A., 80 (37).
SerraltaP., 34 (41).
Servicedes Eauxet Forets,4070 (36).
Seymour,F. C., 6101 (41).
Shafer,J. A., 325 (SOb),342 (SOa),511 (SOb),1726
(SOb),2643 (SOb).
Shattuck,O., 882 (41).
Shepherd,D., 557 (41), 596 (20), 606 (23), 609 (37), 611
(37), 614 (41).
ShilomTon,A., 3075 (41), 3631 (37), 3894 (37).
Sidekbin Kiah,S.60 (44).
Sieber,F. W., s.n. (8, 39).
Silva,A.S.L. da et al., 450 (12).
Silva, I. A., 190 (38).
Silva,J. M. da et al., 3304.596 (51).
Silva,M., 992 (25).
Silva,M. F. et al., 173 (12), 501 (12), 1381 (12).
Silva,M.F.F.da et al., 15 (8), 16 (8), 67 (36), 388 (25).
Silva,M. G. [et al.], 4441 (49), 5724 (39).
Silva, N. T., 956 (52), 1178 (36), 1226 (51), 1340 (36),
3284 (52).
LISTOFEXSICCATAE
Silveira,M. et al., 712 (12), 815 (12), 1310 (25), 1311
(25), 1327 (31), 1329 (31).
Sim6es,J. M. et al., 73 (15).
Simonis,J. E. et al., 134 (38).
Sintenis,P., 250b (50b), 3971 (50b),4805 (50b).
Skutch,A. F., 3986 (37).
Smith,A. C., 2164 (3), 3426 (41).
Smith,C. L., 57 (37), 1040 (41).
Smith,D. N. et al., 465 (2), 1832 (55), 1875 (51), 2422
(59), 2427 (12), 3123 (44), 3663 (55), 3827 (51),
4179 (55), 4196 (55), 4525 (4), 4686 (7), 5225 (33),
5340 (55), 6905 (4), 7608 (55), 7848 (55), 7974 (55),
8555 (55), 12856 (8), 12856 (4), 12897 (31), 13218
(51), 13264(5), 13564(11), 13643(51), 13713(11),
13772 (44), 13989 (53), 14013 (53), 14126 (11),
14336 (8), 14349 (31).
Smith,F. D., 141 (41).
Smith,H. H. et al. 1407 (SOa),2105 (41), 13100 (41).
Smith,J. Donnell,1499 (37), 1675(41), 2024 (37), 4934
(37), 4934 (37), 5494 (41), 6238 (37), 6771 (37),
6772 (41), 6773 (23), 6774 (37), 7164 (41), 7165
(37), 7411 (4), 7666 (37), 7890 (37).
Smith,L. B. et al., 5897 (38).
Smith,L. C., 681 (37).
Smith,N., B.38 (36).
Smith,R. F., V.526(48), V.910(57).
Smith,S. F. et al. 785 (44), 1608 (44).
Sneidem,K. von, A.61 (23), A.71 (37), A.72 (31), A.73
(37), A.74 (37), A.245 (23), A.245 (37), A.246 (41),
A.485 (54), A.563 (13), A.1053 (31), A.1352 (12),
2218 (4), 3070 (47?), 5612 (4).
Snethlage,E. H., 283 (36), 338 (8), 383 (8), 384 (8), 385
(8), 386 (8), 388 (8), 647 (38).
Soares,A. M. et al., 2303.255 (58).
Sodiro,A., s.n. (13).
Soejarto,D. D., et al., 112 (35), 2661 (23), 2662 (41).
Solomon,J. C. [et al.], 3437 (12), 6165 (12), 7982 (51),
12459 (10), 12471 (44), 13999 (44), 14690 (8),
18965 (44).
Soria,N., 2358 (38).
Soto, N. et al., 5064 (37).
Soukup,J., 3017 (12), 3018 (51).
Sousa,M. [et al.], 101 (37), 1180 (37), 2361 (37), 11238
(41).
Souza,S. A. da M. et al., 140 (36).
Souza,W. S. et al., s.n. (15).
Sparre,B., 14534 (26), 15512 (26), 15514 (37), 17358
(47), 18113 (14), 18866 (26).
Spellman,D. et al., 1938 (41).
Sperling,C. R. et al., 5966 (36).
Sperry,O., 1011 (37).
Spichiger,R. [et al.], 1751 (12), 1752 (12), 3049 (12),
3050 (9), 3051 (9), 3053 (9).
Splitgerber,F. L., s.n. (36).
Spruce,R., 1322 (8), s.n. (8).
Sposito,T. C., 82 (15), 83 (21).
Stahel,G., 179 (51).
Standley P.C., 11788 (41), 19614 (37), 22045 (41),
22655 (37), 22665 (37), 23181 (37), 24143 (41),
211
26809 (41), 28477 (41), 54528 (41), 56637 (37),
71248 (4), 90576 (4), 90762 (4).
Stannard,B. L. [et al.], 4 (12), 7271 (38).
Starry,D. E., 308 (37).
Steam,W.T., 977 (41).
Steege,H. ter,290 (3).
Steere,W.C., 1700 (41).
Stehle,H., 1382 (50a), s.n. (50a).
Stehmann,J. R. et al., 1129 (38).
Steinbach,J., 3555 (44), 7267 (44), 7268 (8), 7268bis
(8), 9030 (4), s.n. (4).
Stergios,B. [et al.], 3710 (41), 5372 (41), 9081 (12),
9401 (25), 10800 (41), 10800 (25), 11330 (12),
11784 (51), 12888 (9), 12888 (3).
Stem,W.L. et al., 72 (41), 490 (37).
Stevens,W.D. [et al.], 2703 (41), 2856 (41), 2924 (41),
3341 (41), 3641 (41), 4145 (41), 4158 (41), 6359
(37), 6454 (37), 6925 (37), 7793 (37), 8813 (23),
9556 (41), 9557 (41), 11688 (4), 12193 (37), 17212
(41), 17423(41), 17599 (41), 19926(41), 22520 (4),
24592 (37).
Stevenson,J. A., 2772 (SOb).
Stevenson,P., 14 (37), 226 (51), 294 (31), 361 (32), 545
(32), 779 (9), 889 (12).
Stewart,A., 291 (42).
Steyermark,J. A. [et al.], 758 (24), 48888 (4), 55817a
(57), 56101 (4), 59016 (24), 60416 (24), 60540 (51),
62754 (24), 75387 (51), 86723 (41), 87270 (51),
87693 (41), 87693 (25), 87808 (25), 89976 (41),
90526 (41), 90526 (3), 90526 (9), 90614 (51), 91246
(41), 91272 (41), 91644 (4), 91899 (4), 92863 (24),
93643 (24), 94778 (4), 94971 (4), 97107 (57),
101275(57), 102717(12), 102718(51), 104065(24),
105784(57), 108727(32), 109842(24), 109843(24),
113020(41), 114804(25), 118157(4), 118640(57),
119050(48), 119082(41), 120308(41), 121434(4),
122299(12), 123221(41), 129060(12), 103564(57).
Stier,F., 67 (41).
Stimson,W.R. [et al.], 1337 (50a), 1901 (SOa),4057
(50a), 5143 (41), 5143 (41), 5182 (41), 5230 (41),
5251 (37), 5306 (41), 5378 (20).
Stoffers,A. L. [et al.], 159 (3), 317 (51), 339 (36), 457
(32), 516 (32), 4230 (SOa).
Stork,H. E., 4621 (37).
Strang,H. E., 343 (38), 601 (38), s.n. (39).
Strelnikow,I., 13 (38), 76 (38), 86 (38).
Strube,L. B., s.n. (SOa).
Strudwick,J. J. et al., 3187 (25).
Sturrock,?, 714 (50a).
Sucre,D. [et al.], 795 (38), 1968 (15), 3641 (38), 4923
(38), 5834 (38).
Sugden,A., 1182 (41), 1204 (41).
Sutton,? et al. 152 (41).
Swartz,O., s.n. (41).
Sylvestre,L. et al., 821 (38).
Tamashiro,J. Y. et al., 688 (15), 1255 (15), 18821 (38),
18844 (38).
Tamayo,F.T., 2981 (41).
212
Tameirao Neto, E. [et al.], 747 (38), 898 (15), 899 (38).
Tate, G.H.H., 250 (24), 394 (37).
Taylor, C. M. et al., 8823 (50a).
Taylor, D., 125 (50a).
Taylor, J. [et al.], 12628 (41), 17540 (37).
Taylor, K., 394 (37).
Taylor, N., 235 (50b).
Taylor, R. J., 4393 (4).
Teague, G. W., 397 (38).
Teixeira, L.O.A. et al., 672 (51).
Telles, ?, 49542 (15).
Tellez, 0. et al., 2189 (41), 2510 (41), 3352 (41), 3425
(41).
T6llez V., et al. 11733 (37).
Tello, 242 (59), 506 (51), 1117 (53), 1118 (53), 1583
(11), 1718 (53), 1731 (11), 1752 (53), 1771 (11),
1833 (11), 1837 (51), 1838 (51), 1839 (51), 1840
(51), 1841 (51), 1842 (51), 1872 (53), 2121 (51),
2639 (59).
Templeton, B. C., 8913 (37).
Tenorio L., P. et al., 3450 (37).
Terborgh, J., 6570 (51).
Terceros, W. et al., 368 (8).
Tessmann, G., 3253 (25), 3307 (25), 3312 (11), 3367
(25), 3454 (31), 3455 (11), 3466 (25), 3713 (31),
3713 (11), 3718 (51), 3981 (28), 4033 (31), 4033
(46), 4058 (11), 4278 (51), 4717 (51), 5044 (11),
5444 (25), s.n. (31).
Teunissen, P. A., 12739 (41), 12742 (25), 12743 (25),
12744 (51), 12745 (51), 12746 (51), 12747 (51),
12750 (39), 15274 (25).
Thieme, C., 194 (41).
Thomas, W. W. [et al.], 3351 (12), 5796 (49), 8903 (39),
11347 (38), 11348 (38), 11774 (15).
Thompson, J. B., 390 (38.vs).
Thompson, S. A. et al., 1398 (41).
Thomsen, K., 541 (23), 864 (37), 873 (23).
Tillett, S. S. et al., 45398 (3), 45804 (3).
Timana, M. [et al.], 913 (4), 918 (4), 1406 (25), 1508
(25), 1539 (44), 1595 (44).
Tipaz, G. [et al.], 831 (26), 1258 (14), 1460 (37), 1479
(37), 1480 (47), 2511 (37).
Tirado, M. [et al.], 695 (37), 735 (61), 1104 (14), 1109
(61), 1414 (37), 1443 (23).
Tiritan, 0. et al., 133 (38).
Toledo, C. B., s.n. (15).
Toledo, J., 2 (51).
Tonduz, A., 3999 (37), 10055 (41), 12642 (4), 13870
(41), 13870 (41).
Toro, R. A., 586 (41).
Torres, J. H. et al., 3021 (12), 3093 (31).
Torres, R. et al., 10423 (37).
Tostain, O., 263 (9), 270 (9).
Tovar, O., 4798 (53).
Townsend, W. R., 6 (8).
Tredwell, R. [et al.], 16c (51), 1887 (31).
Tresling, J., 1 (25).
Triana, J. J., 863 (37), 864 (37), 865 (20 and 37), 1865
(37 and 61), 1865 (41).
FLORA NEOTROPICA
Trujillo, B. et al., 15010 (12), 17307 (25), 18553 (57),
20405 (12).
Tsugaru, S. et al., B.635 (8).
Tuerckheim, H. von, 7666 (37), 7890 (37).
Tin Ortiz, R., 385 (41), 1899 (41).
Tunqui, S., 16 (28), 350 (28).
Tweedie, F., s.n. (38).
Tye, H., C.92 (25).
Tyson, E. L.[et al.], 929 (41), 1069 (41), 2927 (23), 3524
(41), 3526 (37), 4773 (41), 4852 (37), 4853 (37),
4854 (27), 4854 (27), 6077 (37), 6170 (37), 6205
(41), 6932 (37), 6950 (37), 6951 (20).
Uhl, C. F., 19 (12), 51 (9), 101 (12), 102 (12), 104 (12),
105 (12), 107 (12), 108 (12), 109 (12), 110 (12), 520
(31).
Ule, E., 1106 (38), 1196 (15),4863 (21),5512 (51),5587
(9), 5588 (12), 5938 (25), 5986 (8), 6845 (33), 7174
(49), 7174 (49), 7891 (41), 8837 (39), 8838 (58),
9311 (44), 9311 (12), 9312 (44), 9313 (11).
Uribe-Uribe, L. et al., 155 (41).
Usteri, P., 393b (38).
Vageler, H., 69 (31).
Valdivia, P. E., 800 (37).
Valencia, R. et al., 375 (9), 2275 (2), 2285 (2), 2304 (2),
2306 (2), 2313 (2), 2336 (2), 2344 (2), 2349 (2), 2386
(2), 2447 (2), 2449 (2), 2478 (2), 2512 (2), 2548 (2),
2564 (2), 2586 (2), 2612 (2), 2631 (2), 2910 (2), 2912
(2), 2913 (2), 2925 (2), 2930 (2), 67779 (51), 67780
(51), 67967 (31), 67967 (9).
Valerio R., J., 3593 (41).
Valeur, E. J., 480 (SOb).
Vareschi, V., s.n. (41).
Vargas, H. et al., 355 (4).
Vargas, I. G. et al., 2695 (9), 3891 (9).
Vargas C., C., 15490 (55), 15760 (51), 16381 (4), 16517
(4), 16900 (8), 18459 (53), 18486 (31), 18572 (12),
21274 (44).
Vasquez, R. et al., 676 (12), 776 (26), 776 (11), 776 (25),
1856 (11), 1995 (51), 2568 (12), 3021 (12), 3276
(51), 4087 (12), 4136 (31), 4591 (12), 6074 (51),
9437 (12), 11631 (25), 12120 (12), 12447 (31),
12449 (25), 12453 (25), 12664 (31), 12685 (11),
14725 (9), 15021 (9), 18822 (28), 19527 (51), 19532
(46), 19807 (51), 19938 (9), 19973 (9).
Vavrek, M. et al., 451 (38).
Vazquez, A. A., 176 (37).
Vazquez, B., 5 (37).
Vazquez Avila, M., 194 (38), 354 (10), 425 (44).
Vazquez Y, C., 870 (37).
Velasquez, J., s.n. (37).
Velasquez, J. [et al.], 78 (41), 79 (41), 80 (41), 81 (41),
82 (32), 83 (32), 84 (32), 86 (41), 87 (41), 88 (41),
89 (32), 90 (32), 91 (32), 92 (4), 93 (4), 94 (41), 95
(41), 97 (41), 98 (41), 101 (48), 102 (41), 244 (4),
245 (41), 246 (41), 247 (41), 249 (4), 250 (4), 252
(3), 252 (41), 253 (3), 254 (3), 255 (51), 256 (41),
257 (41), 258 (51), 259 (3), 260 (3), 261 (48), 262
LIST OF EXSICCATAE 213

(48), 263 (41), 299 (37),5O1(11),506 (11), 614 (51), Wilbert,W., 68325 (3).
2093 (4), 2094 (41). Wiley,J. R., 572 (41).
Velasquez,Ma. P. et al., 238 (41), 293 (23). Williams,D. E. et al., 653 (8), 987 (44).
Velez, M. C. et al., 724 (35), 1693 (4), 2926 (56), 2997 Williams,LI., 96 (25), 2068 (12), 2130 (51), 2373 (53),
(57), 3370 (4). 2557 (12), 4173 (11), 4313 (31), 7372 (12), 8411
Ventura,E. et al., 729 (37), 2705 (37). (37), 8665 (37), 9264 (41), 9431 (37), 9466 (37),
VeraSantos,J., 2891 (37). 10074 (41), 11469 (41), 11533 (41), 11665 (41),
Versteeg,G. M., 418 (25). 11666 (41), 11821 (41), 12689 (25), 12893 (41),
Vester,H. et al., 501 (9), 725 (51), 726 (9), 727 (12). 13300 (41), 13300 (41), 13300 (25), 13983 (12),
Vianna,M. C. et al., 341 (38), 342 (38). 14733 (25 and 31), 14760 (12), 14842 (32), 14878
Vicentini,A. et al., 574 (45). (32), 15516 (11), 15516 (25), 15765 (25).
Vickers,W.T., 91 (9), 92 (18). Williams,L. 0. [et al.], 13438 (41), 16034 (37), 23663
Vieira,C. M. et al., 81 (15), 554 (44). (4).
Villanueva,?, 321 (41). Williams,R. S., 339 (37), 400 (51), 645 (8).
Vinha,S. G. da, 27 (38). Wilson,P., 229 (37).
VivarC., F. et al., 2844 (26). Wood,C. W., 429 (4).
Vogl,C., 787 (41). Woodbury,R. et al., s.n. (SOa).
Voorhies,B. et al., 16-23 (41). Woodson,R. E. et al., 1312 (27), 1313 (37).
"V.V.,"5139 (48). Woodworth,R. H. et al., 644 (37).
Woolston,A. L., 835 (38).
Wachenheim,H., 381 (36). Woronow,S. et al., 4556 (41), 6302 (25).
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Warming,E., 1934 (21), 1935 (21 and38), s.n. (41). Yanez,A. P. et al., 300 (20), 1473 (37).
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Werner,?, 24/80 (51). Zappi,D. C. et al., 10335 (38).
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(25), 2102 (51), 2312 (9), 2313 (12), 2314 (12). 6499 (38), 7162 (38), 7812 (38), 8106 (38), 8311
West,D., s.n. (50b). (38), 8422 (38), 8510 (38), 8758 (38), 9326 (38),
Wester,P.J., 48 (38). 9392 (38), 9982 (38), 10047(38), 10415(38), 15037
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Widgren,J. F., 810 (15). Zuloaga,F. 0. et al., 1421(44),1497 (8),1552 (44),1833
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Wijninga,J., 525 (32), 541 (56). (23), 800 (31).
 CECROPIA AND ITS BIOTIC DEFENSES
D. W. DAVIDSON
In all the world, the genus Cecropia is unrivaled ative to the internodalgroove is thought to be the
for the number of myrmecophytes, or true "ant- product of naturalselection imposed by mutualistic
plants"counted among its species (McKey & David- ants (Schimper,1888). Both the internodalgrooveand
son, 1993). Based on the proportionof Cecropiaspe- prostomaare devoid of fibrovascularbundles,collen-
cies producingMullerianbodies in at least some parts chyma, and lignified parenchyma(Schimper, 1888).
of their distribution, myrmecophytes comprise the Colonizing queens of obligate Cecropia-ants recog-
vast majority(ca. 80%) of species in the genus; most nize and use prostomata as easily excavated sites
nonmyrmecophytesoccur at higherelevationsand on where stems can be entered without rupturingstem
islands, where theirants are missing (Wheeler,1942). transporttissues and flooding internodeswith muci-
Geographically, myrmecophytic Cecropia occur lage.
throughoutthe latitudinalrange of the genus, from The stems of myrmecophyticCecropiaarehollow,
southern Mexico to northernArgentina. Given that lacking the pith that often fills stems of nonmyrme-
association with ants is so widespreadtaxonomically cophytic congeners. The thin, spongy medullarylin-
and geographically,it is likely that relationshipswith ing the hollow stems of myrmecophytesis exploited
ants have been highly influentialin the evolutionary by queens that found their colonies claustrally,i.e.,
diversificationof Cecropia.Here,I reviewthe benefits without foraging, and by using resources from di-
which Cecropia-ants and their hosts receive from their gested wing muscles to produce their first worker
symbiotic partnerships.I then discuss how interspe- broods. Claustrallyfounding queens scrape this ma-
of Cecropiamight terial from the internodewalls to seal the prostoma
cific variationin the ant-attractants
have evolved and then influenced relationshipswith and develop their colonies in seclusion. Foundresses
particularant taxa. Finally, I consider factors influ- of some Cecropia-ants also accumulate and store
encing the diversity of ants, plants, and partnerships medullarytissue near their developingbroods.Water
in these associations. containedwithin this succulentmaterial(and lending
BENEFITS TO CECROPIA-ANTS it the appearanceof "bubblepack"under a stereo-
scope), may nourishdevelopingbroodand/orenhance
Myrmecophytesare plants with "bioticdefenses,"
humidityin the vicinity of brood.
or traitswhich attractants,mainlyas a defense against
In addition,as describedabove by Berg, Cecropia
herbivores and encroaching vines (Davidson &
have evolved to supply two types of food rewardsfor
McKey, 1993). By definition, myrmecophyteshave
theirants,and at least some antassociatesof Cecropia
evolved not only to provision ants with food re-
appear to reject foods typically taken by ants with
sources, but also to grantthem residenceinside either
more generalizedhabits. Mullerianbodies, produced
modified plant parts or preadaptedstructures.Evolu-
at hairy trichilia, located abaxially on the bases of
tion by plants to attractand house ant colonies can
petioles, are harvested by obligate Cecropia-ants, but
often be in doubt in presumedmyrmecophyteswith
not by opportunistically foraging ants with general-
primarydomatia,i.e., naturallyhollow or hollowed-
ized diets (Davidson & Fisher, 1991). These food
out stems that are frequentlyor always occupied by
rewards contain considerable lipid and at least some
ants, because many of these species have no obvious
specializationsfor housing or feeding associatedcol- protein (Rickson, 1973, 1976) but are approximately
onies (Davidson & McKey, 1993). Such is not the 30% glycogen (Rickson, 1971; Marshall& Rickson,
case in Cecropia species, which accommodateants 1973), the form of soluble polysaccharide in which
by widening their naturallyhollow stems at fixed de- animals store excess glucose. Synthesized and
velopmental stages, predictablefrom theories of de- stored in plastids, the glycogen component of Mill-
fensive investment (see below), and temporallycor- lerian bodies lends credence to the hypothesis that
related with the onset of food body production. these rewardsmay be nutritionalmimics of herbivor-
Moreover,as describedabove by Berg, myrmecophy- ous insects or their larvae and thus may attractants
tic Cecropia also produce prostomata,or weakened which typically feed on such prey (Janzen, 1969,
sites in the stem walls beneathinternodalsepta.Pros- 1973).
tomatalie at the distal ends of the internodalgrooves Workerants often standguardat trichiliato await
that occur even in nonmyrmecophyticCecropia,per- the appearancesof Miillerianbodies. In several Ce-
haps due to pressureexertedby axillary buds (Bailey, cropia species for which the diurnal production
1922). However, their evolutionaryenlargementrel- schedules of these food bodies have been studied in
CECROPIAAND ITS BIOTICDEFENSES
the greenhouse,the rewardswere producedin greatest
profusionimmediatelyafterdark(Davidson& Fisher,
1991), coincidentally or not, the time of day when
many predator-waryherbivoresemergefrom theirdi-
urnal hiding places to settle at feeding sites for the
night. The same diurnalschedule of Mullerianbody
productionwas demonstratedfor two Cecropia spe-
cies in a project from an Organizationfor Tropical
Studies field course in Peru, near the junction of the
Rio Sucusori with the Rio Napo (pers. obs.), but a
field study by Belin-Depoux et al. (1997) in French
Guianashowedproductionpeakingthreehoursbefore
sunset in C. obtusa. Additional field studies of food
body productionschedulesin differentspecies should
shed greaterlight on the adaptivesignificanceof these
schedules. Interestingly,both the manualremoval of
previouslyemergedMiillerianbodies (Folgaraitet al.,
1994) and slight downwardpressureon petioles (pers.
obs.) can induce preemergentbodies to be released
early. The significance of pressurechanges in these
prematureemergences is the suggestion that changes
in turgorpressure,coincident with shutdownof pho-
tosynthesis and transpiration,could trigger their nat-
ural release. Fast-growingplants like Cecropiamove
back toward water balance relatively quickly after
dark,and the period immediatelyafterdarkmay rep-
resentthe startof the least expensive time (nighttime)
to producethese rewards.
Cecropiaalso producetiny, lipid-richandprotein-
containing pearl bodies ("pearlglands"to botanists)
on abaxial leaf surfaces (Rickson, 1976). These bod-
ies are not usually apparentin the field, since they are
removedrapidlyand continuouslyby obligate ant in-
habitants,or in the absence of these, by unspecialized
and opportunistic ants. However, pearl bodies are
readily found on greenhouse-grownplants and in the
field, on plants or plant partsprotectedfrom foraging
ants. In contrastto trichilia, pearl glands are simple
structures.As an alternativeor adjunctto extrafloral
nectaries,pearl bodies are taxonomicallywidespread
as a biotic defense of myrmecophilicplant species,
i.e., those that attractants with food rewardsbut not
nest sites (e.g., O'Dowd, 1982; Schupp & Feener,
1991). Their occurrenceboth in nonmyrmecophytic
Cecropia and in closely relatedMusanga and Cous-
sapoa suggests that their presence may be plesio-
morphicin the genus.
Inside host-plantstems, many obligate Cecropia-
ants also tend coccids and mealybugs (Hemiptera:
Coccidae and Pseudococcidae), and they may exca-
vate pits in external layers of nodal diaphragmsto
permitcoccids to feed on the soft internallayers con-
taining strands of conducting tissues. The numbers
215
and biomasses of Hemipteraare typically low com-
pared with those in plants that almost always house
ants but have few or no obviously ant-attractivetraits
(reviewed in Davidson & McKey, 1993). Becerra &
Venable (1989) have argued that myrmecophytes
should be selected to reduce ant-tendedHemiptera,
which not only consume plant resources but also
transmitdiseases, alter plant metabolism and devel-
opment, and are difficult to oppose by chemical de-
fenses. By provisioning symbiotic ants with mainly
carbohydraterewards, myrmecophytes may induce
the ants to balancetheirdiets by consuminga greater
fractionof theirprotein-richassociates.Interestingly,
in insular Cecropiapopulationslacking ants, at least
externallyfeeding Hemipteracan reachoutbreakden-
sities and, combined with overgrowthby vines, may
even limit the species' distribution(Janzen, 1973).
(See also Fiala et al., 1994, for indicationsthat sym-
biotic ants may protect myrmecophyticMacaranga
[Euphorbiaceae],the Asian equivalent of Cecropia,
from damaginginfestationsof pseudococcids.)
BENEFITS TO CECROPIA
Despite earliertheories postulatingno significant
effect of ants on plant fitness (Rettig, 1904; von Iher-
ing, 1907; Ule, 1906; Fiebrig, 1909; Wheeler, 1913;
Andrade& Carauta,1982; but see Muller, 1880, and
Schimper,1888), symbiotic ants appearoften to ben-
efit myrmecophyticCecropiaby protectingtheirhost
plants againstherbivores(Schupp, 1986; Fergusonet
al., 1995; Vasconcelos & Casimiro, 1997; Davidson
& Yu, unpubl.data),and/orvines andothervegetation
that potentiallycompete with the hosts for light (Jan-
zen, 1969; Schupp, 1986; Davidson et al., 1988; but
see Putz & Holbrook, 1988). The occurrenceof ant-
occupied trees with foliar damagecannotbe takenas
evidence against a beneficial role of ants, since alter-
native ant species may occur and differ in the quality
of protectionproffered.Moreover,rates of leaf pro-
duction and investmentin biotic defenses may vary
across habitatsin ways that affect the quality of de-
fense. In a long-term,experimentalstudy in Rio Pal-
enque, Ecuador (Schupp, 1986), herbivoredensities
and herbivoryrates were lower on ant-inhabitedsap-
lings of Cecropia vs. obtusifolia than on plants that
lacked ants either naturally or after experimental
removal. There, Azteca constructorwas effective in
defending its hosts against foliar herbivoryby large-
bodied, chewing insects, including leaf beetles
(e.g., Coelomera atrocaerulea, Chrysomelidae).
Protectionwas effective in the dry season but not the
wet season, when herbivores were more abundant.
Duringthe 15-monthinvestigation,saplingsdefended
216
by ants against herbivores and encroaching vines
grew significantlyfaster than did those lacking ants.
Shorter-termstudies concur that obligate Cecropia-
ants defend againstinsect herbivores.Thus, Rocha &
Bergallo (1992) showed that colony size was posi-
tively relatedto resistanceagainst herbivory.
Leafcutterants can also pose significantthreatsto
Cecropia, especially in habitats not subject to fre-
quent inundationsthat kill colonies of these ground-
nesting species. Vasconcelos& Casimiro(1997) con-
ducted an 18-month, experimental study of such
herbivoryin central Brazil. Frequencyof attackwas
not related to leaf palatabilityas assessed in feeding
trials, but for threeof the four species includedin the
study,to occupationby colonies of Azteca ants.Thus,
although the fastest-growingspecies (Cecropia dis-
tachya)was most likely to be attackedwhen antswere
absent,and to be favoredby leafcutterants in feeding
trials, occupied saplings of this species contained
largerworkerpopulationsthandid stems of a slower-
growing species, and theirleaves also benefitedmost
from ant attendance. (Likewise, in myrmecophytic
Asian Macaranga, species suffering most from ant
removalare those apparentlyproducinggreaterquan-
tities of ant rewards, which in tum support larger
workerpopulations[Itiokaet al., 2000]). The slower-
growing species (C. ulei) was least preferredby leaf-
cuttersin feeding trials and among plants lackingAz-
teca, but it was also least likely to be colonized by
Azteca and most likely to lose its Azteca colonies. It
is possible that slow-growing species invest more in
physical and chemical defenses of leaves (see below).
Across all four host species, more attackedthan un-
attackedsaplings died duringthe observationperiod,
though the difference was not significant at small
samplesizes, andplantsattackedmorethanonce grew
more slowly than did those attackedjust once or not
at all.
Most studies of the effects ants on herbivoryof
myrmecophyticand myrmecophilic plants have fo-
cused on workerprotectionof leaf blades (e.g., Beat-
tie, 1985). However,the location of Miillerianbodies
at the bases of petioles in Cecropia attractsworkers
mainly here and to adjacent stems. The principal
value of symbiotic ants (as opposed to that of oppor-
tunistic, nonresidentspecies that consume just pearl
bodies) could thereforebe the protectionthese sym-
bionts affordpetioles and stems. (See also Fiala et al.,
1994, for a potentiallysimilarsituationin Macaranga
[Euphorbiaceae].)At least some leafcutterants (Az-
teca laevigata) damageor destroyterminalmeristems
of Cecropia and remove entire leaves by severing
FLORANEOTROPICA
theirpetioles (Vasconcelos& Casimiro,1997). More-
over, thick-stemmed(pachycaulous)plants are often
colonized by stem-boringinsects such as the weevils
(Coleoptera,Curculionidae)that are extremely com-
mon in Cecropia stems at higher elevations in the
tropics (H. Hespenhide,pers. comm. for Costa Rica;
pers. obs., for at least 1500-2000 m in the easternAn-
des of southernPeru).At lowerelevations,leaf beetles
(Chrysomelidae)plague young Cecropia throughout
their South American distribution (Andrade, 1981,
1984a; Schupp, 1986; Jolivet, 1989; Davidson &
Fisher, 1991; Rocha & Bergallo, 1992; Jolivet & Sa-
linas, 1993). Some of these beetles (e.g., in the genus
Coelomera)lay eggs inside stems, wherelarvaehatch
and emerge to feed on either leaves or their petioles
(Andrade,1981, 1984a; pers. obs.). Whereherbivory
rates are high, Cecropiaseedlings are hard-pressedto
stay aheadof herbivoryby producingnew leaves just
as the previous ones are either removed entirely or
largely consumed, with only skeletons of veins left
behind (pers. obs. along rivers of southeasternPeru).
The severing or weakening of petioles can be disas-
trous for plants like Cecropia,which invest many re-
sourcesin very largeindividualleaves. Like leafcutter
ants, stem-inhabitingbeetles can also damage termi-
nal meristems,slowing the verticalheightgrowthnec-
essary for Cecropia to overtop other fast-growing
competitorsin light gaps andotherdisturbedhabitats.
For seedlings and saplingswith one or very few meri-
stems, this type of damage would likely convey sub-
stantialmaterialand opportunitycosts.
Given the diversetypes and high levels of damage
attributableto stem-dwellingherbivores,it is possible
to conclude thatthe primaryimportanceof symbiotic
antcolonies (vs. opportunistsfeeding on pearlbodies)
is to precludestem occupancyby herbivorousbeetles,
perhaps especially in vulnerable young Cecropia
seedlings and saplings. In accordwith this argument,
an unusual Cecropia species, C. hispidissima, illus-
trates how protection of leaves and stems may be
fundedby differentant rewardsthat are monopolized
by differentants. Stem-inhabitingcolonies of Pachy-
condyla sp. nov. 1 have been found to date only on
Cecropia hispidissimain Panama,and their workers
apparentlyharvestMiillerianbodies frombeneaththe
stipules and without ever visiting the leaves (T. Kur-
sar,pers. comm.). Nevertheless,atypicallylarge,hard,
andpurpleMillerian bodies of this host would appear
to be adaptedfor use by the large-bodiedPachycon-
dyla ratherthan by tiny Azteca ants. Pachycondyla
workers do not exclude other ants, such as those of
nonresidentAzteca colonies, from frequentopportun-
CECROPIAAND ITS BIOTICDEFENSES
istic foragingon leaves, probablyfor pearl bodies. In
a hypotheticalancestorof myrmecophyticCecropia,
opportunisticants may have harvested pearl bodies
and protectedleaves without alleviatingselection for
ways of ridding the plant of stem-dwelling herbi-
vores. Mullerian bodies and prostomatamay have
evolved under such selection pressures. Moreover,
protection of stems from stem-boring insects may
start as soon as queens begin to colonize myrmeco-
phytic Cecropia;many or most internodesof individ-
ual seedlings and saplings are colonized indepen-
dently by one or more queens, and queens of some
species do feed while developing their first worker
broods (see below).
Accounts of other myrmecophytes also suggest
that ants may protect against stem-boring insects.
Thus, Pheidole ants of myrmecophyticPiper in Cen-
tralAmericahave been shown to protecthosts against
stem-boring weevils (Letourneau, 1998). Replace-
ment of stem-boringplantparasitesby antsis thought
to have been an early stage in the evolution of myr-
mecophytismin other plant taxa with primarydom-
atia (e.g., Ward,1991). Although ants probablyoften
tended Hemipterawithin these domatia, the net ef-
fects of ants and Hemipteraon plants may have been
positive, especially in habitats where abundantlight
allowed rapidcarbongain to offset losses to Hemip-
tera (Davidson & McKey, 1993). Herbivorepressures
may also have been more intense in comparatively
productive,sunny environments(Davidson & Fisher,
1991; Davidson & McKey, 1993; Davidson & Yu, un-
publ. data), reinforcingselection for ant attraction.
Finally, for Cecropia species inhabitedby more
than one species of ant, the quality of defense may
vary with the identity of the partner.Obligate sym-
bionts of Cecropia differ in their diurnal foraging
schedules, the extent of their activity and aggression,
the numbersand sizes of workers,and whetheror not
workersprune vines and other vegetation (Davidson
et al., 1991; Davidson & Fisher, 1991; Longino,
1989b, 1991). Evolution on the partof the plant may
produce adaptationsthat favor one ant species over
another,but to some extent, the predominantinhabi-
tant may be determinedby the outcome of ant-ant
competition and be beyond the plant's control (Da-
vidson & McKey, 1993; Yu & Davidson, 1997).
Poorly defending ants may sometimes gain posses-
sion of their hosts by virtue of rapid colony devel-
opment,or pleometrosis,i.e., colony foundingby two
or more queens, whose combinedfirstworkerbroods
should producelargerand more competitiveincipient
colonies (e.g., Davidson et al., 1991).
217
INTERSPECIFICVARIATION IN
DEFENSIVE INVESTMENT
Early ecological studies of Cecropia were under-
takenmainly in the CentralAmericalowlands,where
species diversityof both the host plantsand theirants
is relatively low. It is not surprising,then, that rela-
tionshipsbetween Cecropiaand its ants were initially
consideredratheruniform.Thatis, all myrmecophytic
Cecropia produced Muillerian(and perhaps pearl)
bodies, and despite very early reportsto the contrary
(Wheeler, 1942), all Cecropiawere implied to house
Azteca ants (Janzen, 1969, 1973; Rickson, 1976,
1977). Not until the early 1990s did researchbegin to
focus on interspecificvariationin the ant-attractants
of Cecropia and the significance of this variationto
ant associates (Longino, 1989a; Davidson et al.,
1991; Davidson & Fisher, 1991; Yu & Davidson,
1997). It now appearsthat there is importantand in-
teresting interspecific variation in the ontogeny,
quantity,and compositionof biotic defenses, andthat
some of this variationboth accordswell with general
theories of plant defensive investmentand is impor-
tant to ants.
Plantdefense theorybegins by postulatingthatde-
fenses are costly and are manufacturedand used only
when benefits, calculated in reduced herbivory,ex-
ceed costs. The costliness of biotic defenses in Ce-
cropia is perhaps most apparentwhen ant-attractive
traits are lost in the absence of selection imposed by
the typical ant associates. It is interesting,then, that
populationsof C. peltata on Jamaicaboth occurin the
absence of their ants and are polymorphic in their
expression of trichilia (see treatmentof C. peltata in
Berg, above).
Several other observationsalso provide evidence
for a cost of biotic defenses in Cecropia.First, food
bodies are producedmainly at stages of leaf devel-
opment and whole-plantdevelopmentwhen they are
aptto do the most good, includingwhen chemicaland
physical defenses arepoorly developed.In the species
surveyedto date, productionof eitherMuillerianbod-
ies, pearlbodies, or both occurs within days of leaves
first beginning to emerge from their stipules, and
peaks relatively early in leaf lifetimes (week 1 for a
pioneerspecies, andweek 5-6 for a small gap species:
Folgarait & Davidson, 1995). Young leaves are es-
pecially vulnerableto herbivores(Davidson& Fisher,
1991), probablydue to theirhigh foliarnitrogenlevels
andpoorly developedphysical and chemicaldefenses
(Folgarait& Davidson, 1995). In addition,the value
of a leaf to the plant is greatest at this stage, since
most of its productivelife span lies ahead (Harper,
218 FLORA NEOTROPICA

"Reclaimable"extrafloralnectar
and pearl bodies

Cumulative "No n-reclaimable" tannin &

cost of defense lignin or Mullerian bodies

Short B Long

Leaf lifetime
FIG. 1. The McKey (1984) model of plant defenses, modified to accommodatethe biotic defenses of Cecropia.
Cumulativecosts of "reclaimable"biotic defenses (e.g., pearlbodies of Cecropia)are lower thanthose of "non-reclaimable"
defenses (e.g., tanninsand lignins, or trichilia and Mullerianbodies), for relatively short-livedleaves and other plant parts.
However, they are higher in species for which "non-reclaimable"defenses can be amortizedover the life spans of long-
lived leaves.

1989). In relation to whole-plant development,tests
of leaf palatabilityto herbivoressuggest thatCecropia
seedlings are better protectedby chemical and phys-
ical defenses before than afterthe plants acquiretheir
ants (Martinelliet al., 1993).
Three final patternsare indicativeof a cost to the
productionof food rewardsfor ants. First, both Mill-
lerian and pearl bodies are responsive to supplies of
key resources (Folgarait & Davidson, 1994, 1995).
Fertilization(N-P-K) and high light intensityincrease
Mullerianbody productionper unit time and per ac-
tive lifetime of trichilia. In contrast,pearl body pro-
duction is enhanced by nutrients but not by higher
light intensities, perhapsbecause these rewardshave
highernitrogencontentthando Muillerianbodies, and
because an increase in carbon fixation under high
light drawsnitrogenaway frompearlbody production
and into growth as the first priority. Second, across
species, there appears to be a trade-off between in-
vestmentin biotic vs. chemical and physical defenses
(Vasconcelos & Casimiro, 1997). Third,both the rate
of Miillerian body production,and the total produc-
tion per lifetime of a trichilium,increase in response
to their manualremoval, simulatingharvestingin the
presence of ants (Folgaraitet al., 1994).
Grantingthen that defense can be costly, two im-
portanttheories have relateddefensive investmentei-
ther to resource availability(Coley et al., 1985) or to
leaf life spans (McKey, 1984). The resource avail-
ability theory startswith the premisethatplantgrowth
rates vary in response to the availability of limiting
resources,typically light in tropicalforests. Plantspe-
cies characteristic of relatively light-rich environ-
ments have evolved rapid growth and low levels of
defense, since costly defensive investmentwould ex-
act opportunitycosts in the form of reduced growth
that could lead to overtopping by fast-growing
competitors. In contrast, slow-growing species of
resource-poor(shaded) habitats would have little to
lose by defending (i.e., low opportunitycosts) and
perhapsmuch to gain, since low resourceavailability
preventsrapidreplacementof lost tissues.
In contrast,McKey's leaf lifetime hypothesissug-
gests thatplantsshoulduse differentkindsof defenses
dependingon whetherleaf lifetimes are shortor long
(Fig. 1). Relatively fast-growing plants with short-
lived leaves should invest mainly in "reclaimablede-
fenses," equated by McKey to biotic defenses (ex-
trafloralnectar and, presumablyalso, pearl bodies).
These defenses have low initial constructioncosts but
significantmaintenancecosts, so cumulativecosts in-
crease linearly through time. As leaves age and ac-
quire other defenses such as toughness, the resources
investedin biotic defenses can be routedto new leaves
at little expense. Therefore,biotic defenses are often
producedon the newest, most valuable,and most vul-
CECROPIAAND ITS BIOTICDEFENSES
nerable foliage. In contrast, McKey postulated that
comparatively slow-growing plants with long-lived
leaves could affordto invest in "non-reclaimable" de-
fenses, such as tannins, lignin, fiber, etc. Once laid
down, these defenses were (theoretically)permanent
andcould not be brokendown andremobilizedto new
tissues. Only in plants with long-lived leaves could
the high initial constructioncosts of non-reclaimable
defenses be amortizedover a sufficientlylong period
so that their cumulativecosts would be exceeded by
the cumulativecosts of "reclaimable"defenses.
Interspecificdifferencesin investmentin the biotic
defenses of Cecropiaare well explainedby these the-
ories if a distinction is made between "reclaimable"
pearl bodies and less reclaimable Muillerianbodies
and trichilia,with the latterhaving higherinitial con-
structioncosts (Folgarait& Davidson, 1994, 1995).
The more light-demanding"pioneer"Cecropia spe-
cies, typical of very open (often riparian)habitats,
have the relativelyrapidgrowthrates and short-lived
leaves characteristicof other tropicalrain forest pio-
neers. In accord with McKey's theory, such species
tend to invest proportionatelymore heavily in pearl
bodies andless heavily in Muillerianbodies (evaluated
as dry weight of rewardper unit of leaf area)thando
related but more shade-tolerantspecies with slower
intrinsic growthrates and longer leaf life spans (Fol-
garait & Davidson, 1994, 1995). Greaterinvestment
in Muillerianbodies is also consistent with the re-
source availability theory of plant defensive invest-
ment, because leaf lifetimes tend to be highly corre-
lated with (and lower in) light-richhabitats(Mooney
& Gulmon, 1982).
Using the McKey (1984) model as originally for-
mulated to compare myrmecophytesand myrmeco-
philes with plants lacking biotic defenses altogether
may also help to explain the distributionof nonmyr-
mecophyteswithin the genus Cecropia.Investmentin
non-reclaimabletannins, fiber, and lignin would be
predicted to replace partially or mainly reclaimable
biotic defenses in plantswith comparativelylong leaf
life spans. It is interestingin this context to note that
nonmyrmecophytic Cecropia sciadophylla, a rela-
tively slow-growingCecropiaspecies of infertilesoils
on higher terraces in Amazonia, produced longer-
lived leaves than did any of eight myrmecophytic
Cecropia species with which it was grown simulta-
neously, or which were grown in other experiments
in temperategreenhouses(D. Davidsonet al., unpubl.
data). Because of the correlation between intrinsic
plant growthrates and mean leaf life spans (Folgarait
& Davidson, 1994, 1995), a decline in myrmecophy-
tism with decreasing intrinsic growth rates at high
219
elevation might also be anticipated,but a lack of ap-
propriateant species could also play a role (Wheeler,
1942). Finally,plant defense theories based on either
leaf life spans or resourceavailability,and developed
explicitly for foliar defenses, should also apply to de-
fenses of other plant parts, such as stems benefiting
from ant occupation;this is particularlytrue consid-
ering that new stems of Cecropia tend to be photo-
synthetic.
Interspecific differences in Cecropia reveal an-
other patternthat was not previously predicted,but
was readily apparentbecause biotic defenses are eas-
ily observableexternally (Davidson & Fisher, 1991;
Folgarait& Davidson, 1994, 1995). The onset of (bi-
otic) defense occurs earlier in seedling development
for relatively slow-growing, shade-tolerantCecropia
than for their faster-growing,light-demandingrela-
tives-a patternthatcould provetypicalin other(e.g.,
chemically defended) genera of tropical trees but is
currentlypoorly explored.Earlyin seedlingontogeny,
leaves are producedwithouttrichilia,and stems both
lack prostomataand are too narrowto accommodate
antcolonies. In commongardenexperiments,the pro-
duction of active trichilia and the swelling of stems
concurrentwith the appearanceof prostomata,occur
earlierin time andearlierin developmentin relatively
slow-growing, shade-tolerantCecropia species than
in their faster-growingrelatives.
Some of the most distinctivemyrmecophyticCe-
cropia occur in small light gaps, uncharacteristically
shadedhabitatsfor this largely "pioneer"genus. Low
light slows growth,increasingthe probabilitythatAz-
teca colonies will fail (Vasconcelos & Casimiro,
1997), yet it also prolongs rates of replacementof
leaves lost to herbivores.In this context, some species
have evolved very early developmentalswitches to
myrmecophytism. For example, in western Ama-
zonia, trichiliafirstappearon medianleaf numbersix
in "Cecropiapungara,"though at leaf number 13 in
the apparentlyclosely relatedCecropiamembranacea
(Davidsonet al., 1991). (Based on gene sequencedata
[S. Cook and D. Davidson, unpubl. data], "C. pun-
gara" appearsto be a distinctspecies but is treatedas
a "form"by Berg & Franco[this volume, see discus-
sion under C. membranacea].)Remarkably,in "Ce-
cropia herrerensis"of northeasternPeru,trichiliaare
produced on the first set of leaves after cotyledons.
From the perspective of the ants that colonize their
seedling hosts as soon as swollen stems with prosto-
mata appear,tiny, slow-growingjuveniles of small-
gap Cecropiaare a far differentresourcethanaretall,
rapidly growing pioneer Cecropia of light-richhabi-
tats. Althoughsmall-gapspecies may supplymorere-
220
sources to theircolonies per areaof leaf surface(Fol-
garait & Davidson, 1994, 1995), rates of resource
supply per plant are the more importantfactor to de-
veloping colonies, and those rates are greaterin com-
parativelyfast-growingpioneer relativesof the small
gap species. Therefore, it is not surprisingthat the
early-defending, shade-tolerantCecropia can have
unusualant associates(Davidson& Fisher,1991). For
example, "C.pungara"is inhabited,not by any of the
Azteca species (subfamily Dolichoderinae)so wide-
spreadon Centraland South American Cecropia,but
by Pachycondylaluteola (Ponerinae)and Campono-
tus balzani (Formicinae),and the typical associate of
"Cecropiaherrerensis"in northeasternPeru is Cre-
matogasteraff. curvispinosa(Myrmicinae).
ANTS SYMBIOTICWITH CECROPIA
Table 1 presentsa list of obligate Cecropia-antsas
we presentlyknow them.Most impressiveis theirtax-
onomic diversity.All ants belong to the same insect
family, the Formicidae,and the majorsystematiccat-
egories of ants are the subfamilies.The associates of
Cecropiarepresentfour ant subfamilies,includingall
but one of those in which at least some species are
known to rely heavily on plant resources.Absent are
only the Pseudomyrmecinae,stem-nestersfound fre-
quently in narrow twigs, where workers can effec-
tively use theirheads to block small entrancesagainst
invasions of competing and predatoryants.
The most common and best-known associates of
Cecropia belong to the dolichoderinegenus Azteca.
This group is endemic to the New World,and its di-
versity and abundance are greatest in the lowland
tropics. Approximately 63 species and numerous
named varieties remainin the genus afterreconciling
Brandao (1991) and Longino (1991) with Kempf
(1972), and 13 of these species are specialists on Ce-
cropia (Table 1). The genus includesmanyfree-living
species as well as specialized plant-antsassociated
with myrmecophytesin at least 16 generaand 9 fam-
ilies (Davidson & McKey, 1997). Azteca ants tend to
have populous colonies, finely divided into many
small workers that tend Hemiptera (Coccidae and
Pseudococcidae),scavenge, and hunt live prey, (usu-
ally insects, and sometimesincludingcompetingants;
Carroll,1983; Adams, 1990a).Based on nitrogeniso-
topic ratios, various Azteca species place among the
most predatory of arboreal ants (Davidson et al.,
2003) and are the most nitrogen-deprivedof neotrop-
ical ants (Davidson, in press). Abundant carbohy-
dratesfrom hemipteranhoneydew may fuel both the
large workerpopulations,which are maintainedprin-
cipally by carbohydratesratherthan by protein re-
sources (Davidson, 1997), and the infamous aggres-
FLORANEOTROPICA
sion exhibited by these ants (Carroll, 1983).
Monogynous (single-queen) colonies are often poly-
domous, i.e., occurring as numerousaffiliatednests
distributedover one or morecrowns.The largecolony
sizes of Azteca are correlated with low population
densities, perhapsreflectinghigh mortalityof incipi-
ent colonies with comparativelyfew hemipteranas-
sociates and thus limited protein resources. Wilson
(1985 & pers. comm.) has arguedthatlow population
densities may be correlatedwith high local extinction
rates that could account for the extinction of Azteca
from West Indianislands, despite theirpresencethere
in fossil ambersfrom the late Oligocene or earlyMio-
cene.
Interspecificand intraspecificspatialterritoriality
have been demonstratedfor a numberof Azteca spe-
cies (Adams, 1990a, 1990b, 1994; see also Leston,
1973, and Carroll, 1979) and may have preadapted
membersof this genus for defense of myrmecophytic
host plants. Cecropia and other myrmecophytespro-
vide carbohydrateswhich fuel large, aggressive, and
territorialworker populations (Davidson & McKey,
1993; Davidson, 1997). Azteca colonies subdividela-
bor by physical subcastes,with (mainly)majorwork-
ers stationedon limbs at key branchintersections,at
the bases of plants, and at colony borders (Adams,
1990a, 1990b, 1994). Defending workersfrequently
exhibit characteristicpostures, with mandibles open
and gasters upraised,ready to release carbon-based
alarmpheromonesshould enemy ants approach.Rel-
atively long-distancerecruitmentof nestmates (over
meters)is apparentlymediatedin Aztecaby trailpher-
omones from the Pavan'sglands. Signalingfood finds
and breaches in colony defense, short distance re-
cruitment (over several centimeters) occurs in re-
sponse to alert/alarmproducts(cyclic ketones) of the
pygidial gland, as well as throughworker-to-worker
contact and tactile displays. Outcomes of territorial
interactionsappearto be determinedby mutual as-
sessment of asymmetriesin colony size, mediatedin
partby recruitmentresponsesthatescalatewith prox-
imity to nests and nestmatereinforcements.With in-
creased aggressiveness at or near nest sites, resident
advantagecan bluntadvancesof numericallysuperior
colonies. Finally,because the traitssummarizedhere
occur in free-living Azteca species, they almost cer-
tainly evolved in the context of territorial defense
against other ants. Although many of the same traits
undoubtedlypreadaptedAzteca to protect their host
plants, arguments that such behaviors of Azteca
evolved by selection on Cecropia for "inducedde-
fenses" (Agrawal, 1998) are currentlynot convincing
because they ignore the basic biology of ants in this
genus.
CECROPIAAND ITS BIOTICDEFENSES 221

TABLE 1
Known or presumedobligate Cecropia-ants.a

Subfamily & Species GeographicDistribution Reference

DOLICHODERINAE
Azteca spp. Longino, 1989, 1991
A. alfarib CentralAmerica throughAmazonia
A. aragua AraguaState,Venezuela
A. australis W Amazonia (Peru,Bolivia), Amazonas,Brasil
A. coeruleipennis S Mexico throughNW Costa Rica
A. constructor GuatemalathroughGuyana
A. isthmica Panamaand Colombia
A. lattke Venezuela(coast ranges & Cordillerade Merida)
A. merida E slopes, Cordillerade Merida,Venezuela
A. muelleri S Brasil
A. ovaticepsb Lowlandtropics of Centraland South America
A. petalocephala SW of Coroico, (La Paz), Bolivia
A. salti SierraNevada de SantaMarta,Colombia
A. xanthochroa Mexico throughPanama
PONERINAE
Pachycondylaluteola W Amazonia Davidson et al., 1991
P prov. dianaec SE Panama W. L. Brown, in ms
P ''insignis''c Mid-elevationwet forest, Costa Rica J. Longino 2000d
P unidentatarugulosae Emery 1902
FORMICINAE
Camponotusbalzani Amazonia Davidson et al. 1989, 1991;
Benson 1985
MYRMICINAE
Crematogasternr. curvispina Descent from E Andean cordillerain SE Peru D. W. Davidson, unpubl.
Pheidole sp.
aOtherants may nest opportunisticallyin Cecropiastems withoutusing prostomataor Miillerianbodies (Longino, 1989a;
Davidson & Fisher, 1991). This phenomenonis especially common in plantswith obligate associatesnestingpolydomously,
only in growing tips. Polydomy,or fragmentationof single colonies over multiplenest sites (here, disjunctstems of a single
host plant) is characteristicof Azteca alfari and A. ovaticeps (Longino, 1989b).
bMembersof the Azteca alfari groupthoughtto have descendedfrom ancestorsthatnested in live stems (Longino, 1989,
1991). The remainingAzteca species are proposedto be descendantsof carton-nestingancestors.
cThese names are conditional proposals (sensu Art. 15 of the 1985 ICZN) by the late W. L. Brown Jr., and thus not
made availablehere. Their appearancehere or in any duplicationof this article does not constitutepublication.
dSee Longino (2000) for naturalhistory, distribution,etc. The species appearsto be a specialist on sapling Cecropia,
and colonies maturebefore saplings grow to tree size. Occupationof saplings by this species may lead to the demise of
seedlings by precludingcolonizationby the more typical Azteca ants.
eThis variantof P. unidentatawas synonymizedwith P. unidentatain an unpublishedmanuscriptby W. L. Brown Jr.,
but sentimentremainsamong myrmecologistsfor consideringit to be a specialist on Cecropia saplings.

Among the usuallypredaciousand scavengingPo- species are arboreal,nesting opportunisticallyin cav-

nerinae,thereappearto have been no fewer thanthree
(and perhaps even four) independent evolutionary
colonizations of Cecropia (Table 1). All of the po-
nerines specialized to live in Cecropia are species in
the genus Pachycondyla,relatively large-bodiedants
occurring in both the New and Old World tropics.
Most Pachycondylaspecies are terrestrial,but some
ities of dead or live stems, or in and aroundepiphytes
and debris, in the upperor lower rainforestcanopies.
Of the arborealspecies, several are known to occupy
abandoned intemodes of seedling and sapling Ce-
cropia. Among the Pachycondylaspecialized to live
in Cecropia, Pachycondyla"insignis"appearsto be
related to P. villosa and P bugabensis, two general-
222
ized arborealpredators(Longino, 2000). Pachycon-
dyla luteola and Pachycondyla (provisionally)
"dianae"(W. L. Brown manuscript),are morpholog-
ically distinctive from one another and from Pa-
chycondylaspecies nov. 2, as well as geographically
disjunct in their distributions.Host-specific on "Ce-
cropia pungara,"Pachycondylaluteola departsmost
notably from other ponerines in behaviorand colony
size, typicallyjust tens to hundredsof workersin the
genus as a whole (Peeters, 1997). One might specu-
late thathemipterantendingmay occur in this species
and provide carbohydratesto subsidize the large
worker populations. However, the fearsome stings
of these aggressive ants-painful over weeks to
months-have deterredinvestigatorsfrom opening P
luteola nests in all but small seedlings and saplings
where Hemiptera are least apt to have colonized.
Workersboth pruneencroachingvines andexhibitab-
solute fidelity to theirhost-plants,feeding principally
on food rewardsprovided by the plant (Davidson &
Fisher, 1991). As is typical of ponerines, queens do
not found their colonies claustrally;instead they col-
lect, store and use Muillerianbodies in the early es-
tablishmentof their colonies. Thus, they must leave
the prostoma open in order to reach the trichilia, a
practicethat makes them susceptibleto loss of brood
to parasitoid wasps (Perilampidae, Davidson &
Fisher, 1991). External foraging by nonclaustral
queens (frequently multiple, competing queens per
colony) may provide some protection to juvenile
plantsimmediatelyaftercolonizationandpriorto col-
ony establishment.
Formicines and Myrmicines are also represented
among the associates of Cecropia. Camponotusbal-
zani (subfamily Formicinae),a host generalist,is ex-
tremely timid and largely nocturnal,and seldom ven-
tures away from stems to patrol leaves (Davidson &
Fisher, 1991). Another formicine, a species of Myr-
melachista, is not listed in Table 1, because rather
than collecting and feeding on Muillerianbodies, it
appearsto feed on the abundantqueens of Azteca spp.
that colonize a variety of Cecropia along the eastern
escarpmentof the PeruvianAndes (pers.obs.). Within
the Myrmicinae,Crematogaster(Table 1) is a genus
of ecologically dominantants thatare most diversein
the Old Worldtropics, where they are frequentlyob-
ligate plant-ants(Davidson & McKey, 1993). Their
possibly late arrivalin the New Worldtropics(Brown,
1973) might account for why small-bodied, stem-
nesting and carton-buildingants in the endemic neo-
tropical genus Azteca were able to diversify and be-
come codominant with this ecologically similar
genus. The species associatedwith "Cecropiaherrer-
ensis" is most closely related to C. curvispinosa, a
FLORANEOTROPICA
generalizedcavity nester,widespreadin the Neotrop-
ics (J. T. Longino, pers. comm.) and found occasion-
ally in Cecropia at high elevations (pers. obs.). Al-
though colonizing queens have occasionally been
found with incipient colonies on "C. herrerensis,"
sometimes cofounding with conspecific queens,
largercolonies are more often resident without their
queens, which may either occupy trunksof adjacent
Cecropiaor live off the host altogether.Nevertheless,
documentationof queen colonization and the collec-
tion, storage, and use of Muillerianbodies suggests
that the ants are specialized associates of Cecropia.
A third myrmicine, in the cosmopolitan genus
Pheidole, is listed here on the basis of a single col-
lection in Cecropiasp. at relativelyhigh elevation(ca.
1500 m) on the descent from the eastermcordilleraof
the Andes in southeasterm Peru(D. Davidson,unpubl.
data). Although the queen was not located, a large
colony occupied numerousinteruodesof a relatively
tall plant, and was harvestingand storing Mullerian
bodies nearits brood.Until morecolonies of the Phei-
dole are located, and queens are shown to recognize
and colonize these hosts, the lone recordmust be re-
garded with suspicion. If the ant is eventually con-
firmedas an obligate symbiontof Cecropia,the find-
ing would be consistent with the pattern of novel
associates occurringat the elevationaland latitudinal
limits of the distributionsof myrmecophytes(David-
son & McKey, 1993; McKey & Davidson, 1993).
Even among the predominantassociates of Ce-
cropia in the genus Azteca, there appearto have been
multiple evolutionarycolonizations of myrmecophy-
tic hosts. Based on both morphologicaland behav-
ioral evidence (Longino, 1991), and gene sequence
data(Ayalaet al., 1996), at least two independentline-
ages of Cecropia-antshave been distinguished.One
lineage, the Azteca alfari group (A. alfari andA. ova-
ticeps), is likely derivedfrom ancestorsthatnested in
live stems. The other,the A. muellerigroup (A. muel-
leri plus A. aragua, A. australis, A. isthmica,A. pe-
talocephala,A. salti, A. xanthochroa,and possibly A.
constructorand A. merida), has probablydescended
from carton-nesters,and species in this lineage still
buildcartonnests withintheirhosts. Cartonnests con-
sist of masticated plant material, sometimes mixed
with ant or vertebratefeces and soil (reviewedin Lon-
gino, 1986; Davidson & Epstein, 1989), all building
materialsthat are availablepracticallyanywhere.Al-
though the occurrenceof stem-nestersmay often be
limited by the availabilityof suitable stems, nesting
in cartonfrees ants to locate theirnests nearabundant
food. Thus, the two Azteca lineages may have had
very differenthistories of associationwith Cecropia.
Carton-nestingancestorsof contemporaryCecropia-
CECROPIAAND ITS BIOTICDEFENSES
ants may have initially sought out Cecropia because
of its pearl bodies (see above), just as other carton-
nesting taxa are sometimes overrepresentedon hosts
supplying extrafloralnectar (Hoildobler & Wilson,
1990; Davidson & Epstein, 1989). In contrast,stem-
nesters would have been restrictedto habitatswhere
suitablehollow stems were available;wide stems with
weak pith are characteristicof pioneer plants in dis-
turbedhabitats.
This scenario is consistent with experimentalev-
idence sortingout the contributionsof host specificity
andhabitatspecificityto the determinationof pairings
between Cecropia and its ants. Yu and Davidson
(1997) cultivatedseveralCecropiaspecies fromseeds
in screen tents in Amazonian Peru, until all of the
species had expandedtheir stems and producedpros-
tomata. They then placed them out in two habitats,
riverine edge and forest gaps (createdor augmented)
and monitoredcolonization over 2-3 months.Azteca
ovaticeps, a member of the stem-nesting A. alfari
group,provedto be habitat-specific(to riverineedge)
and not host-specific, but the reverse was true of A.
australis of the carton-nestingA. muellerigroup.The
relativelysmall thoraxand reducedwing musculature
of A. ovaticeps queens (Davidson et al., 1991) was
consistentwith its inabilityto seek out potentialhosts
in gaps dispersed throughoutthe forest. In contrast,
with its large thorax, presumablycorrelated with a
greaterwing muscle mass, A. australis queens were
successful at locatingseedlings in bothhabitats.How-
ever, in behavioralassays, queens preferredto colo-
nize species with high pearl body production;they
also were underrepresentedamong foundresses on
hosts with low pearlbody productionandsuccessfully
producedbrood during the experimentalperiod only
on the preferredhosts (Yu & Davidson, 1997). (The
dearthof pearlbodies themselves is not likely to have
caused failure of brood production,since the queens
found their colonies claustrally.)
A parsimoniousinterpretationof the history of the
relationshipsbetween the two ant lineages and their
hosts might thereforesuggest that the two were dif-
ferently preadaptedto use Cecropia,and that aspects
of their preadaptationssurvive to influence their use
of these myrmecophytestoday.Thus,Aztecaaustralis
and its allies occur naturallymainly on hosts with
high pearl body production (Davidson & Fisher,
1991; Yu & Davidson, 1997; Folgarait& Davidson,
1994, 1995). In contrast,A. ovaticeps and its sister
species, A. alfari, are restrictedto large-scaledistur-
bances throughouttheirranges(Longino, 1989, 1991;
Davidson & Fisher, 1991). Interestingly,in green-
house experiments, A. alfari did not even feed on
pearl bodies (Baird 1967).
223
This overview of Cecropia-antssuggests thatAz-
teca were likely the original associates of Cecropia.
Not only are they the most diverse and widespread
among the many symbionts of these ant-plants,but
species in the Azteca muelleri group (including A.
australis) may have built cartonnests on these plants
to use pearlbodies, even before Mullerianbodies had
evolved. For this and other reasons, the muelleri
group may have been first among the Azteca species
to inhabit myrmecophyticCecropia. In seeking out
host plants with pearl bodies, queens of carton-
nesting Azteca might have initially colonized larger
plants, not seedlings, and it is difficultto believe that
these foundressescould have initiated their colonies
on plants with preexisting colonies of stem-nesting
species. On the other hand, queens of stem-nesters
may very well have gotten their starton Cecropiaby
colonizing seedlings that had not yet acquiredother
ants. As for the Cecropia-antsin other genera,these
are almostcertainlyall derivedfrom stem nesters,and
with the exception of Camponotus balzani (wide-
spreadin the Amazon basin), are all species with lim-
ited biogeographicand/orhost rangesthatsuggestrel-
atively recent origins of their relationships with
Cecropia.A subset of these ant species, in the genera
Pachycondylaand Crematogaster,also coexist with
much more widespreadand common species that are
their likely progenitors(Table 1).
The richness and diversity of the ants symbiotic
with Cecropiaimply thatcolonizationof Cecropiaby
unspecializedants over evolutionarytime has been a
relatively easy transition.At least two factors could
have facilitated this transition.First, for many ants
with generalized diets, long-lived and secure nest
sites, such as those affordedby cavities in live plants,
may be more limiting than are food resources. Sec-
ond, pachycaulous stems with weak pith probably
evolved in Cecropia as a form of minimal support
structurefor large leaves (Halle et al., 1978; White,
1983; Davidson & McKey, 1993). Once evolved,
these largestems were easily cooptedas desirableand
accessible nest sites for ants of diversebody sizes and
colony sizes. In the context of the susceptibility of
stems to exploitation by damaging herbivores (see
above), such ants may have provided net benefits to
their hosts and exerted selection pressures favoring
the evolution of other ant-attractivetraits.
COEVOLUTIONOF ANTS AND CECROPIA
Coevolution consists of coadaptationand cospe-
ciation (Brooks, 1979). Coadaptionsignifies thateach
species in the partnershiphas evolved one or more
traitsin responseto selective pressuresexertedby the
other.There is little questionthatthis has occurredin
Cecropia and its ants, but since most Cecropia-ants
224
are found on more than one host species, and many
Cecropia species can be inhabitedby more than one
ant associate, coadaptationlikely was diffuse. In dif-
fuse coadaptation,selection pressuresresponsiblefor
the evolutionof a traitin one partnercome from a set
of associates, ratherthan from one species in partic-
ular (Janzen,1980).
What traitsare indicativeof coadaptation?In Ce-
cropia, prostomataand trichilia with Mullerianbod-
ies are surely characteristicsthathave evolved due to
selection pressures exerted by symbiotic ants, and
pearl bodies could have evolved in responseto selec-
tion imposed (originally)by opportunistic,as well as
(later) symbiotic associates. Moreover, once these
traits had evolved, there is evidence that they may
have been modified by selection pressuresgenerated
by particularant species (Davidson et al., 1991; Da-
vidson & Fisher, 1991). Thus, the unusuallylarge and
protruding(ratherthan recessed) prostoma of "Ce-
cropia pungara,"togetherwith its much largerMill-
lerian bodies, may be interpretedas adaptativere-
sponses to its occupation by relatively large-bodied
Pachycondyla and Camponotus species. Pachycon-
dyla luteola is also unique among well-studied
Cecropia-antsin its nutritionaldependencyon Mill-
lerian bodies during the earliest stages of colony
founding, and despite this dependency, P luteola
queens refuse smaller Mullerian bodies transferred
from otherspecies of Cecropia(D. W. Davidson & P.
Hererra,unpubl. data). Although composed of gly-
cogen, as in congeneric species (F. Rickson, pers.
comm.), the Muillerianbodies of "C. pungara"must
be distinctivein some otherway; based on superficial
examination, they are unusually hard and less apt
to degrade during storage. The Miillerian bodies of
C. hispidissima appearto have evolved with Pachy-
condyla prov. dianae, a species even larger-bodied
than P luteola, and are larger and hardereven than
those of "Cecropiapungara."OtherCecropia,for ex-
ample "C. grisita" (a name used for materialnot yet
matchedwith any of the recognizedCecropiaspecies)
in southeasternPeru,have especially small Miillerian
bodies (Folgarait& Davidson, 1994, 1995) or bodies
to which obligate ants from different habitatsreact
abnormally(Davidson & Fisher, 1991, for an Azteca
ovaticeps colony placed on Cecropia ficifolia).
Among the six common species co-occurring in
southeasternPeru, "C. grisita" is the only one that
neverhouses large-bodiedants,e.g., Camponotusbal-
zani (Davidson et al., 1991), perhapsbecause of the
unusually narrow stems at the time when seedlings
are being colonized.
Other ant-relatedtraits of Cecropia might have
originated either through coadaptationwith associ-
FLORANEOTROPICA
ated ants or as preadaptations,arising throughalter-
native selection pressures. For example, among the
Cecropia species present in southeasternPeru, C.
membranaceaand "C.pungara"are distinctivein in-
itiatingMullerianbody productionpriorto the broad-
ening of stems and the production of prostomata.
Since the trait occurs in C. membranacea,on which
Pachycondyla luteola queens fail to establish colo-
nies, it is possible thatit evolved in a differentcontext
and merely preadapted"Cecropiapungara" for use
by this nonclaustral ant species (Yu & Davidson,
1997). The long, relatively sparse stem hairs of "C.
pungara" could either be an adaptationto facilitate
movementsof its relatively large ant species (Pachy-
condyla luteola and Camponotusbalzani) or have
provideda preadaptednest site where urticatinghairs
deterred potential predatorsof ant brood. Rates of
productionof Muillerianand pearl bodies, or leaves
bearing both rewards, could either have coevolved
with or been preadaptedto colony growth rates of
particularant species. Pearl body productionmay be
low in Cecropia membranacea(Folgarait& David-
son, 1995) becauseits most frequentassociate,Azteca
ovaticeps, relies only weakly or not at all on these
food rewards(Baird, 1967), or because of low nitro-
gen levels in the frequentlyinundatedhabitatsof this
species. Glaucous stems, such as those that restrict
climbing by all but the most obligate ant associates
of some myrmecophyticMacaranga (Federle et al.,
1997), occur as a developmentalstage (D. W. David-
son, unpubl.data)or polymorphismin some Cecropia
species. While this trait might have evolved to limit
access to all but beneficial ants, it could also have
originatedas a defense againstinsect herbivores.(Re-
cent evidence [D. W. Davidson et al., unpubl. data]
suggests the latterfor Cecropiavs. strigosa.) Finally,
selection to enhance the integrity of the intemodal
septa inside Cecropia stems (thicker and harderin
myrmecophytesthanin nonmyrmecophytes:see Bai-
ley, 1922) could occur either for structuralreasonsor
to allow for competitionamong multiplefoundresses
and theirincipientcolonies. By assuringthatmultiple
colonies of one or more species can develop in iso-
lation from one another, the plant should both in-
crease its chances for successful establishmentof at
least one colony, andhelp to magnifyselection among
competitorsfor rapidcolony developmentrates.
Among ants, coadapted traits resulting from as-
sociation with one or more Cecropia species would
include the recognition and use of both prostomata
and Muillerianbodies by queens and workers.In both
the introduced(Wetterer,1997) and native ranges of
Cecropia(Davidson& Fisher, 1991), ants thatarenot
CECROPIAAND ITS BIOTICDEFENSES
obligate Cecropia associates tend not to recognize
Miillerian bodies as food. Those in the introduced
rangeshave also not been observedto use prostomata
to nest in Cecropia stems (Putz & Holbrook, 1988;
Wetterer,1997). In contrast, queens of obligate as-
sociates in the Azteca alfari group appear to have
evolved head shapes thatfacilitatestem entry at pros-
tomata (Longino, 1989b). Facing high risk of preda-
tion during colony founding, queens of obligate
Cecropia-antsshouldalso have evolved to locate their
hosts quickly from a distance, by respondingto pos-
sible chemical cues elaboratedby one or more hosts.
At closer range, queen transfer experiments have
shown that both Azteca australis and Pachycondyla
luteola exhibit strongpreferencesfor host species on
which broodproductionis most successful (Yu& Da-
vidson, 1997, and above). The latterspecies colonizes
just "Cecropiapungara,"the only species on which
this ponerine ant succeeds in establishing colonies,
and C. membranacea,believed to be a very close rel-
ative (Yu & Davidson, 1997).
Finally, many Cecropia-ants attack and prune
vines and other vegetation that contacts their host
plants (Janzen,1969; Davidson et al., 1988). This be-
havior, present in the associates of many myrmeco-
phytes, reducesaccess to the plantsby competingand
predatory ants (Davidson et al., 1988). Occurring
among Cecropia-antsin (at least) a numberof Azteca
species (e.g., Janzen, 1969) and in Pachycondylalu-
teola (Davidson et al., 1988), the behaviormay be an
aspect of territorialitythat evolved either before the
relationshipwith Cecropia, or as longer colony life-
times became possible in the long-lived andrelatively
protective nesting environments provided by myr-
mecophytes.
Despite abundantevidence for coadaptation,there
is little supportfor cospeciation or cocladogenesis in
either Cecropia or its ants. Thus, althoughparticular
associates may be highly species-specific (e.g.,
Pachycondylaluteola on "Cecropiapungara,"Pachy-
condyla sp. nov. 1 on Cecropia hispidissima, and
Crematogasteraff. curvispina on "Cecropiaherrer-
ensis"), no substantialradiationof partnerlineages
appears to have occurred through cocladogenesis.
Nevertheless,cospeciationcannotbe firmlyruledout,
and evidence might yet be found as relationshipsof
Azteca and Cecropia come undergreaterscrutiny.
EVOLUTIONARYCOLONIZATION,HOST
SHIFTS, AND HABITAT SHIFTS
De novo or evolutionarycolonization is the alter-
native to cocladogenesis in magnifying the diversity
of Cecropia and its ants over evolutionarytime. As
225
described above, the transitionfrom free-living ants
to symbiotic associates of Cecropiahas occurredfre-
quently and apparentlyeasily, and accountsfor much
of the diversity in these symbionts (Table 1). More-
over, after these species were evolutionarilycommit-
ted to Cecropia, the combinationof habitatshifts in
plants and host switches in ants would have further
enhanced the diversity of ant-plant partnerships.In
turn, the facility with which Cecropia spp. acquired
new ant partnersmay have permittedthe genus to
attainhigh species diversity throughfrequentevolu-
tionary habitat shifts, often correlatedwith changes
in growth rates and defensive investment(Davidson
& Fisher, 1991; Folgarait& Davidson, 1994, 1995).
Although many lowland rain forest Cecropiaspecies
requiresome sort of light gap for establishment,spe-
cies differ widely in their light requirements,as well
as characteristicsoil types. As exemplifiedby species
encountered in southeasternPeru, some are tightly
bound to riparianedge (e.g., C. latiloba and C. en-
gleriana), others to frequently inundated soils (C.
membranacea,"C. pungara,"and C. utcubambana),
some to less fertile terra firme (C. ficifolia, C. poly-
stachya, and "C. grisita"), and still others to high-
elevation cloud forests along the easternAndeancor-
dillera (C. angustifolia and C. tacuna). In their
comparison of ant-plant relationshipsin Africa and
the Neotropics, McKey & Davidson (1993) argued
that a fine-scale habitat mosaic, created by Andean
orogeny and rivers meanderingwidely and at differ-
ent elevations over geologic time (Salo et al., 1986),
has been profoundlyimportantin magnifying diver-
sity within a numberof ant-planttaxa, including Ce-
cropia, and in the partnershipsof habitat-shifting
plant species with novel ant taxa. Thus, comparedto
Africa, where geomorphologyis more monotonous,
the Neotropicshave 3.5-fold greaterspecies richness
of plant-ants,despitejust 1.3 times as many total ant
species.
Yu & Davidson (1997) have attributedto the in-
dependent (horizontal) dispersal of symbionts both
the predominanceof de novo colonization over co-
speciation in the Cecropia-antsystem and the diver-
sity of mechanismsassociatedwith evolutionarycol-
onization and species-specificity. Thus, rather than
being codispersedwith theirpartners,as is the case in
many symbiotic associations between mutualists,or
between parasitesandhosts, colonizationof Cecropia
seedlings and saplings occurs anew each generation,
allowing much opportunityfor evolutionarycoloni-
zation and host shifts that may, in turn, inhibit pair-
wise coevolution and cocladogenesis. In this circum-
stance, historicalcoincidences such as coordinatedor
exclusionary dispersal (to the same or differenthab-
226
itats, respectively) may be as importantto, or more
importantthan,competitionfor partnersin settingthe
stage for future coevolution. Moreover,pairings be-
tween Cecropia and its obligate associates are ex-
plainedlargelyby the effects of local propagulepools,
queen preferencesandhost locating abilities,andcol-
ony performanceson young seedlings and saplings,
i.e., by events early in the life histories of these as-
sociations (Yu & Davidson, 1997).
FLORA NEOTROPICA
Almost certainly, given the species richness of
Cecropia and the widespread distribution of this
importantneotropical genus, additional ant associ-
ates remain to be discovered. Analysis of these new
species, together with continued progress in recon-
structingthe phylogenies of both Azteca and Cecro-
pia with molecularand other characterswill provide
new and informative tests of the ideas developed
here.
INDEX OF SCIENTIFICNAMES 227

INDEX OF SCIENTIFICNAMES
Synonyms are in italics. Names in [brackets]are nomina nuda or doubtfulnames. Names between "quo-
tation marks"are names for Cecropiamaterialwith uncertainor unknownidentityand used in studieson ants.
Page numberswith an asterisk (*) indicatepages with illustrationor maps.

PLANTS boliviana, 151 hondurensis,144

bracteata, 159 hormigana,78
Ambaiba,32 bullata,53*, 64, 65* humboldtiana,142
adenopus, 135 bureauiana,138 ibaguensis, 124
carbonaria, 135 burriada, 130 idroboi, 51*, 99
cinerea, 135 candida, 98 inchuensis, 168
concolor, 68 carbonaria, 135 insignis, 51*, 100, 101*
costaricensis, 128 catarinensis, 135 [javitensis], 188
cyrtostachya,135 caucana, 54 juraniyana, 168
hemsleyana, 128 chlorostachya,48*, 66, 67* kavanayensis,51*, 103
hololeuca, 98 cinerea, 135 klotzschiana,151
latiloba, 104 cobariana, 161 laetevirens, 116
leucocoma, 68 commutata,128 lanciloba, 121
lyratiloba, 135 concolor, 68, 69*, 70* latiloba,48*, 104, 105*, 225
membranacea,116 subsp. englesiana, 76 leucocoma, 68
mexicana, 128 congesta, 90 leucophaea, 151
obtusa, 127 coriacea, 54 levyana, 128
pachystachya, 135 cyrtostachya,135 libradensis, 161
palmata, 138 dabeibana, 130 lisboana, 127
peltata, 142 danielis, 54 litoralis,48*, 107, 108*
sciadophylla, 168 [dentata],188 longipes, 48*, 110
surinamensis, 142 lyratiloba, 135
dielsiana, 144
Cecropia, 32 digitata Klotzsch, 54 var.nana, 135
acutifolia, 34 digitata Miquel, 135 macranthera,86
adenopus, 135 var.grisea, 142 magnifolia, 78
var. lata, 135 diguensis, 150 maranhensis,68
var. lyratiloba, 135 forma albicans, 82 marginalis,48*, 110, 112*
var. macrophylla,135 discolor, 78 maxima, 51*, 113, 114
var. oblonga, 135 distachya,51*, 72, 73*, 216 maxonii, 130
var. vulgaris, 135 elongata, 70*, 75 megastachya,70*, 115
albicans,47, 48* engleriana,51*, 69*, 76, 225 membranacea,48*, 116, 117*,
alborugosa, 177 eximia, 100 219, 224, 225
alexandrina, 121 ferreyrae, 78 metensis, 70*, 120
alvarezii, 130 ficifolia, 51*, 78, 79*, 224, 225 mexicana, 128
[ambaci], 188 flagellifera, 151 var.macrostachya,128
amphichlora,130 [floccosa], 188 mituana,78
andina,49, 50*, 51* francisci, 151 mocoana, 78
angelica, 180 gabrielis, 70*, 82, 83* moniquirana,54
angulata,52, 53* garciae, 48*, 84, 85* monostachya,82
angustifolia,51*, 54, 55*, 56*, [glauca], 188 montana,48*, 121, 122*
57* glaziovii, 53*, 86 multiflora,173
annulata,51*, 63 goodspeedii, 144 multisecta,53*, 124, 125*
antillarum, 167 goudotiana,53*, 88, 89* mutisiana,48*, 124
arachnoidea, 144 granvilleana,48*, 90, 91* [nigra], 188
[arenaria],188 "grisita",224, 225 nivea, 151
arbelaezii, 88 [guanipensis],188 obovata, 68
[argentea],188 hachensis, 54 obtusa, 48*, 127, 215
asperrima, 144 "herrerensis",120, 219, 220, 226 obtusifolia,48*, 128, 129*, 215
auyantepuiana,103 herthae,51*, 90, 92* subsp. burriada, 130
[bella], 188 heterochroma,48*, 93, 94* occidentalis, 116
bicolor, 173 hispidissima,51*, 95, 96, 216 orinocensis, 105
bifurcata, 116 hololeuca, 48*, 98 pachystachya,70*, 135
228 INDEX OF SCIENTIFICNAMES

pacis, 76 silvae, 53*, 172 ANIMALS

palmata,70*, 138 [solanoensis], 188
palmatisecta, 54 standleyi, 78 Azteca, 216, 220
[paludosa], 188 [stenostachya],188 alfari, 221-223
panamensis, 128 steyermarkii,103 aragua,221, 222
paraensis, 105 strigilosa, 54 australis,221-223
pastasana,70*, 140, 141* strigosa, 70*, 173, 224 coeruleipennis,221
peltata, 70*, 142, 143*, 217 subintegra,70*, 174 constructor,215, 221, 222
var. candida, 120 sucrensis, 140 isthmica, 221
philipsonii, 54 surinamensis,142 laevigata,216
pinnatiloba, 151 sylvicola, 54 lattke, 221
pittieri,48*, 150 tacuna,48* merida, 221, 222
plicata, 53*, 150 telealba, 51*, 177 muelleri, 221-223
polyandrophora,100 var.hirsuta, 177 ovaticeps, 221-223
polyphlebia, 54 telealbida, 178 petalocephala,221, 222
polystachya,70*, 151, 152*, 225 teleargentea, 177 salti, 221, 222
porvenirensis,78 teleincana, 180 xanthocroa,221, 222
propinqua, 144 telenitida,70*, 178, 179*
puberula, 182 telenivea, 178 Camptonotus,220, 224
"pungara",120, 219, 220, 225 tessmannii, 116 balzani, 221-224
purpurascens,51*, 155 tolimensis, 124 Chrysomelidae,215, 216
putumayonis,70*, 159, 160* trilobata, 155 Coccidae, 215
radlkoferiana,128 tubulosa,54 Coelomera,216
reticulata,70*, 159, 160* ulei, 53*, 182, 216, 225 atrocaerulea,215
var. alboreticulata, 159 urbaniana, 166 Coleoptera,216
richardii,72 utcubambana,53*, 182, 183* Crematogaster,222, 223
riparia, 72 vageleri, 116 curvispinosa,220, 221, 222
robusta, 116 velutinella,70*, 185, 186* Curculionidae,216
rugosa, 173 villosa, 54
ruiziana, 151 virgusa,48*, 187 Dolichoderinae,220
sandersoniana, 100 yarinensis, 76
santanderensis,180 Cecropiaceae,28 Epitragus,22
sararensis,70*, 161
Cecropioideae,28
saxatilis, 48*, 163 Formicidae,220
Coilotapalus,32
scabra Klotzsch, 151 Formicinae,220, 222
Conocephaleae,28
scabra Martius,188
Conocephaloideae,28
scabrifolia, 144 Myrmecinae,220, 222
Conocephalus,28
schiedeana, 142 Myrmelachista,222
schreberiana,164 Erythroxylum,31
subsp. antillarum,70*, 165*, Ophtalmoborus,22
167 Macaranga,6, 25, 216, 224
subsp. schreberiana,70*, Musanga,6, 25, 28, 215 Pachycondyla,216, 220, 223, 224
165*, 166 cecropioides, 149 bugabensis,221
sciadophylla,53*, 168, 219 Myrianthus,28 prov. dianae, 221
var.decurrens, 168 Ainsignis, 221
var.guamuesensis, 168 Piper, 217 luteola, 220-222, 224
var.juraniyana, 168 Poikilospermum,28 unidentatarugulosa,221
var.pedroa, 168 Pourouma,28 villosa, 221
var.subsessilis, 168 formicarum,8 Pheidole, 217, 221, 222
scutata, 159 guianensis subsp. guianensis, 36, Ponerinae,220
sericea, 167 188 Pseudococcidae,215
setico, 116 myrmecophila,8 Pseudomyrmecinae,220
INDEX OF VERNACULARNAMES 229

INDEX OF VERNACULARNAMES
In additionto seeing the taxa cited below for vernacularnames, also see the section "VernacularNames."
ama'i (36) guarumbo (p. 31)
ama'i-hete (36) guarumo (p. 31)
ama'-y-ata (51) guarumo blanco (51, 60)
ama'-y-puku (8, 39, 51) guarumo moreno (4)
ama-'y-tuwir (8, 36) guarumo negro (4, 18)
amba'i (38) gueremo (41)
amba'y (38)
hormigo (37)
ambahu (38)
hormiguillo (37)
ambaf (38)
huagadeug (37)
ambaiba (p. 31)
huarumo (11, 12, 51)
ambai'ba(p. 31)
ambai'ba-tinga(38) ibaiba (38)
ambaibo (p. 31) imbauba (p. 31)
ambaibo blanco (11, 31, 44, 51) imbauba branca (8, 12, 21, 25, 36, 38, 44, 49)
ambaibo colorado (51) imbautbada varzea (25)
ambaibo negro (5, 8) imbautbagigante (51)
ambati (38) imbautbaroxa (45, 58)
ambay (38) imbau'batorem (I 1)
anduchi dundo (12) imbauba vermelha (9, 21, 38, 39, 51)
arduchina durdu (12) imbaubao (38)
arumo cocedera (37) imbaubarana(25)
arvore da preguica (38) imbaubu,cu(21)
imuisai (51)
bois canon (p. 31)
bois canot (50a) jagrumo (4)
bois trompette (50a). jaibena (12)
bokarawe (31) jucoqui (9)
bospapaja (25) juimekokai (9)
burriada (37) kaa-bona (31)
burriala (37) kama-in-yek (3)
calentano (35) kama-in-yek (9)
camai-yin-yek (24) kiralkai fikai (12)
cetico (p. 31) kooch le' (41)
cetico blanco (12) kulegle (36)
cetico colorado (51) liarumo (61)
chancarro (37, 41) liarumo blanco (4)
chancarro (37) Ilarumo negro (4)
chanchan (2)
maguimehue (51)
changarro (37)
makamakalu (5 1)
cho-otz (41)
manbospapaja (51)
congo pump (3, 36, 41, 51)
mangineowe (51)
congopom (3)
cosedera (14, 47) nig-la (41)
cosedera teu (61)
orumo (4, 41)
diapapaie (51)
pata de gallina (4)
dondo (31)
pau de lixa (38)
dondo (11)
pau de formiga (38)
dundu (51)
purma (4)
dundu (12)
sakaka (51)
emba6ba (p. 31)
sarasara-yek (5 1)
embauba branca (11, 21, 38)
setico (p. 31)
embaubacu'(21)
setico blanco (12, 31)
guanibarae (31) setico colorado (25, 51)
guargiiero (41) setico de altura (51)
230 INDEX OF VERNACULARNAMES

setico negro (9) tselan dundu (28)

setico roja (51) tsenguillachi (37)
shiari (9, 53) tsitica dundu (31)
siari chal (11) tsu (28)
simbrapotro (25) tud teu (61)
snake wood (p. 31)
uaru (25)
su (51, 60)
udagomo (12, 31)
sui (28)
umbadba (p. 31)
suti (31)
sutiik (12, 31) virgusa (61)
suti'k(31)
suu (11, 28, 51) wanasoro (25, 51)
suu (12, 31) waru (3)

taa-bona (31) xkoochle (41)

tacona (p. 000) yagrumo (p. 31)
tacona blanco (53) yagrumo blanco (12)
tacona ojo de venado (51) yagrumo montaniero(51)
tacuma (55) yagrumo morado (9, 51)
tacuna (p. 31) yagrumo rojo (31)
toc'kori (12) yanat (51)
tokori (25) yantan (51)
tokorodek (25) yarumo (31)
torem do igap6 (51) yarumo blanco (35, 56)
torem (38, 39) yarumo corante (31)
torem (38) yarumo de raton (12)
toroc (9, 44) yarumo macho (31)
torog (51) yarumo negro (9)
toronoma (51)
yawa-cl'ki (I 1)
tourem da folha grande (51)
yawatseke (11)
trampy (41)
yawg tseke (46)
trompeta (37)
yemeoba (51)
trumpet tree (p. 31)
yuarumbo (14)
trumpet (37, 41, 50a)
yungapara (4)
trumpeter tree (50a)
yungul (44)
tsake kumpari (28)
yura tsitica dundu (46)
tsake (12)
tseke ( 1, 46) zmuikaz (51)