Organization for Flora Neotropica

Endlicheria (Lauraceae)
Author(s): André S. Chanderbali
Source: Flora Neotropica, Vol. 91, Endlicheria (Lauraceae) (Aug. 24, 2004), pp. 1-141
Published by: New York Botanical Garden Press on behalf of Organization for Flora Neotropica
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 INTRODUCTION
Withinthis book, Endlicheriaand Rhodostemonodaphneare treatedas separatemono-
graphs.Traditionally,Endlicheriawas placednearAnibaandothergeneraof the Lauraceae
with two-locellate anthers.Rhodostemonodaphne, with four-locellateanthers,was recently
segregated from Nectandra. New information,however, suggests that Endlicheria and
Rhodostemonodaphneform a monophyleticgroup of approximately100 species. Never-
theless, the phylogeneticrelationshipsamong these species are still unclearanduntil there
is more conclusive evidence we feel this complex group of species should remainwithin
two genera.
ANDRt S. CHANDERBALI AND

SANTIAGO MADRINAN
FLORANEOTROPICAMONOGRAPH91

ENDLICHERIA (LAURACEAE)
ANDRES. CHANDERBALI

; P _ PCANCER

FLORAL
NEOTROPICA

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 ENDLICHERIA(LAURACEAE)
ANDRE S. CHANDERBALI

CONTENTS
Abstract/Resumen ................................2
Introduction...............................2
Historical Survey................................2
Morphologyand Anatomy ....3...........................3
VegetativeMorphology....3...........................3
Inflorescences...4............................4
Flowers ...............................4
Fruits ...............................5
Wood and Bark Anatomy ...............................5
Floral Anatomy ...5............................5
Embryology...............................6
Karyology................................7
Palynology................................7
Relationships...7............................7
SystematicPosition ....7...........................7
Generic Delimitationand Species Groups ...............................7
Species Concepts............................... 14
Distributionand Biogeography............................... 14
SystematicTreatment............................... 16
Generic Description............................... 16
Key to the Species............................... 16
Species Descriptions ............................... 22
Doubtful Names and ExcludedTaxa ............................... 123
Acknowledgments............................... 125
LiteratureCited............................... 125
NumericalList of Taxa ............................... 126
List of Exsiccatae ............................... 127
Index of Local Names ............................... 137
Index of Scientific Names ............................... 138
2 FLORA NEOTROPICA

ABSTRACT
Chanderbali,A. S. (Departmentof Botany, University of Florida, Gainesville, FL
3261 1, USA). Endlicheria(Lauraceae).FloraNeotropicaMonograph91. 1-141. 2004.-A
taxonomic monographof Endlicheria, a South America-centered genus of the Ocotea
complex, is presented;and with the benefit of moleculardata showing that its members
are nested within Rhodostemonodaphneand Ocotea, its delimitationis assessed in mor-
phological detail. Sixty species of Endlicheria are recognized, of which 16 are newly
described.Two species are transferredto Rhodostemonodaphne.Nine infragenericspecies
groups are informally recognized in Endlicheria. Of these, the Endlicheriapunctulata
species groupis closer to species of the Ocotea cernua species groupthanto its congeners,
in both molecular and morphological aspects. Representativesof each of the eight re-
maining species groups are united with a diverse representationof Rhodostemonodaphne
in a well-supportedbut unresolvedclade. The significanceof this Endlicheria-Rhodoste-
monodaphnealliance is explored from a morphologicalperspective.It is suggested that
the two-locellate anthersthat distinguishEndlicheriafrom Rhodostemonodaphne evolved
repeatedly.

RESUMEN
Chanderbali,A. S. (Departmentof Botany, University of Florida, Gainesville, FL
32611, USA). Endlicheria(Lauraceae).FloraNeotropicaMonograph91. 1-141. 2004.-
Se presentauna monograffataxon6micade Endlicheria,genero pertenecienteal complejo
de Ocotea con centro de diversidaden America del Sur; y con la ventajade datos mole-
culares monstrandoque sus miembrosestainincluidos dentrode Rhodostemonodaphney
Ocotea, su delimitaci6ngenerica es evaluadamorfologicamenteal detalle. Se reconocen
60 especies en Endlicheria,de las cuales 16 se describencomo nuevas. Dos especies son
transferidasa Rhodostemonodaphne.Nueve grupos infragenericosde especies son infor-
malmentereconocidosen Endlicheria.De estos, el grupode especies de Endlicheriapunc-
tulata esta mas cercanamenteemparentadaal grupo de especies de Ocotea cernua que a
las especies congenericas, tanto en los aspectos molecularescomo en los morfologicos.
Los representantesde los ocho gruposde especies restantes,junto con una representaci6n
diversa de Rhodostemonodaphneformanun clado con buen soportepero sin resolucion.
La importanciade la alianza Endlicheria-Rhodostemonodaphne es evaluadadesde una
perspectivamorfol6gica. Es sugerido que las anterascon dos 16culosque distinguenEn-
dlicheria de Rhodostemonodaphnepodrianhaberevolucionadorepetidamente.

INTRODUCTION daphne than to Ocotea (Chanderbaliet al., 2001).

Here I updatethe taxonomy of Endlicheriain mon-
The Lauraceaeform a large pantropicalfamily of
ographic custom and assess its circumscriptionin
about 50 genera and approximately2500-3000 spe-
morphologicaldetail while I present additionalmo-
cies of trees, rarelyshrubs,and one genus of parasitic
leculardata relevantto its systematicposition.
vines, Cassytha L. In Endlicheria Nees all species
This treatmentis based on almost 2500 collec-
with non-involucrate inflorescences, a dioecious
tions. In additionto the substantialmaterialavailable
breedingsystem, andtwo-locellateanthersareplaced.
at MO, many of recent collections with several du-
These characters circumscribe a neotropical taxon
plicates not yet distributedto other herbaria,loans
that differs from sympatric Ocotea Aubl. pro parte
were received from A, AAU, B, C, COL, E, ECON,
and Rhodostemonodaphne Rohwer & Kubitzki
F, G, GB, GH, GOET,HBG, HUA, INPA, K, L, LE,
mainly in having two ratherthan four-locellate an-
MG, NY, P, QAME, R, S, U, UC, US, and VEN.
thers, a distinction of questionablegeneric value in
Lauraceae(e.g., van der Werff, 1984; Burger, 1988;
Rohwer et al., 1991). A recent molecular study of HISTORICAL SURVEY
Lauraceae showed that Endlicheria is at least di- Nees (1833) describedEndlicheriawith two spe-
phyletic, with more species allied to Rhodostemono- cies. CryptocaryahirsutaSchott(= Endlicheriapan-
MORPHOLOGYAND ANATOMY
iculata) was transferredto Endlicheria,andE. sericea
was newly described.In 1836, Nees addedthreemore
species andchangedthe generic epithetto Goeppertia
Nees because of the earlier Endlichera Presl in Ru-
biaceae. Also in 1836. Endlicheria appearedin the
list of Lauraceaegenerapublishedin the second edi-
tion of JohnLindley'sNaturalSystemof Botany.Nees
supplied this list to Lindley in 1835 and, already
awareof Presl's Endlichera,suggested thatSchauera
Nees be used instead of Endlicheria if Presl's name
was to be retainedfor the rubiaceousgenus. Koster-
mans (1936) pointedout unsatisfactoryconsequences
with use of either Goeppertiaor Schauera in Laura-
ceae, and his proposalthat Endlicheriabe conserved
in Lauraceaeover Endlichera in Rubiaceae was ac-
cepted at the VI BotanicalCongress.The generic ep-
ithet honors Stephan Ladislaus Endlicher (1804-
1849).
Meissner (1864) treatedEndlicheria(as Goepper-
tia) in his treatmentof Lauraceaefor De Candolle's
Prodromus.Meissner accepted 14 species in Goep-
pertia, four only tentatively,and transferredG. mul-
tiflora Miq. to AmpelodaphneMeisn., a newly de-
scribed genus with two other species. Mez (1889), in
his treatmentof American Lauraceae,excluded the
four species Meissner tentativelydescribedin Goep-
pertia and included Ampelodaphnein Endlicheria.
Mez apparentlyused the name Endlicheriainsteadof
Goeppertia because he believed Presl's Endlichera
was publishedwithout description,i.e., a nomen nu-
dum.Mez recognized23 species and for the firsttime
provided an infrageneric scheme. Five subgenera
were recognized. Subgenera Hemiajouea Mez, An-
osphaeria Mez, and Ocoteopsis Mez, were mono-
typic, accommodatingE. paradoxa, E. impressa,and
E. anomala, respectively. Subgenus Ampelodaphne
(Meisn.) Mez accommodatedeight species and su-
bgen. EuendlicheriaMez the remaining 12 species.
Kostermans(1937) followed Mez's concept of Endli-
cheria in the only subsequent monograph. He in-
cluded monotypic HuberodaphneDucke in Endli-
cheria and recognized 39 species. In his generic
subdivision Kostermans (1937) maintained Mez's
monotypic subgen. Ocoteopsis and placed the re-
maining species in subgen. EuendlicheriaMez. The
latter was divided into sections Microlocellata Kos-
term. and Macrolocellata Kosterm. on the basis of
size and position of locelli in the outer staminal
whorls.
Since Kostermans's treatment, several authors
have describednew species in Endlicheria.Most no-
tably, C. K. Allen, in a series of paperspublishedin
The New YorkBotanicalGarden'sBotany of the Gui-
3
anaHighlandsseries, addednine species. Priorto this
treatment68 names were validly publishedin Endli-
cheria.
MORPHOLOGY AND ANATOMY
VEGETATIVEMORPHOLOGY
Species of Endlicheria are mostly medium-sized
trees less than 25 meters in height, but they can reach
up to 40 meters; a few are shrubs. Of the shrubby
species, at least E. reflectens and E. vinotincta are
consistentlyreportedto be scandentwith long slender
stems that sprawl across adjacentvegetation.Growth
is apparentlyrhythmic with leaves either produced
along the entire length of flushes or, as typical of the
Ampelodaphnespecies group, clusteredin close spi-
rals at the tips of each new flush.
Leaves are always simple, entire, and petiolate.
Only in Endlicheriadictifarinosa,E. bracteata,E. co-
cuirey,E. melinonii,andE. verticillataareleaves sub-
sessile. Shapevaries fromovateto obovateandis usu-
ally constant within species, but in a few (e.g., E.
ruforamula and E. paniculata) these extremes are
linked by intermediates.Laminae are usually plane,
but are stronglybullatein E. bullata and occasionally
moderatelyso in E. glomerata.Leaf apices areusually
acuminate,but are caudatein E. longicaudataand E.
punctulata,andbluntlyroundedto retusein E. rubra.
Leaf bases are most often acute, but vary from atten-
uate to abruptlycontractedinto the petiole (i.e., sub-
cordate).Marginsareusually curveddownwardat ca.
900 (recurved),at least in the lower half of the lamina,
but they can be plane or recurvedthroughout.
Venationis mostly pinnatewith secondariesalter-
nately arrangedand evenly spaced along the midrib.
Only E. acuminata,E. bracteolata,E. gracilis, andE.
krukoviihave triplinervedleaves, where one pair of
(sub-)oppositesecondariesemerges shortlyabove the
base of the midrib, and others (if present) emerge
from the midrib aroundor beyond midlamina.In E.
bracteolataand E. sericea quintuplinervedleaves ap-
pear when two pairs of (sub-)oppositebasal second-
aries emerge in the lower lamina. Secondariesrange
in numberfrom 2 in triplinervedspecies to over 30
per side in E. bullata,but usually number5 to 7 pairs.
In most species, the secondaries follow arcuate
courses and distal pairs form weak loop connections
near the leaf apex. Secondariesthat form strongloop
connections throughoutthe lamina (i.e., brochidod-
romous secondaries) are restrictedto E. arunciflora,
E. bullata, E. dysodantha,and E. longicaudata. Oc-
casionally, basalmost secondaries are asymmetric,
with one or both merging with the margin near the
leaf base or emergingfrom the junctionof midriband
4 FLORANEOTROPICA
leaf base. This venation patternis typical of E. me-
tallica, E. chrysovelutina,and E. klugii, and appears
occasionally in E. canescens and E. ruforamula.Ter-
tiaries are typically roughly horizontal and parallel,
proceeding undivided or once-forked between adja-
cent secondariesor between midrib and secondaries,
but rarely, e.g., in E. reflectens, they reticulate
throughoutthe intercosta.Midribs, secondaries, and
tertiariesare usually raised on both sides of the lam-
ina, at least in dried specimens, but one, two, or all
vein ordersmay be immersedabove. In E. punctulata
all are immersedbelow.
Variationin color, density, orientation,size, and
shape of hairsprovidesa great arrayof charactersfor
species identification.Apart from Endlicheriapyri-
formis, the vegetativesurfacesof which would be gla-
brous were it not for a marginalfringe of hairson the
youngest leaves (in bud), species have conspicuous,
and often characteristic,indumentum.Hairs can be
straight,crooked,crisped,or tightly crinkled.The lat-
ter type of hairis almostrestrictedto the innersurface
of tepals and rarely achieves the density typical of
Nectandra Rol. ex Rottb. where, as here, they are
termed papillose (Rohwer, 1993a). All other indu-
ment termsare used as definedby Steam (1992). Hair
orientation,whethererect or closely appressedto the
surface, appearsto be stable within species, but both
extremes are found in the circumscriptionof E. acu-
minata, E. anomala, and E. paniculata adoptedhere.
Usually only one type of hairis producedon a surface,
but in E. duotincta and E. williamsii a dense tomen-
tose cover that obscures the lower leaf surface is in-
terspersedwith longer erect hairs, and in E. dysodan-
tha andE. tomentosadense tuftsof longermore stiffly
erect hairs than found elsewhere on the lower leaf
surfaceoccupy the axils of secondaryveins. In the E.
sericea and E. metallica species groups, a dense
golden or silvery sericeous indument persists from
buds to maturity.
INFLORESCENCES
Inflorescencesof Endlicheriaare usually panicles
with terminaldichasia, i.e., thyrso-paniculatein the
sense of Weberling(1989). However,in E. citriodora,
E. acuminata,and E. xerampelaterminalcymes col-
lapse into pseudo-umbellatefascicles; and in several,
terminalbotryoids(sensu Weberling,1989), or deter-
minate racemes, are formed via reductionof subter-
minal lateralcymes to single flowers.The axes of bo-
tryoids are occasionally greatly reduced,and flowers
are groupedin dense clusters along secondaryinflo-
rescence branches(e.g., E. glomerata).Inflorescences
are either borne in the axils of new foliage leaves or,
as typical of the Ampelodaphnespecies group, in the
axils of scalelike bracts(cataphylls)thatprecedeleaf
crops. Often, the axis of the cataphyll-bearingshoot
is greatly reducedand inflorescencesare arrangedin
subterminalclusters above the latest leaf crop. Pistil-
late inflorescencesare usually shorter,less branched,
and fewer floweredthanthe staminateinflorescences,
but can also be equal in these regards.
Inflorescences typically have the indument of
twigs and other vegetative surfaces. However, in E.
arunciflora,E. arachnocome,and E. verticillata,in-
florescences (and outer surfaceof flowers) are subgl-
abrousalthoughthe plants are otherwise densely pu-
bescent. In a few species, inflorescences are more
densely pubescentthanvegetativeorgans(e.g., E.for-
mosa and E. metallica).
FLOWERS
Flowers of Endlicheriaare organizedin the man-
ner typical of Lauraceae,i.e., all are trimerousand
consist of two whorls of (sub-)equaltepals and three
whorls of stamens surroundinga simple pistil. As is
also typical of Lauraceae,stamensare arrangedin al-
temating cycles, such that those of outermost(whorl
I) and innermost whorls (whorl III) stand opposite
each other, with those of the intermediatewhorl II
placed altemately. Rarely, a whorl of staminodes
(whorl IV) is producedopposite whorl II inside the
innermoststaminalwhorl (whorl III). A pair of nec-
tariferous glands is usually attachedto the base of
each whorl III stamen.Locelli areintrorseor introrse-
latrorse in whorls I and II and extrorse-latrorsein
whorl III. With dehiscence, valves that initially cov-
ered locelli areraisedabove antherswith pollen grains
attachedto theirinnersurfaces.The pollinationof En-
dlicheria flowers has not been studied in detail, but
as elsewhere in Lauraceae(Kurz, 1982; Kubitzki&
Kurz, 1984) it is most likely that short-tonguedin-
sects, such as bees andflies, arethe pollinatingagents.
Not surprisingly,the arrangementof stamensand lo-
celli places valves of all threestaminalwhorlsin close
proximityover the nectariferousglandsthatthese vis-
iting insects seek.
Within the limits of this basic organization,the
range of floral variationin Endlicheriais such that a
typical flower structurecannotbe envisioned.Flower
shape ranges from rotatewith horizontallyspreading
tepals, through cyathiform or infundibuliformwith
erect or ascending tepals, to globose or obconical
where tepals open only enough to provide a terminal
pore (terminology after Steam, 1992). The androe-
cium is also variable.Stamensof the outerandroecial
whorls (whorls I and II) are (sub-)equal.Hereanthers
MORPHOLOGYAND ANATOMY
are two-locellate, and, in most species, the filaments
are appreciablynarrowerthan anthers;i.e., stamens
are stipitate.In several species however, whorl I and
II stamens are sessile with fleshy filaments that are
broaderthan the anthers,and in E. lorastemonfila-
ments equal anthersin width. Also, as noted by Kos-
termans(1937), two basic anthertypes occur in whorl
I and II stamens. In species he assigned to sect. Mi-
crolocellata, locelli are small and distallyplaced (Fig.
IA-), while in sect. Macrolocellata locelli occupy
almost all available space, and sterile tissue, if pres-
ent, is located mostly above ratherthan below locelli
(Fig. ID-I). These Macrolocellata-typeanthers ap-
pearin eitherof two forms. More commonly,they are
ovate with apiculateconnectivesthatemergebetween
andbeyond locelli (Fig. ID-F). Here,locelli areoften
obliquely hemispherical (as though sliced out of a
sphere along a diagonal path) with the deeper part
adjacentto the main axis of the anther(e.g., E. brown-
iana). Less frequently,Macrolocellata-typeanthers
are depressed-oblongto elliptic in shape,andconnec-
tive tissue is either reduced between, level with, or
broadabove the two locelli (Fig. 1G-I). Here, locelli
are suborbicularin shape, or in other words, hemi-
spherical with the deeper part towardthe outside of
the anther.Such anthers are typical of the Ampelo-
daphne species group, and occur also in E. anomala,
E. paniculata, and their allies. Stamens of the third
androecialwhorl (whorl III) are usually sessile with
indistinct filamentsequal to or broaderthan anthers.
Narrowlystipitatewhorl III stamensoccur only in E.
anomala, E. coriacea, E. punctulata, E. williamsii,
and the Ampelodaphne species group. Anthers of
whorl III stamensare consistentlyfour-locellatein E.
anomala, but occasionally so in E. williamsii. In both
cases, the second pairof locelli appearson the adaxial
surfaceof extrorselybowed anthers.WhorlIII anthers
are two-locellate in all other species. Nectariferous
glands of whorl III stamensare eithersessile or raised
on shortstalks (stipitate)andrangein size fromfilling
the space between stamens to inconspicuous.These
glands are lacking in E. chrysovelutina,E. longicau-
data, E. metallica, andE. mishuyacensis,but arepres-
ent at the base of all nine stamensin E. vinotincta.
Apartfrom a slightly deeperreceptaclein pistillate
flowers, sexual dimorphismis usually limited to ru-
dimentarydevelopment of male and female organs.
The gynoecium is rarelyabsent in staminateflowers,
but pistillate flowers always have staminodes, and
these may almost equal their fertile counterparts.In
such cases the tri-lobed or reniform stigma that
spreadsabove the staminodesmust be noted to deter-
mine gender (but see discussion of E. gracilis).
5
FRUITS
In most species, fruitsmatureinto a blackishdrupe
subtended by a reddish cupule. In Lauraceae this
structureis typical of most neotropicalgeneraand, as
elsewhere, the colors are classic omithochoroussig-
nals. In Endlicheria, drupes are either ellipsoid, ob-
ovoid, ovoid, or spheroidin shape, and cupules range
from deeply hemispherical with at least the lower
third of drupes included to patelliform with drupes
completely exposed. Cupulemarginsare often entire,
but in severalspecies undulateto stronglylobed mar-
gins result from a crown of persistenttepals or tepal
bases. Pedicels increase during fruit developmentto
varying degrees and range from terete and distinct
from cupules to distally swollen and graduallymerg-
ing with cupules, i.e., claviform.
Cupules are usually glabrous or at least glabres-
cent when developing from densely pubescentflow-
ers. Only the cupules of E. chalisea have a dense in-
dumentumthatcompletely covers the outersurfaceat
maturity.In this, and several other species however,
cupules are densely pubescentinside.
WOOD AND BARK ANATOMY
In terms of wood and bark anatomyEndlicheria
belongs to a large group of genera centered around
Ocotea, where it appearsto be most similarto Aiouea
and Pleurothyrium(Richter, 1981). As reportedby
Richter(1981), the wood of Endlicheriahas a specific
gravity of 0.5-0.65 g/cm3 and is diffuse-porouswith
vessel diametersranging from 100 ,um to 180 gim.
Perforationplates are simple, and intervascularpits
are 9-13 ,um diameter;fibers are libriform and sep-
tate, with simple pits; axial parenchymais paratra-
cheal, and vasicentric to vasicentric-confluent;rays
aremultiseriate,2-3 cells wide, 0.4-1.4 mm high, and
heterogeneous,but rarelyhomogeneous;and oil cells
areprevalent.Crystalsarepresentin the wood of 50%
of the species Richter examined,but the only report
of silica in the bark is based on a misidentifiedrep-
resentative, Capucho 565 (F), of Aiouea or Cinna-
momum.
FLORAL ANATOMY
Species of Endlicheria have not been examined,
but from the accounts given for Persea Mill. (Reece,
1939), Laurus L., Umbellularia(Nees) Nutt. (Kasa-
pligil, 1951), andLinderaThunb.(Boyle, 1980), there
appears to be a uniform pattem in Lauraceaethat
should also apply to Endlicheria.As summarizedby
Rohwer(1993a, 1993b):First,the six medianbundles
of the tepals branch off the stele of the floral axis
eitherat the same level or slightly differentlevels. The
6 FLORA NEOTROPICA

4~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~

-Az-A

FIG. 1. Whorl I stamens of Endlicheria. All seen from within except D which is seen from above. A. E. rubriflora
(Palacios et al. 4473). B. E. sprucei (Prance et aL 18037). C. E. metallica (Moretti 1443). D. E. browniana (Palacios
13750). E. E. canescens (Tillett & Tillett 45789). F. E. klugii (Jaramillo et aL 1159). G. E. paniculata (Heringer et al.
3062). H. E. arunciflora(Davidse 27495). I. E. anomala (Vdsquezet al. 4778).

remainingvasculartissue providestwo carpelbundles of traces in carpels, tepals, and stamensappearsto be

(the ventralone supplyingthe single apical ovule, the correlatedwith the size of these organs.
dorsal one supplying the ovary wall), style, and
stigma. Stamen traces arise from the ventral side of
the tepal bundles, and when not anastomosing,it is EMBRYOLOGY
clear thattracesto the staminalglands arise fromtheir Heo et al. (1998) recendlyimproved the under-
respective stamen traces. An increase in the number standing of the embryology in Lauraceae,but a rep-
RELATIONSHIPS
resentativeof Endlicheria is yet to be examined in
this respect. Still, the conservativenatureof embryol-
ogical charactersin the family (Heo et al., 1998) sug-
gests thatfeaturesfound in closely relatedgeneraalso
apply to Endlicheria.If so, it is expected that the an-
therwall developmentis of the basic type, the tapetum
is amoeboid; tapetal cells are 2 or 4 nucleate; cyto-
kinesis is successive; microsproretetrads are tetra-
hedralin shape and maturepollen grainsare 2-celled.
The ovule is anatropous, bitegmic, and crassinu-
cellate, the archesporiumis 1-celled; megesporetet-
rads are linear;a single embryo sac is formedvia the
Polygonum-typepattern;antipodalcells are ephem-
eral, the matureembryosac is elongateandpositioned
within the nucellus, the nucellarcap is 2-6 cell layers
thick, the ovule is pachychalazalwith ramifiedvas-
cular bundles at the chalaza; and endospermforma-
tion is of the nuclear-type.
KARYOLOGY
Chromosomecounts have not been done for En-
dlicheria, but unless it differs from most genera of
Lauraceaethat have been investigated(Okada& Ta-
naka, 1975), a base numberof x = 12, and 2n = 24
chromosomes is expected. In Lauraceae,polyploidy
is knownonly in Cassytha,Laurus,SassafrasJ. Presl.
(all with 2n = 48), and Neolitsea (Benth.) Merr.(2n
= 72). Cassytha has the largest metaphasechromo-
somes in the family (5-7 gm long) but otherwise
these range from 1 Rtmto 3 Rm in length. The cen-
tromere may be medial to subterminal,and hetero-
chromatinis found only near the ends of the arms.
PALYNOLOGY
Six species of Endlicheria-E. canescens (cited
as E. endlicheriopsis),E. glomerata,E. paniculata,E.
lhotzkyi(cited as E. sericea), E. tessmannii, and E.
verticillata-were examinedin a recentpalynological
study of neotropicalLauraceae(Raj & van derWerff,
1988). As is typical of the family, the pollen grains
in Endlicheriaareinaperturateand spheroidalwith an
extremely thin intine but ratherthick and stratified
extine. In Endlicheria,pollen size rangedfrom 15 tm
to 27 rim,the extine was 0.5-1 ,im, and scatteredwith
<.5-1 ,umlong spinules that are 0.5-2 gm apart,and
the intine was 1.5-2.5 trn. Accordingto Raj and van
derWerff(1988), such pollen is similarto thatof other
neotropicalLauraceaewith two-locellate anthers,but
among these, densely spaced processes that surround
and hide the basal cushion of spinules are found only
in Endlicheria.
7
RELATIONSHIPS
SYSTEMATICPOSITION
Relationships among Lauraceae have recently
been investigatedon the basis of wood andbarkanat-
omy (Richter,1981), inflorescencestructure(van der
Werff & Richter, 1996), embryology (Heo et al.,
1998), and molecular sequence data (Rohwer,2000;
Chanderbaliet al., 2001). These studies show that
published classifications(i.e., Nees, 1836; Meissner,
1864; Bentham, 1880; Pax, 1889; Mez, 1889; Kos-
termans,1957;Rohwer,1993b;van derWerff& Rich-
ter, 1996), do not wholly reflect phylogenetic rela-
tionships. Although differing in level of resolution
provided,these studies approacha consensusover the
main groups of genera in the family.
Cryptocaryeae(sensu van der Werff & Richter,
1996) receives supportfromall five studies.This tribe
includes all genera with cupules that enclose drupes
except for a terminalpore (e.g., AspidostemonRoh-
wer & Richterand CryptocaryaR. Br.), plus noncu-
pulateBeilschmiediaNees andits allies. Hypodaphnis
Stapf, Cassytha, CaryodaphnopsisAiry Shaw, and
NeocinnamomumH. Liu, occupy relativelybasal and
unstablepositions. A small clade of neotropicalgen-
era including Anaueria Kosterm., Chlorocardium
Rohweret al., MezilaurusKuntzeex Taub.,Sextonia
van derWerff,and Williamodendron Kubitzki& H. G.
Richt., lies sister to a large clade that resolves into a
noncupulategeneric complex centeredaroundPersea,
and a cupuliferous clade that separates into two
groups correspondingfavorablywith the tribes Lau-
reae of most systems and Kostermans's(1957) Cin-
namomeae (Chanderbaliet al., 2001). In Cinnamo-
meae, Endlicheriabelongs to a largegenericcomplex
centered aroundOcotea where it shares a dioecious
lineage with Ocotea s.str. and Rhodostemonodaphne
(Chanderbaliet al., 2001).
GENERIC DELIMITATIONAND SPECIES
GROUPS
To better assess generic delimitation,ITS rDNA
sequences of 13 species of Endlicheriaand 4 of Rho-
dostemonodaphnewere producedand analyzed with
the 94 ITS sequences of higherLauraceaepreviously
producedby Chanderbaliet al. (2001). The molecular
techniquesand data analyses are briefly summarized
here. Total DNA was obtainedfrom silica gel-dried
or herbariumleaves using DNeasy (QIAGEN)extrac-
tion kits. PCR amplificationof the ITS region was
usually conductedusing the forwardprimerLAUR 1
designedby Chanderbaliet al. (2001) andITS B from
8 FLORANEOTROPICA
Blattner(1999), but for some poor quality templates
it was necessary to amplify the region in sections
(ITS1 & ITS2) by combiningthe appropriateuniver-
sal primersof White et al. (1990) with LAUR 1 and
ITS B. The ITS/5.8S sequences producedinclude all
but the first ca. 10 bp of ITS 1 and include the entire
ITS 2 and 5.8S regions. PCR productswere purified
following the protocol providedby QlAquick gel ex-
tractionkits (QIAGEN)and sequencedusing the dye
terminatorcycle sequencing protocol (Applied Bio-
systems). Sequence reactions were analyzed on an
ABI 377 automated sequencer (University of Mis-
souri-St. Louis, D. E. Lee andFamilySequencingFa-
cility). Both strandsof DNA were readandconsensus
sequences generatedusing Sequencherver. 3.1 (Gene
Codes Corp., 1998). Sequences were manually
aligned using the sequence editing facilities of Seq-
pup version 0.6 (D. Gilbert,IndianaUniversity,Bloo-
mington, 1996).
Initial Neighbor Joining (NJ) analyses of the 111
taxa ITS dataset placed all new EndlicheriaandRho-
dostemonodaphnesequences with previously sam-
pled congenersin the dioecious lineage of the Ocotea
complex identifiedby Chanderbaliet al. (2001). The
data set was thereforereducedto these dioecious taxa
and two outgroup lineages for parsimony analyses.
Table I provides Genbankinformationfor these ac-
cessions. Heuristic searches for most parsimonious
trees were conductedwith 10 randomtaxon additions
and TBR branch swapping using PAUP* ver. 4.0b4
(Swofford, 1998). Both the MULPARS and COL-
LAPSE options were in effect, but the STEEPEST
DESCENT option was not employed. Characters
were assumedto be unordered(i.e., Fitchparsimony),
equally weighted, and gaps were treatedas missing
data. Parsimony uninformativecharacterswere ex-
cluded. Parsimonyanalyses surpassedmemory limi-
tations in the first addition replicate and 10,000
equally parsimonious topologies were retained for
branchswapping(L = 138, CI = 0.682, RI = 0.849).
The majority rule topology with clade supportesti-
mated from bootstrap analyses (Felsenstein, 1985)
with the above heuristic search settings but with
MAXTREESset to 100, is shown in Figure 2.
In the dioecious lineage of the Ocotea complex,
Rhodostemonodaphneand Ocotea s.str. are techni-
cally separatedby the arrangementof the four locelli
in whorl I and II stamens;locelli are in a shallow arc
or horizontalrow in Rhodostemonodaphnebut in su-
perimposed pairs in Ocotea. Endlicheria is distin-
guished from dioecious relatives solely by its two-
locellate anthers.The limited value of locelli number
in generic delimitationhas often been noted (e.g., van
der Werff, 1984; Burger, 1988; Rohwer et al., 1991),
and the two-locellate condition apparentlycircum-
scribes a polyphyletic assemblage of species in En-
dlicheria.
Moleculardataplace Endlicheriapunctulatawith
Ocoteapauciflora(Nees) Mez, a representativeof the
Ocotea cernua species group (sensu Rohwer, 1986),
in a clade that lies sister to remainingdioecious taxa
of the Ocotea complex (Chanderbaliet al., 2001; Fig.
2 herein). With members of the 0. cernua species
group E. punctulata shares pauciflorous inflores-
cences with sparselypubescentflowers and glabrous
concolorous leaves with immersed venation below.
Endlicheria coriacea differs from E. punctulata
mainly in habit and is certainlya second example of
a two-locellate species in the Ocotea cernua species
group.Rhodostemonodaphne elephantopusMadrifiin
most likely belongs here as well. Fourlocelli arranged
in a shallow arc refer this species to Rhodostemono-
daphne,but it is isolatedthere(Madriiinn,1996a),has
the leaf morphologyof the 0. cernua species group,
and is vegetatively indistinguishablefrom E. cori-
acea. The two species of Endlicheria here allied to
the Ocotea cernua species group, comprise the En-
dlicheriapunctulataspecies group.
Remaining Endlicheria sampled for molecular
charactersrepresent all species groups here recog-
nized (see NumericalList of Taxa),and all place with
a diverse representation of Rhodostemonodaphne
(Fig. 2). Although this Endlicheria-Rhodostemono-
daphne alliance is unresolved,it is itself significant.
Severalspecies of Endlicheriaaremorphologicallyso
similar to species of Rhodostemonodaphnethat they
seem misplacedin theirpresentgeneric station.With
these two genera in a strongly supported clade,
morphology-basedsuspicions (e.g., Rohwer et al.,
1991; Madrifinan, 1996a; Madrifiain,1996b), that sep-
arate lineages in Endlicheria are derived from
ancestorsthatwould fit the generic charactersof Rho-
dostemonodaphne,now have molecularsupport.
Among the species of Endlicheria clearly closer
to Rhodostemonodaphnethan to congeners are those
Kostermans(1937) assignedto sectionMicrolocellata
(E. bullata, E. longicaudata, E. rubriflora,E. spru-
cei), and a new species, E. ferruginosa, here
described. These five species comprise the Microlo-
cellata species group herein. As in Rhodostemono-
daphne, the locelli in these species are distally posi-
tioned in the anther (Fig. 1A, B), and flowers are
rotatewith fleshy tepals. At least E. rubrifloraandR.
antioquensis Madriiainappearto be sister taxa, and
it is likely that the others are each independentlyde-
rived fromRhodostemonodaphne.As I do not believe
RELATIONSHIPS 9

TABLE I.
Sources of plant material.Accessions markedby an asterisk (*) were previously submittedto Genbankby
Chanderbaliet al. (2001)

Genbank
accession
Taxon Provenance Voucher numbers
Endlicheriaanomala (Nees) Mez Brazil, Amazonas Lohmann343 (MO) AF363371

Endlicheriabracteolata (Meisn.) Brazil, Amazonas Ribeiro 885 (MO) AF363372

C. K. Allen
Endlicheriachalisea Chanderbali Guyana,Iwokrama Chanderbali252 (MO) AF272271*
Reserve

Endlicheriacitriodoravan der Peru, Iquitos Va'squez25231 (MO) AF272272*

Werff
Endlicheriadysodantha(Ruiz & Bolivia, Santa Cruz Vargas1098 (MO) AF363373
Pav.) Mez

Endlicheriagracilis Kosterm. Guyana,Iwokrama Chanderbali250 (MO) AF363374

Reserve

Endlicherialongicaudata (Huber) Brazil, Amazonas Assunqao366 (MO) AF363375

Kosterm.
Endlicheriametallica Kosterm. Ecuador,Napo Tirado 1010 (MO) AF363376
Endlicheriamultiflora(Miq.) Mez Guyana,Karanambo Chanderbali206 (MO) AF363377

Endlicheriapaniculata (Spreng.) Brazil, Parana Silva & Abe 2883 (MO) AF363378
J. F. Macbr.
Endlicheriapunctulata(Mez) C. K. FrenchGuiana de Granville1448 (MO) AF272273*
Allen

Endlicheriapyriformis(Nees) Mez Ecuador,Napo Neill 9842 (MO) AF363379

Endlicheriareflectens(Nees) Mez Guyana,Karanambo Chanderbali208 (MO) AF272274*

EndlicheriarubraChanderbali Peru, San Martin van der Werff 15436 (MO) AF363380

EndlicheriaruforamulaChander- Peru, San Martin van der Werff 15736 (MO) AF363381
bali
Endlicheriasprucei (Meisn.) Mez Brazil, Amazonas Vicentini 1224 (MO) AF363382

EndlicheriatessmanniiO.C. Peru, Loreto Vasquez 25237 (MO) AF363383

Schmidt
NectandraamazonumNees Guyana,Iwokrama Chanderbali217 (MO) AF272289*
Reserve
NectandracuspidataNees & Mart. Guyana,Kamarang Chanderbali279 (MO) AF272291*

NectandrapsammophilaNees & C. Brazil, Sao Paulo 5595 (MO)

Lorea-Hermandez AF272292*
Mart.
Nectandraturbacensis(HBK) Nees Puerto Rico, Rio Taylor 11746 (MO) AF272295*
Grande,El Verde
10 FLORANEOTROPICA

TABLEI.
Sources of plant material.Accessions markedby an asterisk (*) were previously submittedto Genbankby
Chanderbaliet al. (2001) (Continued)

Genbank
accession
Taxon Provenance Voucher numbers

Ocotea botranthaRohwer U.C. Riverside Scora 99-1 (UCR) AF272297*

Ocotea ceanothifolia (Nees) Mez Guyana,Demerara, Chanderbali(MO) AF272299*
MaburaHill
Ocotea guianensis Aubl. Guyana,Demerara, Chanderbali232 (MO) AF272302*
MaburaHill

Ocotea helicterifolia (Miesn.) Mexico, Oaxaca Torres11911 (MO) AF272303*

Hemsl.

Ocotea heydeana(Mez & Donn. Honduras,Yoro, Evans 1760 (MO) AF272304*

Sm.) Bernardi Pico Pijol
Ocotea nigra Benoist Guyana,Iwokrama Chanderbali162 (MO) AF272308*
Reserve

Ocotea pauciflora (Nees) Mez Guyana,Demerara, Chanderbali219 (MO) AF272310*

MaburaHill

Ocotea percoriacea (Mez) Kosterm. Brazil, Goias Lorea-Hernandez5584 (MO) AF272311*

Ocotea pulchella Mart. Brazil, Minas Ger- Lorea-Hernandez5575 (MO) AF262312*

ais

Ocotea schomburgkiana(Nees) Guyana,Iwokrama Chanderbali286 (MO) AF272315*

Mez Reserve

Ocotea spixiana (Nees) Mez Brazil. Minas Ger- Lorea-Hernandez5574 (MO) AF272316*
ais

Ocotea tomentellaSandwith Guyana,Kamarang Chanderbali284 (MO) AF272317*

Ocotea tristis (Nees & Mart.)Mez Brazil, Minas Ger- Lorea-Hernandez5577 (MO) AF272318*
ais

Pleurothyriumcinereumvan der Peru, San Martin van der Werff 15325 (MO) AF272329*
Werff

Pleurothyriuminsigne van der Ecuador,Napo, Ja- Neill 9033 (MO) AF272330*

Werff tun Sacha

Rhodostemonodaphnecrenaticupula Guyana,Iwokrama Chanderbali265 (MO) AF272331*

Madrifian Reserve

Rhodostemonodaphnekunthiana Guyana,Iwokrama Chanderbali257 (MO) AF363384

(Nees) Rohwer Reserve

Rhodostemonodaphnenegrensis Brazil, Amazonas Vicentini 628 (MO) AF363385

Madrifian
Rhodostemonodaphneparvifolia Brazil, Amazonas Assun,cao327 (MO) AF363386
RELATIONSHIPS
TABLE I.
Sources of plant material.Accessions markedby an asterisk (*) were previously submittedto Genbankby
Chanderbaliet al. (2001) (Continued)
Taxon Provenance
peneia Mad-
Rhodostemonodaphne Brazil, Amazonas
rii~ian
Rhodostemonodaphnepraeclara Guyana,Iwokrama
(Sandwith)Madrinan Reserve
Rhodostemonodaphnerecurvavan Brazil, Amazonas
der Werff
Rhodostemonodaphnescandens Guyana,Iwokrama
MadriiaTn Reserve
these species form a monophyleticgroup,their affin-
ities are discussed individually in the Systematic
Treatment,below.
A likely monophyletic group crossing the Endli-
cheria-Rhodostemonodaphneboundaryis comprised
of Endlicheriachrysovelutina,E. metallica, Rhodos-
temonodaphnegrandis, R. praeclara (Sandw.)Mad-
rifia'n,R. peneia MadriiaTn,R. saiulensisMadrifian,
and R. recurva van der Werff. All share the peculiar
asymmetric basal secondaries described in the leaf
venationsection above, sericeous indument,andlarge
fruits with robusthemisphericalcupules.They assign
to differentgeneraon the basis of locelli number,but
R. recurvais conceivably intermediatein combining
four-locellateanthersin whorl III stamenswith two-
locellate anthersin whorls I and II. Althoughflowers
are rotate in species assigned to Rhodostemonoda-
phne, but campanulatein the two assigned to Endli-
cheria, intermediateconditions are found in material
that may representundescribedspecies. Thus, Stey-
ermark59001 from Bolivar,Venezuela(F, MO) com-
bines campanulateflowerswith four-locellateanthers,
while in Prance et al. 14616 from Amazonas,Brazil
(INPA, NY), and SdnchezVegaet al. 9345 from San
Martin,Peru(MO), two-locellateanthersarefoundin
rotate flowers. The latter two collections may repre-
sent additionaltwo-locellate species in this transge-
neric species group, but are not described here be-
cause anthers are poorly developed and devoid of
pollen, possibly indicating hybrid origin. They key
out to E. metallica aff. sp. 1 and E. metallica aff.
sp. 2, respectively. Pending better material of these
entities, E. metallica and E. chrysovelutinaare the
11
Genbank
accession
Voucher numbers
Ramos 2834 (MO) AF363387
Chanderbali256 (MO) AF272332*
Vicentini 653 (MO) AF272333*
Chanderbali271 (MO) AF272334*
only described members of the E. metallica species
group.
The Endlicheriabrownianaspecies group(species
10-18) may also find its closest relatives in Rhodos-
temonodaphne.Here, stamensof all whorls arefleshy
and sessile, andanthersareovate with obliquelyhem-
isphericallocelli between andbeyond which an apic-
ulate connective protrudes (Fig. ID). In most, the
flowers are trumpet-shapedwith slenderpedicels that
widen distally to merge with an infundibuliformre-
ceptacle. Especially in E. browniana,E. jefensis, and
E. mishuyacensis, flowers are densely papillose in-
side. Such papillosityis rarein Endlicheriabuttypical
of Rhodostemonodaphne.,wherein similarly shaped
flowers are found in its type species, R. laxa (Meisn.)
Rohwer and, among others, in R. cyclops Madriiain
where whorls I and II antherscan be three-locellate,
conceivablyillustratingthe transitionfromfourto two
locelli.
Remaining species of Endlicheria can be sorted
into groupsthat appearto be linkedby intermediates.
Still, ratherthan constitutinga single lineage, these
species groups appear to intermingle freely with
counterpartsin Rhodostemonodaphne.
One close-knit group of species (19-34) includes
those previously assigned to Ampelodaphne(Meis-
sner, 1864; Mez, 1889), herein, the Ampelodaphne
species group. It is characterizedby subverticillate
leaves and bracteate inflorescences with numerous
fragile flowers that have a deep, almost tubular,re-
ceptaclebelow spreadingto recurvedmembranaceous
tepals. Here, all stamens have slender distinct fila-
ments, andtransverselyoblong anthersaredominated
12 FLORA NEOTROPICA

100 Endlidsefl nUIOmIaTI

53 Enck*_ biatodsta
.. . _~anma
100 E d
EdIdeapanbiata
L---Ersdtwi baIs
gmn
100

86 ioEcoeipffni

100
94
RhdX

. _S~~~~~Peu
100~~~~~10
66~ 610 - cm-1 I p e group

_10
73 r co ni
~81

10 1ade0100 . group0p

. 2. e ase a. T oit e po

810
100 r EfltnpvMtlaraw
00W.C
0 dibta0 Iva 0
paired w all h s oE and
ue

....e0..C - - - - e "
.100 , anb00 - f p

100p 10*--- p q
. 0. ocd R
ills-
FI. 2. Phloeneiffnte of.EndichriaindctdbIT seuenc dt. T..is.... majriymetployo
th .00 equa legt tre -L = 138 C- = 0.82 RI' =F.4)rtie o rac wpigsosE dlceipntaa
clades areindicated.~ ~ ~ ~ ~ ~ ~ ~ ~~~nd

pairewit Ocyoteanetuifr affndtiesoftndiherseiesofindicheianted 50% majrt

wlSsqunedta.Rhodstmndpe wellesuppologyeo

but poorly resolved clade. Majorityrule percentagesare indicatedabove, and bootstrapvalues >50% below, branches.Main
clades are indicated.
RELATIONSHIPS
by suborbicularlocelli (Fig. lH). Two subgroupsare
suggested by venation and leaf texture.In the Endli-
cheria multiflorasubgroup,also with E. arenosa, E.
dictifarinosa,E. levelii, E. macrophylla,and E. mel-
inonii, tertiaryand higher-orderveins are always im-
mersed above and thus inconspicuousagainstusually
coriaceouslaminae.In the E. bracteatasubgroup,ter-
tiary and fourth-orderveins are prominent above,
forming a conspicuousreticulatepatternover usually
chartaceouslaminae. Othermembersof the E. brac-
teata subgroupare E. chalisea, E. cocuirey,E. direc-
tonervia, E. glomerata, E. tessmannii, and E. verti-
cillata. Representatives of both subgroups, E.
chalisea, E. multiflora,and E. tessmannii,constitute
one of the few clades resolved in the Endlicheria-
Rhodostemonodaphnealliance by ITS sequence var-
iation (Fig. 2). This clade lies sister to a pair of Rho-
dostemonodaphnespecies while Endlicheria refiec-
tens, a species somewhat intermediatebetween the
two subgroupsin having the adaxiallyprominentter-
tiaries of the E. bracteata subgroupbut the rusty to-
mentose indument typical of the E. multifiorasub-
group, appearsto share molecularcharacterswith E.
sprucei (Fig. 2). This resolutionis weakly supported,
but it is intriguing from a morphological point of
view. The two species of Rhodostemonodaphne,R.
crenaticupulaMadrinia'n and R. negrensis Madrinian,
placed close to core members of the Ampelodaphne
species group are morphologicallyvery similar to E.
sprucei. Indeed, fruiting specimens of R. crenaticu-
pula are sometimes mistaken for E. sprucei (Madri-
nian,1996b). Further,the sessile outer stamens of E.
sprucei (e.g., Fig. 1B) are typical of R. crenaticupula.
Therefore,if the molecular signal for close relation-
ship between E. reflectensand E. sprucei has phylo-
genetic significance, it is conceivable that the Ampe-
lodaphne species group relates to members of
Rhodostemonodaphnevia E. sprucei.
Like members of the Ampelodaphne species
group, Endlicheria anomala and E. williamsii have
stipitatestamensin all staminalwhorls,buttheirflow-
ers are otherwise quite different.In both, the flowers
are rotate with horizontally spreading tepals, and
whorl III anthershave four-locellate anthers(incon-
sistently so in E. williamsii) althoughwhorls I and II
are two-locellate. Rhodostemonodaphnerecurvaand
R. revolutifoliashow similarvariationin locelli num-
ber among androecialwhorls, but close relationship
with these is not supportedby other characters.ITS
sequence data places Endlicheriaanomala in the En-
dlicheria-Rhodostemonodaphne alliance without
clearerindicationsof affinities.Endlicheriaanomala
13
and E. williamsii comprise the E. anomala species
group.
In all remaining species of Endlicheria, whorl I
and II stamens are stipitate,to a greateror lesser de-
gree, but whorl III stamens are broadly stipitate or
sessile with filamentsequal to or wider than anthers.
On the basis of indument and stamen morphology,
three groups of similar, and presumablyclosely re-
lated, species can be recognized.
One such group is comprised of Endlicheria ca-
nescens, E. citriodora, E. duotincta, E. lorastemon,
E. oreocola, E. rubra,E. szyszylowiczii,E. vinotincta,
andE. xerampela(species 37-45). Most have a dense
reddish tomentose indumentumcovering branchlets,
and whorls I and II stamenshave densely greyish to-
mentose filamentsand ovate antherswith an apically
prolonged connective (Fig. lE). These stamens re-
semble those of Aniba Aubl., and at least flowers of
E. citriodora are strikingly similar to those of that
genus. However,moleculardataunambiguouslyplace
E. citriodora (and E. rubra)in the Endlicheria-Rho-
dostemonodaphnealliance (Fig. 2) and associateAn-
iba with Licaria Aubl. (Chanderbaliet al., 2001).
Endlicheriaxerampelais placed here on the basis of
overall resemblance to E. canescens, although its
whorl I and II anthersare truncateabove.
Members of the Endlicheria canescens species
group appearto be related to Rhodostemonodaphne
species with similar dense reddish indumentum.In
particular,E. vinotincta is almost indistinguishable
from R. celiana C. K. Allen and R. steyermarkiana
(C. K. Allen) van der Werffwithoutcounting locelli;
in venation and leaf shape E. citriodora and R. kun-
thiana (Nees) Rohwer are extremely similar;E. ca-
nescens shows the asymmetricbasal secondaries of
R. rufovirgataMadriiainand R. mirecolorata(C. K.
Allen) Rohwer;and E. rubrais vegetativelyindistin-
guishablefrom material,Garc(a-Barriga13944 from
Amazonas, Colombia (NY, US), representingan un-
describedspecies of Rhodostemonodaphne. Relation-
ships among these and dioecious species of Ocotea
with a similarvestiture,e.g., 0. discrepensC. K. Al-
len, 0. endlicheriopsisMez, 0. indirectinerviaC. K.
Allen, and 0. rufovestitaDucke, should be investi-
gated further.
The Endlicheria sericea species group (species
46-55) comprises species with stamens similar to
those in the E. canescens species group but with a
dense sericeous indumentthat usually persists on the
lower leaf surface.Here, leaves are usually ovate and
coriaceous with arcuatesecondariesthat tend to fol-
low asymmetriccourses.Close relationshipwith End-
14 FLORANEOTROPICA

licheria metallica, E. chrysovelutina,and their allies all anthersare four-locellatewith the locelli arranged
in Rhodostemonodaphneis suggested by shared se- in superimposedpairs and species either have bisex-
riceous indument, and occasional asymmetricbasal ual flowers or are dioecious. Rhodostemonodaphne
secondaries in E. klugii and E. ruforamula,but the differs from dioecious Ocotea only in having its four
stipitatewhorlI and II stamensin the E. sericea group locelli arrangedin a shallow arc, while Endlicheria
speak against it. Instead, a closer link to Rhodoste- differs from Rhodostemonodaphneand dioecious
monodaphnemay be providedby material,e.g., Ac- Ocotea only in having two-locellate anthers. Since
evedo et al. 1370 (F, MO, US), of an undescribed Endlicheria and Rhodostemonodaphnenest in dioe-
species. There, the flowers are similar in size and cious Ocotea (Chanderbaliet al., 2001; Fig. 2 herein),
shape to those of E. griseo-sericea but anthers are a rather broad generic concept would conceivably
three-locellatein whorls I and II and four-locellatein treatthis dioecious lineage as one genus; viz. Ocotea.
whorl III. This unusualandroeciumis more like that This would make Ocotea monophyletic,providedthat
of RhodostemonodaphnethanEndlicheria,but in leaf its current bisexual members are removed, but
shape and venation this species appearscloser to the whetherdemotingthe delimitationof Endlicheriaand
E. sericea species group. It is not described for rea- Rhodostemonodaphneto infrageneric significance
sons given in the discussion of E. griseo-sericea. improves the current situation is questionable.Per-
Endlicheriaacuminata,E. gracilis, and E. kruko- haps the Endlicheria-Rhodostemonodaphne alliance
vii have triplinervedleaves andappearto form a loose alone can be consideredfor generic status.However,
association aroundE. paniculata. All have stipitate such a groupis too heterogeneousto be circumscribed
whorl I and II stamens, and in E. acuminata,E gra- in morphologicalterms,and might yet be found to be
cilis, and usually also E. paniculata, antherapices are paraphyleticwith respect to species of Ocotea. An
truncateratherthan apiculate.Endlicheriakrukoviiis opportunityto revise generic conceptswill arisewhen
close to E. acuminata, althoughits antherapices are relationshipsamong the membersof these threegen-
apiculate.Endlicheria nilssonii is poorly known but era are more fully elucidated. Until then, this treat-
is placed here on the basis of its stipitatestamenswith ment addresses the great practicalnecessity of pro-
truncateantherapices. viding modem revisions at the species level in
The above species groups are informal concepts Lauraceae, even before all the generic concepts are
meant to parcel infragenericheterogeneity into more solidified.
homogeneous units that can each be considered in
more detail. In the case of the Endlicheriapunctulata SPECIES CONCEPTS
species group (species 1-2), affinitiesare confidently Specific status is here grantedto the smallest as-
assumedto lie with Ocotea, while the Microlocellata semblage of individuals(collections) that is morpho-
species group (species 3-7) and E. metallica species logically distinguishable from other such assem-
group(species 8-9) areclearlybest comparedto Rho- blages. Species so circumscribedpossess at least one
dostemonodaphne.In the case of the E. browniana unifyingcharacteror charactercombinationthatis not
species group (species 10-18), counterpartsin Rho- found in other species, and may either be internally
dostemonodaphneare conceivably provided by R. uniform or variable. Infraspecific categories are
laxa and allies. The Ampelodaphne species group avoided because the causes of infraspecificvariation
(species 19-34) has deviated from ancestralcondi- are unknown.
tions and become very distinctive, but may link to The source of species charactersvaries with spe-
Rhodostemonodaphnevia E. sprucei. The E. sericea cies and species groups.Vegetativemorphologyis al-
species group (species 46-55) is also very distinctive most uniformin the Endlicheriasericea species group
a
but may include species with the generic characters but floral variationis important,while flowers are al-
of Rhodostemonodaphne.The E. canescens species most identical in theAmpelodaphneandE. punctulata
species groups but there is significantvegetativevar-
group (species 37-45) is heterogeneous.At least E.
vinotinctaandE. rubrahave clearfour-locellatecoun- iation. In the other species groupsdistinguishingchar-
acters are provided by both floral and vegetativevar-
terpartsin Rhodostemonodaphne,but the affinitiesof
and E. pan- iation.
remainingmembers,like the E. anomala
iculata species groups, are unclear.
DISTRIBUTION AND
Given the numeroustransgenericalliancesinvolv-
ing species of Endlicheria, Rhodostemonodaphne,
BIOGEOGRAPHY
and to a lesser extent Ocotea, the currentconcepts of Endlicheria is a neotropical genus centered in
these threegenerawarrantcarefulrevision.In Ocotea South America, while reachingCosta Rica in Central
DISTRIBUTIONAND BIOGEOGRAPHY
America, Guadeloupein the CaribbeanIslands, and
its southernmostextreme in the Atlantic coastal for-
ests of SE Brazil (Fig. 3). This distributionis almost
paralleledby Rhodostemonodaphne(lacking only in
the Caribbean).Ocotea s.str. is similarly distributed,
but more widespreadin CentralAmerica and the Ca-
ribbean. Closely related Nectandra s.str. and Pleu-
rothyriumare also similarly distributed.The biogeo-
graphic significance of the commonality in these
generic distributions was partially addressed by
Chanderbaliet al. (2001).
According to the biogeographichistory of Laura-
ceae reconstructedthere, the lineage to which these
genera belong, namely the Ocotea complex, has an
ancestralMadrean-Tethyandistribution(sensu Axel-
rod, 1975) and radiatedin the New Worldduringthe
Miocene. Furthermore,they constitutea South Amer-
ica-centeredclade thatdivergedfrom a CentralAmer-
ica-centered sister group, the Ocotea helicterifolia
species group, ca. 20 million years ago (Chanderbali
et al., 2001). As this divergence time coincides with
-----------
,'----- ----- ....f
------
~~~~~X----- -
---------------------
e~~~~~~~IDistibuio
30
S.I2
FIG. 3. Distributionof Endlicheria.
15
increasedAndean uplift in the Early Miocene, Chan-
derbali et al. (2001) envisaged that the resulting ele-
vational barriers to migration isolated the ancestors
of Endlicheria and its sympatric relatives in South
America. The subsequent biogeographic history of
these genera and Endlicheria itself has not been in-
vestigated in detail but conceivably involves a com-
plex scenarioof diversificationalong edaphicandele-
vational gradientsin South America and dispersalto
CentralAmerica and the CaribbeanIslands.
The species of Endlicheria occur in moist forest
habitatsfrom elevations of aroundsea level to 2500
m in the Andean and Guianianhighlands. The limi-
tations imposed by low water availabilityare evident
in the absence of Endlicheriafrom the caatingaveg-
etation of eastern Brazil and the Chaco region of
southernSouth America (Fig. 3). Species diversityis
greatestin the easternAndeanfoothills wherelowland
and lower montane elements intermingle. Central
Amazonia and the Guianas are not nearly as rich in
diversity,andin the Atlanticcoastal forests of SE Bra-
-------------- ----- ------- - --- ------- -
of Endlih;r;a
W , k
16 FLORANEOTROPICA

zil, only two species are found. The distributionof E.
paniculata almost equals the generic range of Endli-
cheria (lacking only in NE South America and the
CaribbeanIslands)but all otherspecies havenarrower
ranges.Severalspecies arewidespreadthroughoutthe
lower slopes of the eastern Andes to western, and
rarely central, Amazonia, e.g., E. metallica and E.
rubriflora.Of these, E. formosa and E. ruforamula
rangewest of the Andes to CostaRica andColombian
Choco, respectively.Endlicheriabracteolata,E. chal-
isea, E. pyriformis,and E. szyszylowicziirange from
the Guianasthroughthe Amazon basin to lower mon-
tane eastem Andean slopes, while E. canescens and
E. gracilis range from the Guianasto the easternAn-
dean slopes without a central Amazonian presence.
Lowlandspecies with wide distributionsall appearto
be elements of riparianforests, e.g., E. anomala, but
narrowdistributionsfor species of terrafirmemay be
collecting artifacts(see Nelson et al., 1989). The pos-
sible exceptions, as suggested by relatively well-
collected species, are rare.Endlicheriaglomerataap-
pearsto be endemicto the coastalforestsof SE Brazil;
E. sericea is essentially an element of the lower
montane slopes of the Lesser Antilles and Trinidad
butpossibly also in adjacentVenezuela;andE. brown-
iana is restricted to the western side of the Andes
from Ecuadorto Panama.
SYSTEMATIC TREATMENT
GENERIC DESCRIPTION
Endlicheria Nees, Linnaea 8: 37. 1833, nom. cons.
Type. Endlicheria hirsuta (Schott) Nees (lecto-
type, designated by Kostermans,1937; = Endli-
cheria paniculata (Spreng.)J. F. Macbr.).
Goeppertia Nees,Syst.laur.365. 1836.Type.Goepper-
tia hirsuta(Schott)Nees(lectotypeheredesignated;
= Endlicheria paniculata(Spreng.)J. F. Macbr.).
Ampelodaphne Meisn., DC. Prodr.15(1): 81. 1864.
Type.Ampelodaphne macrophylla(lectotypehere
designated).
Huberodaphne Ducke,Arch.Jard.RiodeJaneiro 4: 191.
1925. Type. Huberodaphne longicaudata Ducke
(lectotypeheredesignated).
Endlicheriasubgen. Ampelodaphne(Meisn.) Mez,
Jahrb.Konigl.Bot.Gart.Berlin5: 125. 1889.Type. nym of Aiouea Aubl.
Ampelodaphnemacrophylla (lectotype here desig-
nated).
Endlicheria subgen. EuendlicheriaMez, Jahrb.Konigl.
Bot. Gart. Berlin 5: 114. 1889. Type. Endlicheria
brownianaMez (lectotype here designated).
Endlicheria subgen. Hemiajouea Mez, Jahrb.Konigl.
Bot. Gart. Berlin 5: 114. 1889. Type. Endlicheria
paradoxa Mez (lectotype here designated).
Endlicheriasubgen.OcoteopsisMez, Jahrb.Konigl.Bot.
Gart.Berlin 5: 132. 1889. Type. Endlicheriaanom-
ala (Nees) Mez (lectotype here designated).
Endlicheriasect. MicrolocellataKosterm.,Recueil Trav.
Bot. NMerl.34: 511. 1937. Type. Endlicheriabullata
Ducke (lectotype here designated).
Endlicheria sect. Macrolocellata Kosterm., Recueil
Trav. Bot. NMerl.34: 517. 1937. Type. Endlicheria
gracilis Kosterm.(lectotype here designated).
Trees or shrubs. Leaves alternate,widely spaced
along branchletsor arrangedin close spirals;mostly
penninerved,occasionally asymmetricallytrinerved,
tripli- or quintuplinerved.Inflorescences mostly in
axils of foliage leaves, but often in cataphylls,thyrso-
paniculate, the terminal dichasia rarely reduced to
pseudo-umbelsor botryoids.Flowersunisexual,trim-
erous. Tepals 6, in 2 whorls of 3, usually spreading,
rarely inflexed, erect, or reflexed. Stamens 9 in 9
whorls of 9; whorl I and II stamens ? equal, the
filamentsdistinctor undifferentiated,the antherstwo-
locellate; whorl III stamens usually sessile with in-
distinctly differentiatedfilaments,rarelystipitate,the
antherstwo-locellate, rarely four-locellate,the basal
glands sessile or stipitate,rarelyabsent;fourthwhorl
of staminodesoccasional;pistillodepresentor absent.
Pistillate flowers usually deeper with smaller sterile
stamens;ovary superior;stigma tri-lobed, discoid or
reniform.Fruitsborneon tereteor claviformpedicels;
cupules usually hemispherical, occasionally flat,
rarely reflexed, the margins entire, undulateor with
persistenttepal bases; drupeselliptic, obovoid, ovoid,
or spheroid.
The name Schauera was only tentativelyoffered
in place of Endlicheria by Nees (in Lindley, 1836)
and is thereforenot validly published.
Endlicheriasubgen. AnosphaeriaMez is a syno-

Key to the species of Endlicheria

1. Lower surfaceof matureleaves concealed by the indumentcover.
2. Indumenton lower leaf surfacecomprisedof a dense creamishtomentose cover interspersedwith
longer reddish, straight,erect hairs.
3. Leaves alternate,widely distributedalong branchlets.Flowers campanulate,densely pubescent
outside. Outeranthersapiculateabove. E Andean foothills.......................................................... 44. E. duotincta
SYSTEMATICTREATMENT 17

3. Leaves clusteredat tips of branchlets.Flowers rotate, sparselypubescentoutside. Outeranthers

truncateabove. W Amazonianlowlands................................................................. 36. E. williamsii
2. Indumenton lower leaf surface comprisedof straightappressedhairs, these silvery-greyto golden,
uniformin length and orientation.
4. Flowers sparselypubescentoutside, the surfaceclearly visible.
5. Flowers rotate;tepals spreadinghorizontally.Anthersof whorl III stamensfour-locellate.
Flooded Amazonianlowlands .............................................................. 35. E. anomala
5. Flowers obconical or urceolate;tepals inflexed or erect. All antherstwo-locellate. Nonflooded
forests of E Andean foothills and SW Amazonia................................................................... 55. E. robusta
4. Flowers densely pubescentoutside, the surfacenot visible.
6. Basalmost secondariesasymmetric,one or both mergingwith leaf marginnear base or emerging from
junction of midrib and leaf base.
7. Tepals broaderthan long. Whorls I & II stamens sessile, their anthersover toppedby a broad
sterile truncateapex.
8. Branchletssericeous, the hairs closely appressedto the surface, silvery grey. E Andes to central
Amazonia.............................................................. 8. E. metallica
8. Branchletsdensely velutinous,the hairs erect, golden to yellowish brown.White sands around
Iquitos, Peru.............................................................. 9. E. chrysovelutina
7. Tepals longer than broad.Whorls I & II stamens stipitate,their anthersovate with narrowly
apiculateapices.
9. Branchletssericeous, the hairs appressed.Tepals spreadinghorizontally;receptaclenarrowly
infundibuliform,longer than broad;whorls I & II filamentsS-shaped.Cupules shallow, drupes
exserted.W Amazonia to E Andean slopes.............................................................. 53. E. klugii
9. Branchletstomentellose,the hairs ascending.Tepals ascending;receptacleas broadas or
broaderthan long; outer filamentsstraight.Cupulesdeeply hemispherical,drupesat least 1/4
included. CentralAmazoniato W Andean slopes of Colombia and Ecuador.................... 50. E. ruforamula
6. Basalmost secondaries ? symmetric,emergingfrom midrib above leaf base.
10. Leaves tripli- or quintuplinerved.The lowermostpair(s) of secondariessuboppositeshortlyabove
the leaf base, arcuate,ascending to beyond midlamina,the othersemergingbeyond midlaminaor
absent.
11. Flowers infundibuliformor campanulate,the tepals ascendingat anthesis. Guianas,
Amazonia, Pacific coast of Colombia.................................................................... 47. E. bracteolata
11. Flowers rotate,the tepals spreadinghorizontallyat anthesis.Lesser Antilles, NE
Venezuela.................................................................... 46. E. sericea
10. Leaves penninerved.Secondaryveins ? uniformlyspaced along midrib.
12. Both branchletsand inflorescenceaxes tomentelloseor tomentose,the hairs ascendingor
sprawling.
13. Hairs on branchletsca. 0.2 mm long or shorter.Tepals erect to ascending,the inner
surfaceglabrous.CentralAmazoniato W Andean slopes of Colombia and Ecuador
..................................................................................................................................50. E.ruforamula
13. Hairs on branchletsca. 0.5 mm long. Tepals spreading,the inner surfacepubescent.
14. Leaves ovate. Hairs on branchletspale to rust red. Tepals chartaceous,sparsely
pilose inside. Mato Grosso, Brazil....................................................... 52. E. lhotzkyi
14. Leaves obovate. Hairs on branchletsgreenish yellow. Tepals fleshy, densely
papillose inside. Amazonas,Brazil..................................................... E. metallica aff. sp. 1
12. Eitheror both branchletsand inflorescenceaxes silvery to rusty sericeous, the hairs closely
appressedto the surface.
15. Receptacle narrowlyinfundibuliform,longer than broad.Pedicels claviform,indistinctin
flower and fruit. Plants mostly from lowland Amazonia at ca. 100-200 m.
16. Midrib prominentabove. Filamentsof whorl I & II stamens S-shaped.W
Amazonia to E Andean slopes ......................................................... 53. E. klugii
16. Midrib impressedabove. Filamentsof whorl I & II stamens straight.W Amazonian
lowlands.................................................... 54. E. argentea
15. Receptacle campanulateor shallowly infundibuliform,broaderthan long. Pedicels ?
terete, distinct in flower and fruit. Plants mostly from lower montaneAndean slopes
above 500 m.
17. Leaves drying darkolive-brownabove. Receptacle constrictedbelow tepals. Whorl
I and II anthersobovate, the apex broadlytruncateto retuse over the locelli. Lower
montaneAndes .................................................... 49. E. griseo-sericea
18 FLORA NEOTROPICA

17. Leaves drying greenish above. Receptacle graduallymerging with tepals. Whorl I
and II stamensovate or depressed-ovate,the apex apiculatebetween the two locelli.
18. Indumenton branchlets,leaves, and flowers golden to rust brown. Anthersof
all whorls depressed-ovate,the locelli ? apical. Cupule marginsstrongly
lobed. E Andes ............................................ 51. E. aurea
18. Indumenton branchlets,leaves, and flowers tawny to silvery grey. Anthersof
whorls I & II ovate, the locelli introrse-latrorse; anthersof whorl III oblong,
locelli extrorse-latrorse.Cupule marginsentire or weakly undulate,not
stronglylobed.
19. Flowers rotate,the tepals spreading,densely grey-tomentoseinside.
Cupules deeply hemispherical,densely sericeous inside. Lesser Antilles,
NE Venezuela ........................................ 46. E. sericea
19. Flowers campanulateor infundibuliform,the tepals erect to ascending,
glabrousinside. Cupules shallow, glabrousinside. NW South America,
Panama..................................................................... 48. E. tschudyana
1. Lower surfaceof matureleaves clearly visible throughindumentcover.
20. Leaves and inflorescencesclusteredin close spirals at the tips of branchlets.Inflorescencebractsand
bracteolespersistentat anthesis.
21. Tertiaryand higher-orderveins immersedabove, inconspicuousagainstthe lamina.
22. Inflorescenceaxes and exteriorof flowers glabrous.
23. Hairs on branchletsand lower leaf surfacereddishto rust brown, erect. Upper Rio
Negro, Brazil, Venezuela................................................... 19. E. arunciflora
23. Hairs on branchletsand lower leaf surfacecreamishwhite, weak, sprawling,silky.
PeruvianAmazonia .......................................................... 20. E. arachnocome
22. Inflorescenceaxes and exteriorof flowers moderatelyto densely pubescent.
24. Hairs on sides of midriband secondariesbelow stiffly straight.
25. Hairs on lower leaf surfacevery short, ca. 0.2 mm. Well-drainedwhite sand soils.
CentralAmazonia............................................... 21. E. arenosa
25. Hairs on lower leaf surfacerelativelylong, ca. 0.5 mm. Flooded forest, NE South
America............................................... 26. E. melinonii
25. Hairs on sides of midriband secondariesbelow curved or crooked,not straight.
26. Leaves ovate to elliptic. Basal glands large, surroundingfilamentsof the
inner staminalwhorl. CentralAmazonia ........................................... 22. E. macrophylla
26. Leaves obovate to obovate-elliptic.Basal glands minute, inconspicuous,
leaving the filamentsof the inner staminalwhorl exposed.
27. Hairs on branchlets,midribbelow, and inflorescenceaxes reddishto rust
brown. Guianas....................................... 23. E. multiflora
27. Hairs on branchlets,midribbelow, and inflorescenceaxes greyish to
white.
28. Leaves subsessile, the base (sub-)cordate,abruptlycontractedinto
the petiole. SE Venezuela,Roraima,Brazil ......................... 25. E. dictifarinosa
28. Leaves petiolate, the base acute, graduallytaperinginto the petiole.
Centralto W Amazonia.................................... 24. E. levelii
21. Tertiaryand higher-orderveins raised above, forming a prominentnetworkover the lamina.
29. Tertiaryveins reticulatingbetween secondaries.Shrubsfrom Guianashield savanna
formations....................................................... 27. E. reflectens
29. Tertiaryveins undividedor once-forkedbetween adjacentsecondaries.Trees from wet
forests.
30. Hairs on lower leaf surface appressed.Iquitos, Peru, and vic............................... 31. E. tessmannii
30. Hairs on lower leaf surfaceerect.
31. Branchietsand midribbelow tomentose or tomentellose,the hairs ca. 0.5 mm
long or shorter,if longer then sprawling.
32. Leaves subsessile, the base subcordate,abruptlycontractedinto the petiole.
W Amazonia........................................... 30. E. cocuirey
32. Leaves petiolate,the base acute or obtuse, graduallytaperinginto the petiole.
CentralAmazonia to E Andean slopes ........................................... 32. E. directonervia
31. Branchletsand midribbelow hirsute,the hairs ca. 1 mm or longer, straight,stiff.
33. Hairs on branchletsreddish.Distal branchesof inflorescencesand exteriorof
flowers glabrous.CentralAmazonia.......................................... 29. E. verticillata
SYSTEMATICTREATMENT 19

33. Hairs on branchletsgreyish white or yellow. Inflorescenceaxes and exterior

of flowers densely pubescent.
34. Indumenton vegetative surfaces greyish white. Flowers pedicellate,
distantalong terminalcymes. CentralAmazonia to E Andes.......... 28. E. bracteata
34. Indumenton vegetative surfacesyellowish. Flowers subsessile,
clustered.
35. Laminaeplane and cupules densely pubescentoutside, the surface
obscured.Guianas,Amazonia, and E Andean slopes ................ 33. E. chalisea
35. Laminaebullate or plane, but cupules always glabrousoutside. SE
Brazil ......................................... 34. E. glomerata
20. Leaves and inflorescencesarrangedevenly along branchlets.Inflorescencebractsand bracteoles
caducousby anthesis.
36. Terminalbuds glabrousexcept for a distal ciliate fringe of hairs.Young leaves and branchlets
completely glabrous.Guianasto E Andes ................................................................... 14 .E. pyriformis
36. Terminalbuds densely pubescent,the surfacecompletely covered.Young leaves and/or
branchletspubescent.
37. Indumenton leaves below closely appressedto the surface.
38. Branchletsmidway along flush sparselypubescent,the surfaceclearly visible.
39. Basalmost secondariesasymmetric,one or both mergingwith leaf marginnear
base or emerging fromjunction of midrib and leaf base.
40. Flowers campanulate,the tepals erect, glabrousinside. CentralAmazoniato
E Andes ................................................ 8. E. metallica
40. Flowers rotate,the tepals spreadinghorizontally,densely pubescentinside.
San Martin,Peru................................................ E. metallica aff sp. 2
39. Basalmost secondaries ? symmetric,emergingfrom midrib above leaf base.
41. Axils of secondaryveins below with barbellatetufts of hairs. SW Amazonia
and E Andeanfoothill ................ ..................................... 17. E. dysodantha
41. Axils of secondaryveins below withoutbarbellatetufts of hairs.
42. All staminalwhorls stipitate,the filamentsdistinct and narrowerthan
anthers.Basal glands large, filling the space between stamens.Inner
surface of tepals glabrous.
43. Midrib and secondaryveins immersedbelow; tertiaryand higher-
orderveins immersedabove; small trees or shrubs.NE South
America......................................... 1. E.punctulata
43. Midrib and secondaryveins prominentbelow; tertiaryand higher-
orderveins prominentabove. Large trees to 30 m. Central
Amazonia, Brazil......... ................................ 2. E. coriacea
42. Outerwhorls of stamenssubsessile, the filamentsinconspicuous,or if
obvious as broadas or broaderthan anthers.Basal glands
inconspicuous.Innersurfaceof tepals pubescent.
44. Flowers depressed-globose,the tepals inflexed, revealingonly a
small terminalpore at anthesis.
45. Tertiaryveins straightor forked between adjacentsecondaries.
Flowers ca. 2 mm diam. CentralAmazonia and uplandsfrom
Peru to Costa Rica..................................... 15. E. formosa
45. Tertiaryveins reticulatingbetween adjacentsecondaries.
Flowers 5-7 mm diam. Cajamarca.Peru.......................... 16. E. paradoxa
44. Flowers hypocrateriform,campanulate,or rotate,the tepals
ascending,horizontallyspreading,or reflexedat anthesis.
46. Anthersof whorl I & II stamensreniform,the apex truncateto
rounded.Locelli suborbicular,minute,just below apex. W
Amazonia to E Andes .............. ....................... 4. E. rubriflora
46. Anthersof whorl I & II stamensovate, the apex apiculate.
Locelli obliquely hemispherical,occupying entire anther.
47. Tertiarieslaxly reticulatingbetween adjacentsecondaries.
48. Indumenton branchletsrust brown.W
Amazonia.............. ................... 13. E. mishuyacensis
48. Indumentsilvery grey. CerroJefe, Panama.......... 11. E. jefensis
20 FLORANEOTROPICA

47. Tertiariesstraightor once-forkedbetween adjacent

secondaries.
49. Receptacleconstrictedbelow tepals. Lowland and
lower montaneslopes on Pacific side of Andes in
Ecuadorand Colombia,Panama..................... 10. E. browniana
49. Receptaclegraduallymerging with tepals.
Antioquia,Colombia ......... ............ 12. E. colombiana
38. Branchletsmidway along flush densely pubescent,the surfacebarely or not visible.
50. Basalmost secondariesasymmetric,one or both merging with leaf marginnear
base or emerging fromjunction of midriband leaf base.
51. Flowers campanulate,the tepals erect at anthesis, glabrousinside. Stamens
included. CentralAmazonia to E Andes........................................ 8. E. metallica
51. Flowers rotateor hypocrateriform,the tepals spreadingat anthesis, densely
pubescentinside. Stamensexserted.
52. All antherstwo-locellate;whorls I and II stamens stipitatewith S-curved
filaments.W Amazonia to E Andean slopes .................................... 53. E. klugii
52. Whorl III anthersfour-locellate;whorls I and II stamenssessile,
petaloid. Manaus,Brazil..................................... Rhodostemonodaphnerecurva
50. Basal secondaries ? symmetric,the basalmostpairs emerging from midribabove
leaf base.
53. Leaves triplinerved.The lowermostpair of secondariessuboppositeshortly
above the leaf base, the others emerging after midlaminaor absent.
54. Anthersof whorl I & II stamenstruncateto emarginateabove. W
Amazonia.................................... 58. E. acuminata
54. Anthersof whorl I & II stamens apiculate,the connective prolonged
above. E Andes to W Amazonia..................................... 59. E. krukovii
53. Leaves penninerved.The secondaries ? uniformlyspaced along midrib.
55. Flowers obconical or urceolate,the tepals incurvedor erect at anthesis.
Stamensincluded. Cupules seemingly hemisphericalwhen seen from
outside but actually shallow, the base thickened.W Amazonia to E
Andes..................................... 55. E. robusta
55. Flowers rotateor hypocrateriform,the tepals patentor reflexedat
anthesis. Stamensexserted. Cupules actually deeply hemisphericalor if
shallow, this not due to a thickenedbase.
56. Apex of whorl I & II anthersflat, emarginate,or with a hornlike
lobe above each locule.
57. Tepals chartaceousto membranaceous,sparselypale-pilose
inside. SE Brazil, lower montaneAndes to Panama,W
Amazonia............................. 56. E. paniculata
57. Tepals fleshy, densely grey-tomentoseinside. E Venezuelan
Highlands............................. 60. E. nilssonii
56. Apex of whorl I & II anthersapiculate,prolongedbetween locelli.
58. Whorl I & II stamens stipitate,the filamentsnarrowerthan
anthers,S-shaped,tomentose.E of Andes in South America
................................................................................................. 53. E. klugii
58. Whorl I & II stamens sessile, the filamentsas broadas
anthers,densely papillose. W of Andes from Ecuadorto
Panama................................................................. 10. E. browniana
37. Indumenton leaves below either erect or at least ascendingat angle of ca. 300.
59. Indumenton branchletsmidway along flush sparse and scattered,the surfaceclearly
visible.
60. Leaves triplinerved,the lowermostpair of secondariessuboppositeshortly above
the leaf base, the others appearingafter midlaminaor absent.
61. Tertiaryand higher-orderveins immersedabove. Secondaryvenation
brochidodromous.Innersurfaceof tepals and filamentsof outer whorls of
stamenspubescent.Guianasto E Andes...................................................... 57. E. gracilis
61. Tertiaryand higher-orderveins prominentabove. Secondaryvenation
eucamptodromous.Inner surfaceof tepals and filamentsof outer whorls of
stamensglabrous.W Amazonia............................. ..................... 58. E. acuminata
SYSTEMATICTREATMENT 21

60. Leaves penninerved,the secondaryveins ? uniformlyspaced along midrib.

62. Axils of secondaryveins below with barbellatetufts of hairs. Apex of outer
anthersapiculate.Cajamarca,Peru ................................. .............. 18. E. tomentosa
62. Axils of secondaryveins below withoutbarbellatetufts of hairs. Apex of
outer anthersrounded,truncate,or emarginate.
63. Leaves densely pubescentbelow. Outerstamens stipitate,the filaments
distinct;locelli filling anthers.SE Brazil, lower montaneAndes to
Panama,W Amazonia ............. ............................... 56. E. paniculata
63. Leaves subglabrousbelow. Outerstamenspetaloid, sessile, the filaments
as wide as or wider than anthers;locelli minute, occupying only upper
half of anthers.
64. Branchletsgreenishyellow; leaves oblong with caudatetips.
Secondariesloop-connected.Centralto E Amazonia .......... 6. E. longicaudata
64. Branchletsdarkbrown;leaves obovate to elliptic, tips acuminate.
Basal secondariesending freely, the distal pairs weakly loop-
connected.W Amazoniato E Andes ........................................ 4. E. rubriflora
59. Indumenton branchletsmidway along flush dense, the surfacebarely visible or
concealed.
65. Laminanarrowlyelliptic or lanceolate,ca. 10 times longer than broad,strongly
bullate. Secondariesmore than 50. Hairson all surfacesca. 2 mm long. SW
Amazonia................................................... 3. E. bullata
65. Laminaovate to obovate, less than 3 times longer than broad,plane. Secondaries
less than 15. Hairs on all surfacesless than 1 mm long.
66. Leaf surface areolateabove, the depressionscorrespondingto the fourth-
orderreticulations.All antherstwo-locellate.
67. Branchletsreddishtomentose. Innertepals densely pubescent,the
surface not visible. Tepuisof Venezuelaand Colombia...................41. E. vinotincta
67. Branchletsgreenish yellow tomentose. Innertepals sparselypubescent,
the surfaceclearly visible. Andes of Ecuadorand Peru.................... 42. E. oreocola
66. Leaf surface smooth or minutelypunctulateabove. If with areolate
depressions,then at least whorl III anthersfour-locellate.
68. Flowers globose, the tepals erect or inflexed at anthesis. Stamens
included.Amazonia, Guianas,E Andes .................................... 43. E. szyszylowiczii
68. Flowers campanulateto rotate,the tepals ascendingto spreadingat
anthesis. Stamensexserted.
69. Whorl I and II antherstruncate,roundedor emarginateabove;
connectives broad above, level with, or reducedbetween
suborbicularlocelli.
70. Outerstamens stipitate,the filamentsmuch narrowerthan
anthers,locelli large, filling anther.
71. Leaves triplinerved,the lowermostpair of secondaries
suboppositeshortlyabove the leaf base, the others
emerging after midlamina.Guianasto E Andes.......... 57. E. gracilis
71. Leaves penninerved,the secondaries ? evenly spaced.
72. Flowers infundibuliform;tepals ascendingat
anthesis. Cupule marginsentire, the tepals or tepal-
bases not persisting.Pacific coast of Colombia
..........................................................................38 .E. xerampela
72. Flowers rotate;tepals spreadingat anthesis.Cupule
marginslobed, the tepals or tepal-basespersisting.
73. Whorl III anthersfour-locellate.Centralto W
Amazonia.Flooded forest ........................... 35. E. anomala
73. All antherstwo-locellate. SE Brazil, throughout
Andes to Panama,W Amazonia.Terrafirme
................................................................. 56. E. paniculata
70. Outerstamens (sub-)sessile, the filamentsbarely or not
narrowerthan anthers;locelli small, occupying distal half of
anther.
22 FLORA NEOTROPICA

74. Branchletsrusty or grey-tomentose.Tertiariesimmersed

above. Amazon basin ............... ..................... 7. E. sprucei
74. Branchletsrusty tomentellose.Tertiariesraised above.
75. Laminaeplane with flat margins.All antherstwo-
locellate. AmazonianEcuador.......................... 5. E. ferruginosa
75. Laminaewith revolutemargins.Whorl III anthers
four-locellate.FrenchGuiana
..............................................Rhodostemonodaphnerevolutifolia
69. Whorl I and II anthersapiculateabove; connectivesprolonged
between obliquely hemisphericallocelli.
76. Tertiariesimmersedabove. Flowers sparselypubescent
outside, the surface visible. Guianas,W Amazonia,E Andes
.......................................................................................... 37. E. canescens
76. Tertiariesraised above. Flowers densely pubescentoutside, the
surfacebarely or not visible.
77. Leaves obovate, the apex blunt, rounded.Indumenton
branchletsdarkred. W Amazonia and E Andes............. 40. E. rubra
77. Leaves ovate to elliptic-oblong,acuminate.Indumenton
branchletsbrownto yellowish.
78. Branchletstomentellose,the hairs erect. Filaments
of whorl I and II stamensnarrowerthan anthers.W
Amazonianlowlands ............................... 39. E. citriodora
78. Branchletssubsericeous,the hairs appressedto
ascending.Filamentsof whorl I and II stamens
equal anthers.W Amazonia and lower montaneE
Andean slopes ........ 45. E. lorastemon

1. Endlicheria punctulata (Mez) C. K. Allen, Mem.
New YorkBot. Gard. 15: 68. 1966. Ocotea punc-
tulata Mez, Jahrb.Konigl. Bot. Gart. Berlin. 5.
379. 1889. Type. FrenchGuiana.Withoutlocality
and date, Melinon 204 (lectotype, designated by
Allen, 1966: P; isolectotype:P).
Treeletsor shrubsto 4 m. Branchletsslender,mid-
way along flush 1.5-2.5 mm diam., angular,sparsely
strigillose, the surface clearly exposed, reddish
brown,the hairs short,to 0.2 mm, straight,appressed,
greyish white; terminalbuds slender, 3 X 0.6 mm,
silvery sericeous. Leaves alternate,widely andevenly
spaced along currentflush;petioles slender,to 1.5 X
0.1 cm, canaliculate,the indumentas on branchlets;
laminae stiff-chartaceousto coriaceous, plane to sub-
bullate, obovate, 7-11 X 3-5 cm, the base attenuate,
the apex caudatefor up to 1.5 cm, the marginsmin-
utely recurvedthroughout;upper surfaceolive green
to light brown,minutelypunctulate,the midribprom-
inent, the higher-ordervenationimmersed;lower sur-
face glabrous, all vein orders immersed, the midrib
dark,conspicuous againstthe lighter lamina;second-
ary veins 5-6 per side, ? evenly spaced, slightly
more distantaroundmidlamina,ascending at 50-60?
(moreobtusely towardsapex), arcuate,the distalpairs
loop-connected;tertiarieslaxly reticulatingbetween
secondaries. Staminateinflorescencesevenly spaced
along currentflush in the axils of foliage leaves, to 6
cm long with 6 lateral branches,branchorders 2-3,
the highest order dichasial, lax, the flowers distant,
the axes sparsely grey-strigillose;bracts and bracte-
oles caducous by anthesis, triangular,with indument
as on axes; pedicels terete, to 2 mm long, those sup-
porting secondary flowers slightly shorter.Flowers
tubiform,sparselygrey-strigilloseoutside;receptacle
infundibuliform,0.6 x 1 mm, densely rusty tomen-
tose inside. Tepals chartaceous,triangular,0.6 X 0.3
mm, erect, surroundingandroeciumat anthesis, the
outer and inner surfacessparselygrey-strigillose,the
marginsand apex inside sparsely papillose. Stamens
of whorls I and II 0.6 mm tall, stipitate,the anthers
ovate, 0.4 x 0.3 mm, glabrous,the apex rounded,the
connectives broad above the 2 locelli, these suborbi-
cular, introrse, the filaments laminar,narrowerthan
anthers,sparsely grey-pilose;whorl III stamens stip-
itate, 0.7 mm tall, the anthersoblong, 0.4 x 0.3 mm,
erect, locelli 2, extrorse-latrorse,the filaments nar-
rower than anthers, slender, laminar,sparsely grey-
pilose, the basal glands sessile, globose, relatively
large, filling the space between filamentsof innerand
outer whorls; whorl IV wanting;pistillode fusiform.
Pistillate inflorescences unknown. Fruits borne on
claviformpedicels of up to 1 X 0.3 cm; cupuleshem-
ispherical,to 0.5 X 1 cm, glabrousinside andoutside,
the marginsentire;drupesellipsoid, to 1.5 X 1 cm.
SYSTEMATICTREATMENT 23

Distribution (Fig. 4) and ecology. Understory Endlicheria punctulata is distanced from most
treeletsor shrubsof lowlands and lower montanefor- species of Endlicheriaby its concolorousleaves, pau-
ests (200-800 m) in French Guiana, adjacent Suri- ciflorous inflorescences, and distinct filaments in all
name, and the state of Amapd in Brazil. Flowering threestaminalwhorls. This combinationof vegetative
August and September, fruiting material collected and floral features is otherwise only found in E. cor-
from Januaryto October. iacea, from which E. punctulata is readily distin-
guished by the minute pin-prickpatternon its upper
Representative specimens examined. SURINAME.
leaf surface and the immersedvenationbelow. These
Vic. of Ulemarie R. ca. 150 km upstreamfrom confluence
with LitaniR., 175 m, 18 Apr 1998 (fr), Hammeletal. 21557 two species areisolated in Endlicheriaandapparently
(MO). more closely related to species in the Ocotea cernua
FRENCH GUIANA. Saul, Monts La Fumee, 200-400 species group (see Generic Delimitation and Species
m, 25 Sep 1982 (fl 6), Mori et al. 14995 (HBG, NY, US); Groups,above).
Tumac Humac, Mitiraka, 1 Sep 1972 (fl 6), de Granville Mez (1889) cited five fruitingcollections (M6linon
1448 (MO, P); Route de Belizon, MontagueTortue,km 25, 204, 216, 227 & 551, and Jenman 1702) in the pro-
23 Apr 1992 (fr) Acevedo et al. 4834 (F, MO). tologue of the basionymof this species, Ocoteapunc-
BRAZIL. AMAPA:Rio Araguari,Porto Platon, 21 Sep tulata. I
have seen the Paris (P) duplicatesof Me'linon
1961 (fl d), Pires et al. 51168 (B, HBG, K, NY, US); Rio
204, 216, and 551, and the Copenhagen(C) sheet of
Iaue, first cachoeira, 22 Aug 1960 (fl d), Irwin & Westra
47706 (GH, NY). Melinon 227. All are from French Guiana and are
clearly conspecific. However,Jenman1702, collected
Local name. Brazil: louro. in Guyana and seen at BRG, likely deposited also at

-4-------
---------

. . . . . . . . .. .....

4"

10 ~ ~ f

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

-- -- -- -- .. .. .. .

~ ~ ~ -. ' ~?- A
~ ~ ~ ~ ~ ~
A~~~~~~~~~~~~~~~~~~~~~~~~x ~ ~ *~*

FIG. 4. Distributionof Endlicheriapunctulata, E. coriacea, E. bullata, and E. rubriflora.

24 FLORANEOTROPICA

K, belongs to either Ocotea pauciflora or 0. cernua. locelli, these suborbicular,introrse,the filamentslam-

Both are very similar to E. punctulata, but with inar, narrower than anthers, sparsely grey-pilose;
smoothratherthanpunctulateupperleaf surfaces.Al- whorl III stamens stipitate,0.7 mm tall, the anthers
len's (1966) lectotypification on Melinon 204 pre- oblong, 0.4 X 0.3 mm, erect, locelli 2, extrorse-
vents futureconfusion. latrorse,the filamentsnarrowerthan anthers,slender,
laminar,sparselygrey-pilose,the basalglands sessile,
globose, relativelylarge, filling the space between fil-
2. Endlicheria coriacea Chanderbali,sp. nov. Type. aments of inner and outer whorls; whorl IV wanting;
Brazil. Amazonas: Manaus, Agropecuairio,Re- pistillode filiform. Pistillate inflorescencewith indu-
serve 1501 (km 41) of the WWF/INPABiological ment, color, and branchingas in staminateplants,the
Dynamics of ForestFragmentsProject,50-150 m, flowers similar in size and shape; stamens sterile,
21 Sep 1989 (fl d), Lepsch da Cunha et al. 391 smaller; ovary glabrous;style slender, distinct from
(holotype:NY; isotypes: NY, MO). Fig. 5 ovary; stigma tri-lobed, 0.3 mm diam. Maturefruits
Endlicheriae punctulatae affinis, inflorescentiiset flori- unknown, immaturefruits borne on terete pedicels,
bus similis,sed staturapermajoreet foliis subtusnervis the cupules glabrousinside and outside, the margins
prominentibus recedit. entire.
Trees to 30 m. Branchletsslender,midway along Distribution (Fig. 4) and ecology. Tall trees
flush 2-3 mm diam., distally weakly angularsoon te- knownonly fromlowlandforests (50-150 m) on well-
rete, sparsely strigillose, the surfaceclearly exposed, drained sand and clay soils aroundManaus, Brazil.
dark brown, the hairs short, to 0.1 mm, straight,ap- Flowering from July to Septemberwith fruit set ini-
pressed,greyishwhite; terminalbuds slender,3 X 0.6 tiated by September.
mm, silvery sericeous. Leaves alternate,widely and
evenly spaced along currentflush;petioles slender,to Additional specimens examined. BRAZIL. AmA-
1.5 X 0.1 cm, canaliculate,glabrous;laminae coria- ZONAS: Estrada km 135,22 Aug 1973,
Manaus-Itacoatiara,
ceous, plane, obovate, 4-6 X 8-12 cm, the base at- (fl Y), Rodrigueset al. 8523 (HBG,INPA,US), 10Jul1974
tenuate,the apex acute, narrowlyacuminate,or cau- (fl Y),Rodrigues& Loureira9454 (F,INPA,MO),13 Sep
1972 (fr), Pires & Coelho175 (INPA);Reserva2107 do
date for up to 1 cm, the margins minutely recurved
ProjectoDinamicaBiologicados FragmentosFlorestais,
throughout;uppersurfacelight to darkbrown,waxy, et al. INPAI
Arv.No. 388, 30 Jul 1982 (fl Y), Mackenzie
the primaryto fourth-orderveins raised, their prom- WWF2107.388(MO).
inence decreasingwith rank;lower surfaceglabrous,
all vein orders raised, their prominence decreasing Local name. Rodrigues et al. 8523 suggest
with rank; secondary veins 7-9 per side, ? evenly "Louropirarucu"is the local name, but this may have
spaced, slightly more distantaroundmidlamina,pat- been misapplied.Accordingto Vicentiniet al. (1999),
ent, diverging at 70-85? (more obtusely towards "Louropirarucu"refers to Licaria cannella (Meisn.)
apex), arching after midcourse, weakly brochidod- Kosterm.and materialof Endlicheriacoriacea can be
romous; tertiaries once-forked or laxly reticulating easily mistakenfor this species. The strong smell of
between secondaries.Staminateinflorescencesevenly fish from the inner bark of L. cannella, surely re-
spaced along current flush in the axils of foliage sponsible for association with the pirarucu,or ara-
leaves, to 6 cm long with 6 lateralbranches,branch paima,is yet to be reportedfor species of Endlicheria.
orders2-3, the highest orderdichasial, lax, the flow-
ers distant, the axes sparsely grey-strigillose;bracts Endlicheria coriacea is distinguishablefrom E.
and bracteoles caducous by anthesis, triangular,the punctulata by its smooth upper leaf surface, promi-
indumentas on axes; pedicels terete, to 2 mm long, nent venationbelow, and large stature,but flowersof
those supportingsecondary flowers slightly shorter. the two are indistinguishable.Both are apparently
Flowers tubiform, sparsely grey-strigillose outside; two-locellate membersof a species group in Ocotea
receptacle shallowly infundibuliform,0.6 X 1 mm, centeredaround0. cernua(see Relationships,above).
densely silvery pilose inside. Tepalsmembranaceous, Among the treelets and shrubstypical of this group,
ovate, 0.6 X 0.3 mm, ascending,the androeciumsur- E. coriacea and Rhodostemonodaphneelephantopus
rounded at anthesis, the outer and inner surfaces are remarkablefor theirarborescence.These two spe-
sparsely grey-strigillose, the margins sparsely papil- cies differ in locelli number and, with E. coriacea
lose. Stamensof whorls I and 110.6 mm tall, stipitate, known only from centralAmazonia and R. elephan-
the anthersdepressed-ovate,0.4 X 0.5 mm, glabrous, topus from French Guiana, they may be allopatric.
the apex rounded,the connectives broadabove the 2 However,they are vegetativelyindistinguishable.
 FIG. 5. Endlicheria coriacea (Lepsch da Cunha et al. 391). A. Habit. B. Single leaf showing venation. C. Flower
with facing tepal turneddown to reveal androecium.D. Flower ls. E. Whorl I stamen seen from within. F. WhorlIII stamen
seen from without.
26
3. Endlicheria bullata Ducke, Arch. Jard.Bot. Rio
de Janeiro4: 190. 1925. Type. Brazil. Amazonas:
Bom Logar,25 Jul 1903 (fl d), Hubers.n. R 18359
(holotype: R-n.v.; isotypes: B-n.v., U).
Shrubsto 6 m. Branchletsslender,midway along
flush 2-3 mm diam., terete, densely rusty to reddish-
hirsute, the surface barely visible throughindument
cover, the hairs long, to 2.5 mm, straight, rigidly
erect;terminalbudsdensely rustypubescent,the hairs
as on branchlets,ascending.Leaves alternate,widely
and evenly spaced along currentflush; petioles slen-
der, to 1 X 0.1 cm, terete, the indumentas on bran-
chlets; laminae chartaceous,bullate, lanceolate, 15-
35 X 3-4 cm, the base acute to rounded, the apex
acute, acuminatefor up to 2 cm, the marginsminutely
recurvedthroughout;uppersurfacelight green,waxy,
the midribprominulous,the secondariesflat, the ter-
tiaries immersed;lower surfacedensely rustyhirsute,
the hairs as on branchlets,denser on main veins, all
vein ordersraised, their prominencedecreasingwith
rank;secondary veins numerous,20-35 per side, +
evenly spaced, slightly more distantaroundmidlam-
ina, divergingat 850, abruptlyascendingnearmargin,
brochidodromous;tertiaries laxly reticulating be-
tween secondaries. Staminateinflorescences evenly
spaced along current flush in the axils of foliage
leaves, to 8 cm long with 8 lateralbranches,branch
orders3-4, the highest orderdichasial,lax, the flow-
ers distant,the axes densely rusty hirsute;bractsand
bracteoles caducous by anthesis, lanceolate, the in-
dument as on axes; pedicels terete, to 5 mm long,
those supportingsecondary flowers slightly shorter.
Flowers rotate,8 mm diam., rustytomentoseoutside,
the indumentsparsertowardstepals;receptacleinfun-
dibuliform,2 X 2 mm, rustyvelutinousinside. Tepals
fleshy, obovate, 3 X 2 mm (the inner whorl slightly
narrower),spreadingat anthesis,the innersurfacegla-
brous. Stamens of whorls I and II sessile, stout, 1.3
X 1.3 mm, the anthersdepressed-ovate,0.6 X 1 mm,
glabrous, the apex rounded,the connectives slightly
incurved over the 2 locelli, these suborbicular,just
below apex, minute, introrse, the filaments fleshy,
broaderthananthers,the base rustytomentose;whorl
III stamens stout, sessile, 0.8 mm tall, the anthers
depressed-oblong, 0.4 X 1 mm, erect, locelli 2,
extrorse-latrorse,the filamentsbroaderthan anthers,
fleshy, the base rustytomentose,the basal glands ses-
sile, relatively large, globose-apiculate; whorl IV
wanting; pistillode fusiform. Pistillate inflorescence
not seen; detachedpistillateflowerssimilarin size and
shape to staminateflowers; stamens sterile, smaller;
ovary glabrous, ovoid; style stout, weakly distin-
FLORA NEOTROPICA
guished from ovary; stigma discoid, 0.8 mm diam.
Fruitsunknown.
Distribution (Fig. 4) and ecology. Small shrubs
from terra firme forest of SW Amazonia in Brazil.
Flowering in July and September.
Additionalspecimensexamined.BRAZIL. ACRE:
NearmouthRio Macuahan of Rio Yaco),3 Sep
(tributary
5780(A, G, K, S, U, US).
1933(fl d & Y?),Krukoff
Endlicheriabullata is a poorly known but highly
distinctive species. Nowhere else in Endlicheria are
narrowbullateleaves with numeroussecondariesand
stiff hirsute vestiture combined with rotate flowers
and fleshy sessile stamens. Similar flowers combine
with plane laminae in E. ferruginosa and E. rubri-
flora, while similarleaves and vestiturecombine with
campanulateflowersand stipitatestamensin E. glom-
erata. However,outside of Endlicheriathese charac-
ters all appear in Rhodostemonodaphnescandens
Madrinian,albeit with four-locellateanthers.
Although only two collections are cited here (the
type and Krukoff5780), they representthreeindivid-
uals. Type material is clearly staminate since the
protologue is based on a staminate plant, and at
least the isotype deposited in Utrecht(U) shows sta-
minate inflorescences attachedto twigs. Of Krukoff
5780, sheets at Geneva (G), Stockholm (S), and
Utrecht(U) lack inflorescences,while that at the Ar-
nold Herbarium(A) shows intact staminateinflores-
cences, and the Kew (K) sheet has detached stami-
nate flowers. However, the US duplicate of Krukoff
5780 has detached pistillate flowers that appear to
be con-specific with staminateones. Since monoecy
is unknownin Lauraceae,the source of these pistil-
late flowers is unknown unless Krukoff5780 is a
mixed collection.
4. Endlicheria rubriflora Mez, Jahrb.Konigl. Bot.
Gart.Berlin 5: 494. 1889. Type. Colombia.With-
out locality and date, Triana1032 (holotype:P).
AnibareticulataA. C. Sm.,Bull.TorreyBot. Club58:
99. 1931.Type.Peru.Loreto:Alto Amazonas,Yu-
rimaguas, lowerRioHuallaga,138m,22 Aug-9Sep
1929 (fl Y), Killip & Smith 28050 (holotype: NY;
isotype: US).
Endlicheria trianae 0. C. Schmidt, Repert. Spec. Nov.
Regni Veg. 31: 175. 1933. Type. Colombia. Andes
de Antioquia,Triana2040-1 (holotype:K-n.v.).
EndlicheriawurdackianaC. K. Allen, Mem. New York
Ama-
Bot. Gard.10(5):67. 1964.Type.Venezuela.
zonas:RioSiapabetweenRaudalGallinetaandSalte
Gallineta, 130-150 m, 23 Jul 1959 (fr), Wurdack&
Adderley43592 (holotype:NY; isotypes: GH, U).
SYSTEMATICTREATMENT
Trees to 15 m. Branchletsslender,midway along
flush 2-3 mm diam., distally weakly angular,soon
terete, sparsely pubescent, the surface clearly ex-
posed, dark to reddish brown, the hairs relatively
short, to 0.3 mm, straight, appressed to ascending,
yellowish green; terminalbuds slender,2 X 0.6 cm,
densely pubescent, the hairs as on branchlets, ap-
pressed. Leaves alternate,widely and evenly spaced
along currentflush;petioles slender,to 1.5 X 0.2 cm,
terete, the indumentas on branchlets;laminae char-
taceous, plane, elliptic to obovate, 7-20 X 3-8 cm,
the base obtuse to rounded,briefly attenuate,the apex
obtuse to acute, acuminatefor up to 2 cm, the margins
flat throughout;upper surface greyish green, waxy,
the midribsunkentowardspetiole, otherwiseprimary
to fourth-orderveins raised, their prominence de-
creasing with rank;lower surfacesparselypubescent,
the hairs appressed, ascending, or erect, slightly
denser on main veins, all vein orders raised, their
prominencedecreasingwith rank;secondaryveins 6-
9 per side, + evenly spaced, slightly more distant
around midlamina, patent, diverging at 70-85O (more
obtusely towards apex), arcuate, distal pairs loop-
connected; tertiaries laxly reticulatingbetween sec-
ondaries. Staminate inflorescences evenly spaced
along currentflush in the axils of foliage leaves, to
10 cm long with 8 lateralbranches,branchorders3-4,
the highest order dichasial, lax, the flowers distant,
the axes sparselypubescent,the hairsgreenishyellow
to grey, ascending, denser on pedicels; bracts and
bracteoles caducous by anthesis, lanceolate, densely
pubescent, the hairs as on axes, appressed;pedicels
terete, to 2 mm long, those supporting secondary
flowers slightly shorter.Flowersbroadlyrotate,5 mm
diam., sparingly rusty strigillose outside; receptacle
shallowly infundibuliform,0.6 X 1.3 mm, densely
yellowish tomentose inside. Tepals fleshy, broadly
ovate, 2 X 1.6 mm, spreadingto recurvedat anthesis,
the innersurfaceandmarginsdensely rustyto reddish
papillose. Stamensof whorlsI and II broadlystipitate,
0.8 m tall, the anthersreniform,0.6 X 0.8 mm, whorl
II slightly narrower,glabrous,the apex rounded,the
connectives slightly incurvedover the 2 locelli, these
suborbicular,just below apex, minute, introrse, the
filamentslaminar,0.2 X 0.6 mm, glabrous;whorl III
stamens stout, sessile, 0.6 mm tall, the antherstrans-
versely oblong, 0.3 X 0.6 mm, erect, locelli 2,
extrorse-latrorse,the filamentsas broadas or broader
thananthers,columnar,glabrous,the basalglandsses-
sile, globose, relatively large, filling the space be-
tween inner and outer whorls of stamens; whorl IV
wanting; pistillode filiform. Pistillate inflorescence
with indumentand color as in staminateplants, but
27
shorterand with fewer lateral branches,the flowers
slightly deeper; stamens sterile, smaller, the anthers
auriculate;ovary glabrous,ovoid; style stout, weakly
distinguishedfrom ovary; stigma almost sessile, dis-
coid, 0.6 mm diam. Fruits borne on stout claviform
pedicels of up to 3 X 0.5 cm; cupules discoid to re-
flexed, to 4 cm diam., glabrousoutside andinside, the
marginslobed; drupesellipsoid, to 3 X 2 cm.
Distribution (Fig. 4) and ecology. Small trees
from westernAmazonianlowlands and adjacenteast-
ern Andean slopes at 100-1900 m. Found in both
flooded and nonfloodedsoils. Flowering and fruiting
throughoutthe year.
Local names. Colombia:jigua de mierda. Peru:
palta amarilla,roble palta, roblecillo puchirin.
Representative specimens examined. COLOMBIA.
ANTiOQUIA: San Luis, AutopistaMedellin-Bogota, 790 m,
15 Jan 1983 (fl d), Cogollo & Brand 394 (MO). CAUCA:
El Tambo,VeredaLa Romelia, ParqueNaturalMunchique,
23 Aug 1994 (fr), Jarvenpaa27 (COL).META: Villavicen-
cio, Andes de Bogota, Jan 1856 (fr), Triana2060 (K).QUIN-
DIO: Genova, 1870 m, 17 Mar 1988 (fl d), Arbelaez et al.
2490 (HBG, MO). TOLIMA: Mariquita,near Hondaon Rio
Magdalena,s.d. (fr), Karstens.n. (LE).VALLE: Sevilla, via
Sevilla-La Raquelita,a orilla de la quebradala Raquelita,
1500 m, 31 Jul 1985 (fr), Devia 1052 (MO).
VENEZUELA. APURE: Reserva Forestal San Camilo,
280-300 m, 27 Mar 1968 (fr), Steyermarket al. 101408
(HBG, NY). BARINAS: Pedraza,El Velador,14 Dec 1954 (fl
Y), Bernardi1789 (G, NY).
ECUADOR.NAPO: Tena,Estaci6nBiol6gica JatunSa-
cha, 400 m, 22 Sep 1989 (fl d), Palacios 4473 (G, MO,
NY). ZAMORA-CHINCHIPE:Zamora,JamboeBajo, E bor-
der of PodocarpusNational Park, 1100 m, 5 Nov 1996 (fl
6), Clarkeet al. 3274 (MO).
PERU.MADRE DE DIOs: Tambopata, Cuzco Amaz6n-
ico, 200 m, 14 Jun 1989 (fl 6), Nuiiez et al. 10804 (MO).
PASCO: Oxapampa,Iscozacin, 26 Jun 1986 (fl Y), Pariona
& Sebastian46 (F, MO); Palcazd, 300-600 m, 18 Nov 1985
(fr), Hartshornet al. 2827 (MO).
BRAZIL.AMAZONAS:Humaitd,PortoVelho, km 30 da
estradaHumaita-Labrea, 10 Jun 1982 (fl Y), Filho 82-7 N
(HBG); Rio Curuquete,halfway between Cachoeiras Sao
Paulo and Republica, 22 Jul 1971 (fl Y), Prance et al. 14498
(HBG, K, MO, NY).
Endlicheria rubriflora has green-drying leaves
similar to those of E. longicaudata and E. pyriformis,
but its rotate flowers with densely papillose tepals
arounda centralhub of large globose glands andred-
dish stamens are unmistakable.The cupules of this
species arealso unmatchedby congeners.The cupules
appearto start off as shallow patelliformstructures,
and as fruits maturethey flatten to a thick disc and
ultimately the margins become recurved,becoming
28 FLORA NEOTROPICA

umbrellalikein aspect. Similar cupules are produced
by Rhodostemonodaphne antioquensis(see also Mad-
rinain,1996b), the flowers of which would also be
identicalbut for the four-locellateanthers.Besides the
added pair of locelli in R. antioquensis, its ovate
leaves with a more coriaceoustexturehelp to separate
it from E. rubriflora.
Lowland Amazonian collections, including the
type of Aniba reticulata,have slightly smaller flow-
ers, but otherwise Endlicheria rubriflorais morpho-
logically uniform.The type of E. trianae is not avail-
able to me, but since Smith's (1933) descriptiondoes
not provide distinguishingcharacters,I follow Kos-
termans's(1937) synonymy underE. rubriflora.The
third synonym accepted here, E. wurdackiana, is
based on a fruitingspecimen that has the cupule and
vegetative charactersof E. rubrifiora.
5. Endlicheria ferruginosa Chanderbali,sp. nov.
Type. Ecuador. Napo: La Joya de los Sachas,
Parque Nacional Yasuni, Maxus highway and
pipeline under construction, km 27, 230 m, 16
Aug 1993 (fl d), Dik 203 (holotype: MO; isoty-
pes: COL, MO, QCNE). Fig. 6
branches,branchorders3-4, the highest orderdicha-
sial, lax, the flowers distant, the axes sparsely pale
rusty tomentellose, distalmostbranchesand pedicels
more densely so; bracts and bracteolescaducous by
anthesis, lanceolate, rusty sericeous; pedicels terete,
to 4 mm long, those supportingsecondary flowers
slightly shorter.Flowers rotate,5 mm diam., sparsely
rusty strigillose outside; receptacle shallowly infun-
dibuliform, 1 X 2 mm, densely rusty tomentose in-
side. Tepals fleshy, ligulate, 2 X 2 mm, spreadingto
recurved, the androecium exserted at anthesis, the
outer surface sparselyrusty strigillose, the inner sur-
face more densely so, the margins and apex inside
rustypapillose. Stamensof whorls I and 11I1mm tall,
sessile, the antherstransverselyoblong, 0.6 X 0.8 mm
(whorl II slightly smaller), sparsely minutely papil-
lose, the apex truncate,the connectiveslevel with the
2 locelli, these minute, suborbicular,introrse,the fil-
aments clavate, slightly narrowerthan anthers,rusty
tomentose;whorl III stamenssessile, 1.3 mm tall, the
anthers depressed-oblong,erect, locelli 2, extrorse-
latrorse,the filamentsbroaderthananthers,columnar,
glabrous,the basal glands sessile, globose, apiculate,
filling the space between inner and outer whorls of
stamens;whorl IV wanting; pistillode wanting. Pis-
tillate plants unknown.

A speciebusquasKostermans sectionisMicrolocellatae Distribution (Fig. 7) and ecology. Known only

ascripsitramulisferrugineo-tomentellisdistinguenda. from the type materialtakenfrom a small tree of terra
firme lowland forest in Amazonian Ecuador found
Trees to 12 m. Branchletsslender,midway along floweringin August.
flush 2-3 cm diam., distally weakly angular,soon te-
rete, densely tomentellose, the surface concealed by The ratherlarge rotate flowers with fleshy tepals
the indument cover, the hairs short, to 0.1 mm, and sessile stamenswith minutelypapilloseindument
straight,erect, rust brown;terminalbuds plump,4 X are all characteristicof Rhodostemonodaphne,but
2.5 mm, rusty sericeous. Leaves alternate,widely and two-locellate anthers assign this species to Endli-
evenly spaced along currentflush;petioles slender,to cheria. Therein, similar flowers are found in E. bul-
1.5 X 0.15 cm, semi-terete,the indumentas on bran- lata and E. rubriflora, from both of which E.
chlets; laminaechartaceous,plane, obovate, 12-17 X ferruginosa can be easily distinguishedby its rusty
3-6 cm, the base acute to cuneate, briefly decurrent, tomentellosebranchlets.As in those species, the tran-
the apex obtuse, acuminatefor up to 2 cm, the mar- sition to two-locellateanthersfromthe ancestralfour-
gins flat; upper surface olive-brown,minutely punc- locellate condition is complete but in other respects
tulate, the primaryto fourth-orderveins raised, their the androeciumis best matchedby membersof Rho-
prominence decreasing with rank; lower surface dostemonodaphne.In the case of Endlicheriaferru-
sparsely tomentellose, the hairs as on branchlets,ap- ginosa it is Rhodostemonodaphnerevolutifoliathat
pressed and denser on main veins, all vein orders comes closest. In staminateflowersof the latter,whorl
raised, their prominence decreasing with rank; sec- I and II anthersaretwo-locellateandbroadlypetaloid,
ondaryveins 5-7 per side, ? evenly spaced, slightly muchlike those of Endlicheriaferruginosa,butwhorl
more distantaroundmidlamina,ascending at 50-60O III anthersare four-locellate,and in pistillate flowers
(more obtusely towards apex), arcuate, distal pairs all anthersarefour-locellate(Madrifinan, 1996a).Even
loop-connected; tertiaries roughly horizontal, be- if pistillate plants of E. ferruginosa turn out to be
tween secondariesstraightto once-forked.Staminate four-locellate,this species has alreadyadvancedfur-
inflorescences evenly spaced along current flush in ther in locelli reduction than Rhodostemonodaphne
the axils of foliage leaves, to 6 cm long with 8 lateral revolutifolia,and more likely in parallelthan from a
SYSTEMATICTREATMENT 29

co

FIG.6. Endlicheniaferruginosa(Dik 203). A. Habit.B. Single leaf showing venation.C. Leaf base below. D. Rlower.
E. Rlower15. F. Whorl I stamen seen from within. G. Whorl III stamen seen from without.
30
W,
~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~
4.
.......... ----------
~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
- r~~~~~~~~~~~~'~Y
ofEndlicheriaferruginosa,E longicaudata and E
FIG.7. Distribution
common ancestor.The indumentof R. revolutifoliais
much longer and more yellow in color, and the rev-
olute leaves to which its name alludes contrastwith
the plane laminae of Endlicheriaferruginosa.
6. Endlicheria longicaudata (Ducke) Kosterm.,Re-
cueil Trav. Bot. Neerl. 34: 515. 1937. Hubero-
daphne longicaudata Ducke, Arch. Jard.Bot. Rio
de Janeiro4: 191. 1925. Type. Brazil. Para:Rail-
roadbetween Belem and Braganca,near SantaIz-
abel, Jun 1908 (fl cd), Huber s.n. MG 9431 (lec-
totype, designated by Kostermans,1937: R-n.v.;
isolectotypes: S, U).
Treeletsor shrubsto 5 m. Branchletsslender,mid-
way along flush 1-1.5 mm diam., at first sharplyan-
gular, soon terete, sparsely pubescent, the surface
clearly exposed, light or yellowish green, the hairs
short,to 0.2 mm, straight,erect to ascending, pale or
yellowish brown; terminalbuds slender, 2 X 1 mm,
FLORA NEOTROPICA
-
K~~~~~~~~~A
W
~ ~ ~~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ I"
, .. .
sprucei~~~~~~~~~~~~~~~~~~~
densely pubescent, the hairs yellowish green, ap-
pressed. Leaves alternate,widely and evenly spaced
along currentflush; petioles slender, to 2 X 0.2 cm,
terete, the indument as on branchlets;laminae char-
taceous, plane to subbullate, oblong, 10-15 X 4-7
cm, the base acute, attenuate,the apex obtuse, then
abruptlycaudatefor up to 4 cm, the marginsminutely
recurvedthroughout;uppersurfacedeep green, waxy,
the primaryto fourth-orderveins raised, their prom-
inence decreasing with rank, creamish yellow, con-
spicuous against the lamina; lower surface sparsely
pubescent, the hairs scattered,erect, persisting only
on the midvein, all vein orders raised, their promi-
nence decreasingwith rank;secondaryveins 4-5 per
side, ? evenly spaced, slightly more distant around
midlamina,patent,divergingat 70-85' (more acutely
towards apex), abruptly ascending after midcourse,
brochidodromous; tertiaries laxly reticulating be-
tween secondaries. Staminate inflorescences evenly
spaced along current flush in the axils of foliage
SYSTEMATICTREATMENT
leaves, to 5 cm long with 4 lateralbranches,branch
orders2-3, the highest orderdichasial, lax, the flow-
ers distant, the axes sparsely grey-tomentellose,the
hairs 0.1 mm long, scattered, erect to ascending;
bractsandbracteolespersistentat anthesis,lanceolate,
sparsely grey-strigillose, the hairs as on axes, ap-
pressed;pedicels graduallyincreasingin diameterap-
ically, to 2 mm long, those supporting secondary
flowers slightly shorter. Flowers rotate, to 5 mm
diam., sparsely grey-strigillose outside; receptacle
shallow,infundibuliform,0.5 X 1 mm, densely tawny
tomentose inside. Tepals chartaceous,ligulate, 3 X
1.2 mm (the inner whorl slightly broader),horizon-
tally spreading,the inner surface glabrous, the mar-
gins and apex inside lightly papillose. Stamens of
whorls I and II ligulate, sessile, 1 X 0.6 mm, the an-
thersminute,transverselyoblong, 0.1 X 0.3 mm, ster-
ile (lacking) in whorl I, glabrous,the apex retuse,the
connectivesreducedbetween the 2 locelli, these min-
ute, introrse,suborbicular,the filamentsligulate, gla-
brous; whorl III stamens slender,columnar,1 X 0.3
mm, the anthers0.3 X 0.3 mm, depressed-triquetrous,
incurved,locelli 2, minute, introrse-latrorse,the fila-
ments as broad as anthers,triquetrous,glabrous,the
basal glands absent; whorl IV wanting; pistillode
wanting.Pistillateinflorescencewith indument,color,
and branchingas in staminateplants,the flowerssim-
ilar in size and shape;stamens sterile, smaller;ovary
glabrous,ovoid, 0.6 X 0.5 mm; style slender,distinct
from ovary, about 0.6 mm long; stigma strongly tri-
lobed, 0.7 mm diam., spreadingabove sterile androe-
cium. Fruitsborne on claviformpedicels of up to 2.5
X 0.5 cm; cupules patelliform,to 0.3 X 1 cm, gla-
brous outside, sparsely grey-strigoseinside, the mar-
gins lobed, tepals persisting;drupesellipsoid, to 2.5
X 1 cm.
Distribution (Fig. 7) and ecology. Small treelets
or shrubs from terra firme lowland forests (50-200
m) in centralto easternAmazonia.FloweringJuneto
October,fruitingAugust to June. 7. Endlicheria sprucei (Meisn.) Mez, Jahrb.Konigl.
Representative specimens examined. BRAZIL.
AMAPA: Canteiro de obrasdeHidroeletrica deBalbina,Mar
1986 (fr), Cid Ferreira et al. 6713 (INPA, K, MO, NY).
AMAZONAS:Vic. of Rio Uatuma,Rio Pitinga,Aug 1979 (fl
6), Cid Ferreira et al. 881 (F, HBG, INPA, MG, NY, R,
US); Tapuruquara,Oct 1971 (fl d), Prance et al. 15817
(HBG, INPA, MG, NY). PARA:Belem, Bosque Municipal,
Jun 1943 (fl 6), Ducke 1232 (MG, MO, NY, US); lands of
InstitutoAgron6mico do Norte, Jul 1944 (fl i), Silva 282
(F, K); Tome, Aqu, Rio Acara, Jan 1978 (fr), Nascimento
386 (F, HBG, MG, NY). RONDONIA: Sao Loren,o mines,
Nov 1968 (fr), Prance et al. 8892 (HBG, INPA, NY, US);
31
trail N of Rio Madeira from 2 km below confluence with
Rio Abuna,Nov 1968 (fr), Prance et al. 8343 (INPA,NY).
Local names and uses. Called"azywa'yw-pihun"
by the Tembe Indiansof Brazil. The wood is used to
make axe handles and chairs and as a firewood.
Endlicheria longicaudata was described as the
only species of Huberodaphne,a genus Ducke (1925)
separatedfrom Endlicheriaby lack of basal glands in
whorl III stamensand shallow cupules with persistent
tepals. After Kostermans's (1937) treatment these
charactersappearin other species of Endlicheria,but
the androeciumof E. longicaudatais still unmatched
therein.Its sessile outer stamens with small, distally
positionedlocelli are much thinnerand morepetaloid
than others of the Microlocellata species group. In-
deed, in some staminateflowers,whorl I stamenscan
be sterile with empty shallow locelli. Whorl III sta-
mens are also remarkable.These are fleshy, account-
ing for most of the androecialtissue, and anthersare
incurved and locelli introrse-latrorse.This androe-
cium is reminiscent of Dicypellium and Phylloste-
monodaphne. However these genera have bisexual
flowers and double-rimmedcupules, and are closely
related to Licaria (Chanderbaliet al., 2001), while
ITS sequence dataplace Endlicherialongicaudatain
the Endlicheria-Rhodostemonodaphne alliance (Fig.
2). Therein, close relationshipwith Rhodostemono-
daphne scandens lacks statisticaland morphological
support.However, R. parvifolia Madriinanhas shal-
low cupules with persistenttepals and rotate flowers
with greenishsessile stamensvery similarto those of
Endlicherialongicaudata.Interestingly,both species
arepartof a weakly supportedclade (Fig. 2), also with
Rhodostemonodaphnescandens, Endlicheriapyrifor-
mis, and E. citriodora, which, given the morpholog-
ical similarities between E. longicaudata and Rho-
dostemonodaphneparvifolia, may have phylogenetic
significance.
Bot. Gart.Berlin 5: 125. 1889. Goeppertiasprucei
Meisn., DC. Prodr.15(1): 172. 1864. Type.Brazil.
Amazonas:Rio Uaupes, Nov 1852 (fl d), Spruce
2769 (lectotype,designatedby Kostermans,1937:
K; isolectotypes:B-n.v., BM-n.v., E, G, K, NY-
n.v., OXF-n.v., P).
Small trees or shrubs,3-7 m. Branchletsslender,
midway along flush 2-3 mm diam., distally weakly
angular,soon terete, densely greyish green to rusty
hirsute,the surfacebarely visible, the hairsrelatively
long, to 0.6 mm, straight,erect;terminalbuds plump,
32
1 X 0.6 mm, densely pubescent, the hairs as on
branchlets, ascending. Leaves alternate,widely and
evenly spaced along currentflush;petioles slender,to
1 X 0.3 cm, terete, the indument as on branchlets;
laminaechartaceous,plane, elliptic to obovate, 11-23
x 4-8 cm, the base obtuse, or acute, the apex acute,
narrowlyacuminatefor up to 5 cm, the marginsmin-
utely recurved throughout;upper surface dull grey,
minutelypunctulate,the midribprominulous,the sec-
ondaries flat, the tertiariesimmersed,inconspicuous
againstthe lamina;lower surfacesparselyhirsute,the
hairs as on branchlets,slightly denser on main veins,
all vein orders raised, their prominence decreasing
with rank; secondary veins 4-6 per side, ? evenly
spaced, slightly more distant aroundmidlamina,as-
cending at 50-60o (moreobtusely aroundmidlamina),
arcuate,brochidodromous;tertiarieslaxly reticulating
between secondaries.Staminateinflorescencesevenly
spaced along current flush in the axils of foliage
leaves, to 8 cm long with 5 lateralbranches,branch
orders2-3, the highest orderdichasial, lax, the flow-
ers distant, the axes sparsely rusty to grey-hirsute;
bracts and bracteoles caducous by anthesis, lanceo-
late, densely pubescent, the hairs as on axes, ap-
pressed; pedicels terete, to 1.3 mm long, those sup-
porting secondary flowers slightly shorter.Flowers
rotate, 3 mm diam., densely grey-pubescentoutside,
the hairs ascending; receptacle infundibuliform,0.5
X 0.6 mm, densely grey-tomentose inside. Tepals
chartaceous,ligulate, 1.2 X 0.6 mm (the inner whorl
slightly broader), the inner surface densely grey-
strigose, the indument soon restricted to the base,
eventuallylost, the marginsand apex inside minutely
papillose. Stamensof whorls I and II sessile, 0.7 mm
tall, the antherstransverselyoblong, 0.4 X 0.5 mm,
glabrous,the apex truncate,the connectiveslevel with
the 2 locelli, these suborbicular,latrorse-introrse,the
filamentsfleshy, as broadas anthers,the base sparsely
grey-pilose; whorl III stamens sessile, 0.8 mm tall,
the anthersdepressed-oblong,0.3 X 0.4 mm, slightly
bowed towards the outer whorls, locelli 2, extrorse-
latrorse,the filamentsas broadas anthers,clavate,ta-
pering towards base; sparsely grey-pilose, the basal
glands sessile, globose; whorl IV wanting;pistillode
wanting.Pistillateinflorescencewith indument,color,
and branchingas in staminateplants,the flowerssim-
ilar in size and shape; stamens sterile, smaller;ovary
glabrous, ovoid; style slender, distinct from ovary;
stigmabroadlytri-lobed, 1 mm diam. Fruitsborneon
claviform pedicels of up to 1.5 X 0.1 cm; cupules
hemispherical, to 1 X 1.5 cm, glabrous inside and
outside, the margins sharplylobed; drupesellipsoid,
to 2 X 1 cm.
FLORANEOTROPICA
Distribution (Fig. 7) and ecology. Small trees or
shrubs from nonflooded forests throughoutlowland
Amazonia (50-300 m). Flowering from May to No-
vember with fruits maturingby Novemberand avail-
able to July of the following year.
Representativespecimensexamined.COLOMBIA.
CAQUETA: Araracuara,5 Nov 1991(fl d), Duivenvoorden
738 (MO); Morelia, 150-300 m, 19 Oct 1941 (fl d), Snei-
dern 1191 (COL).
ECUADOR.PASTAZA: Lorocachi,200 m, 26 May 1980
(fr), Jaramilloet al. 31147 (K,MO, NY).SUCUMBiOS:Lago
Agrio,ReservaFaunfstica
Cuyabeno,Chiritza,230 m, 13
Nov 1991 (fl d), Palacios et al. 8845 (MO, NY).
PERU.LORETO:Maynas,Mishuyacu, nearIquitos,100
m, Oct-Nov1929(fl i), Klug272 (F, NY,US); Estaci6n
Biol6gicaRfoBlanco,150m, 16 Sep 1985(fr),Vdsquez
et
al. 6742 (F,MO,NY);Rio Nanay,Mishana,80-110 m, 30
Sep 1990 (fl d), Pipoly et al. 12633 (MO).
BRAZIL.AcRE: Cruzeirodo Sul, Estradado Ale-
manha, 4 Nov 1966 (fr), Prance et al. 3008 (HBG, INPA,
NY);Rio Jurua& Rio Moa,Serrade MoaVillage,25 Apr
1971 (fr), Prance et al. 12102 (HBG,INPA,NY). AMA-
ZONAS:Agropecuario, FazendaDimonaof WWF/INPA
MCSProject,50-125 m, 23 Oct 1988(fl d), Boom& Pa-
checo 8511 (INPA,MO);Humaita,EstradaHumaita-La-
brea,km70, 15Jun1982(fl Y), Teixeiraet al. 1025 (INPA,
K, MG, MO, NY). MATO GROSSO:Novo Aripuana,Rod.
do Estanho km 120, 21 Apr 1985 (fr), Cid Ferreira 5682
(INPA,MO,NY,US). PAR":Rio Mapuera, Cach.de Ma-
damee Cachede Ilhas,15Aug 1986(fl 6), CidFerreiraet
al. 7776(F,INPA,K, MO,NY,US).
Local name. Peru:muena.
Among species of Endlicheria with sessile outer
stamens, E. sprucei alone has a dense pin-prickpat-
tern (minutelypunctulate)and immersedtertiaryve-
nation on the upper surface of greyish green drying
leaves. Elsewherein Endlicheria,the leaves of E. gra-
cilis are similar in color, texture,indument,and sur-
face features,but ovate and triplinerved.
Indumentcolor varies from greyish green to rusty
red, but Endlicheriasprucei is otherwisehomogene-
ous. Mez (1889) and Kostermans (1937) both re-
ported that all stamens are provided with a pair of
basal glands; however, as in most species of Laura-
ceae, and all of Endlicheriaexcept E. vinotincta,only
the inner stamens (whorl III) are thus equipped.Fil-
amentsof whorl I (Fig. 1B) and II stamensareindeed
fleshy, but the tissue does not appearto be glandular,
nor does it appearto representa separatestructure.
Mez (1889) assigned Endlicheria sprucei to su-
bgen. Ampelodaphneand ITS sequencedatasuggests
close relationshipwith one of its members,E. reflec-
tens (Fig. 2). As discussed above (GenericDelimita-
tion and Species Groups),E. sprucei may occupy an
SYSTEMATICTREATMENT 33

intermediate position between the Ampelodaphne inside minutelypapillose. Stamensof whorls I and II
species group and species of Rhodostemonodaphne, 0.6 mm tall, sessile, the anthersovate, 0.3 X 0.4 mm,
including R. crenaticupula.Fruiting material of the the apex truncate,the connectives broad above the 2
latterhas been mistakenfor Endlicheriasprucei, but locelli, these ellipsoid, introrse,the filamentsligulate,
its upperleaf surfaceis traversedby a reticulatepat- as broad as anthers, slightly narrowerin whorl II,
tern of prominenttertiaryveins. densely grey-tomentose;whorlIIIstamenssessile, 0.6
mm tall, the anthers oblong, 0.4 X 0.3 mm, erect,
locelli 2, occupying the lower half of the anther,
8. Endlicheria metallica Kosterm., Recueil Trav. extrorse-latrorse,the filamentsbroaderthan anthers,
Bot. Neerl. 34: 543. 1937. Type. Brazil. Ama- fleshy,the basal glands absent;whorlIV wanting;pis-
zonas:Near mouthof Rio Embira(tributaryof Rio tillode filiform.Pistillateinflorescencewith indument
Tarauaca),19 Jun 1933 (fl Y), Krukoff4932 (ho- and color as in staminateplants, but shorterand with
lotype: NY; isotypes: A, K-n.v., MO, S, U). fewer lateral branches, the flowers slightly deeper;
stamens sterile, smaller;ovary glabrous,ovoid; style
Trees to 35 m. Branchlets stout, midway along
stout, weakly distinguishedfrom ovary; stigma min-
flush 3-5 mm diam., striate, angular, silvery seri-
utely tri-lobed, 0.3 mm diam. Fruits borne on stout
ceous, the surface barely visible to concealed by the
claviformpedicels of up to 2 X 0.7 cm; cupulesheavy
indumentcover, the hairs short, to 0.3 mm, straight,
walled, hemispherical,to 1.5 X 3 cm, glabrousinside
appressed;terminalbuds plump, 3 X 2 mm, silvery
and outside, the marginsentire;drupesovoid, to 4 X
sericeous. Leaves alternate,widely andevenly spaced
2 cm.
along currentflush; petioles slender,to 3 X 0.3 cm,
semi-terete, the indument as on branchlets;laminae Distribution (Fig. 8) and ecology. Trees from
coriaceous to chartaceous,plane, elliptic to oblong, well-drainedsoils in centralto westernAmazoniaand
11-20 X 4-9 cm, the base acute to obtuse, briefly adjacent eastern Andean foothills at 100-1100 m.
decurrent,the apex acute to obtuse, acuminatefor up Flowering from March to October,with fruits avail-
to 2 cm, the margins minutely recurvedthroughout; able by August until June of the following year.
upper surface greenish grey, waxy, the primary to
fourth-orderveins raised, their prominencedecreas- Representativespecimensexamined.COLOMBIA.
ing with rank;lower surface sparsely to moderately ANTIOQUIA:SanLuis,Quebrada LaCristalina,550-690 m,
pubescent, the hairs appressed, silvery grey, uni- 25 May 1987 (fr), Ram(rez& Cdrdenas1016 (MO); Alto
formly distributed,all vein ordersraised, theirprom- Rico,RioClaro,600m,9 Oct1982(fr),Renterfa et al. 2821
inence decreasingwith rank;secondaryveins 3-5 per (MO).
side, ? evenly spaced, slightly more distant around ECUADOR. MORONA-SANTIAGO: Pozo petrolero
midlamina, ascending at 50-60? (more obtusely 'Garza'de TENNECO, 35 km al NE de Montalvo,260 m,
2-12 Jul 1989 (fl Y, fr juv), Zak & Espinoza 4671 (MO).
aroundmidlamina),arcuate,the lowermostpairasym-
NAPO: Aguarico,
ReservaEtnicaHuaorani,
carretera
y oleo-
metric (one or both merging with margin at base),
ductode Maxus,km 54, 250 m, 26-30 Sep 1993 (fl d),
distal pairs loop-connected, or distal pairs weakly Aulestia&Andi841 (MO);Estaci6nExperimental INIAP-
loop-connected;tertiariesclose, roughlyhorizontalto Napo,Payamino, ReservaFloristica"ElChuncho,"250 m,
oblique to midrib, between secondaries straight or 5 Apr 1986 (fr), Jaramillo 8349 (B, MO). SANTIAGO-
once-forked.Staminateinflorescencesevenly spaced ZAMORA: Taisha, 500 m, 1 Feb 1962 (fr), Cazalet & Pen-
along currentflush in the axils of foliage leaves, to nington 7629 (B, K, US).
15 cm long with 8 lateralbranches,branchorders3-4, PERU. AMAZONAS: Bagua, Imaza-Yamayakat,400 m,
the highest order dichasial, lax, the flowers distant, 25 May 1996 (fl d), Vdsquez& Vdsquez20969 (MO), 450
the axes silvery strigose, distalmost branches and m, 15 Nov 1997 (fr), Vdsquezet al. 24898 (MO). Cuzco:
pedicels more densely so; bracts and bracteoles ca- La Convenci6n, 910 m, 12 Jun 1968 (fr), Dudley 10072
ducous by anthesis, ovate, sericeous; pedicels terete, (MO). HUANUCO: Pachitea, Puerto Inca, Llullapichis, 280
m, 16 May 1989 (fl Y), Kroll Saldafia 326 (MO). JUNiN:
to 2 mm long, those supporting secondary flowers
Juaja,Rio Negro, N of Satipo, 800 m, 20 Aug 1960 (frjuv),
slightly shorter.Flowersurceolate,up to 2 mm diam.,
Woytkowski5864 (US). LORETO: Requena, 140 m, 11 Aug
densely silvery sericeous outside; receptacle deeply 1988 (fl d), van der Werifet al. 10084 (MO); Maynas,Ca-
cyathiform, 1.3 x 1 mm, constricted below tepals, huide, 11 Oct 1984 (fr), Vdsquez& Jaramillo 5697 (MO).
silvery tomentose inside. Tepals chartaceous, MADRE DEDios: Tambopata,Inca, PuertoMaldonado,car-
depressed-ovate, 0.4 X 0.7 mm, the inner whorl reteraMaldonado-Cuzco,km 16, 240 m, 22 Jul 1989 (fr),
slightly smaller, erect at anthesis, the inner surface Vdsquezet al. 12435 (MO). PASCO: Oxapampa,Palcazd,
sparsely tomentose near base, the margins and apex 300-600 m, 26 Mar 1986 (fl d), Hartshorn et al. 2916
34 FLORA NEOTROPICA

~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
-..-F-
;

I~~~~~~~~~~~~~~~

0 peX 1 C1) ...S

i aS
X
--
------F-- Dis o of E r m a E c

<~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~CA
FIG.8. Dstriutin ofEridicheia etalica,E. hrysveluina,arl E. rownana

(MO). SANMARTiN: Rioja, PardoMiguel, 1100 m, 16 Jun two-locellate staminodesremnanton cupule rims. Af-
1998 (fl d), Sdnchez Vegaet al. 9363 (MO). ter this sorting exercise, it became apparentthat En-
BRAZIL. AMAZONAS: Manaus,Igarapedo Passarinho, dlicheria metallica is a predominantlywestern Am-
14 Mar 1956 (fr), Co6lho 3611 (INPA); Rio Purus, Cach-
azonian species (reaching the lower slopes of the
oeira Uby, 22 Jun 1903 (fl 6), Goeldi s.n. MG 3911 (U).
Andes) that consistently has elliptic or oblong leaves
BOLIVIA. LA PAz: Alto Madidi, 370 m, 26 May 1990
(fr), Gentry & Estensoro 70632 (MO); SANTACRUZ:Es-
that assume a greenish grey color in the dry state,
taci6n del Valle de Sacta, Regi6n del Chapare,5 May 1989 while these Rhodostemonodaphnespecies are re-
(fl d3), Moretti 1443 (MO). stricted to northeasternSouth America and have ob-
ovate leaves that dry darkbrown (see also Madrifinn,
Local names. Peru: mautaga, mantaga, moena 1996b).
callhuangiaamarilla,moena negra.Bolivia: laurelne-
gro.
Endlicheriametallica shares campanulateflowers 9. Endlicheria chrysovelutina Chanderbali,sp. nov.
and an androeciumof nine sessile stamens with E. Type. Peru. Loreto: Maynas, Iquitos, Puerto Al-
chrysovelutina,but differs by its closely appressed mendras,130 m, 27 Jul 1988 (fl d), van der Werff
silvery-grey indument. Rlower shape provides easy et al. 9815 (holotype: MO; isotypes: F, GH, NY-
contrastwith vegetatively similar Rhodostemonoda- n.v.). Fig. 9
phne grandis, R. praeclara, R peneia, and R. saulen- Endlicheriae metallicae absque dubio proxima, sed ra-
sis, but all fruiting specimens (25 of the 39 known mulis chrysovelutinis et foliis subtus praeclaro pannosis-
collections) had to be initially identified by locating sericeis differt.
 co

__ G *D -- E
0.5mmrHn 1mm:
FIG. 9. Endlicheria chrysovelutina(AXG, van der Werifet al. 9815; H, Vdsquez& Jaramillo 9183). A. Habit. B.
Close-up of lower leaf surface. C. Leaf base below. D. Rlower with facing tepal turneddown to reveal androecium.E.
Rlowerl.s. F. Whorl I stamen seen from within. G. Whorl HIIstamen seen from without. H. Fruitingbranchlet.
36
Trees to 20 m. Branchlets stout, midway along
flush 4-7 mm diam., striate, angular,densely veluti-
nous, the surface concealed by the indument cover,
the hairs relatively long, to 0.6 mm, straight,erect,
golden to reddish yellow; terminalbuds plump, 2 X
1 cm, densely pubescent, the hairs as on branchlets,
ascending. Leaves altemate, widely and evenly
spaced along current flush; petioles robust, to 4 X
0.4 cm, semi-terete, striate, the indument as on
branchlets;laminae coriaceous, plane, ovate to ellip-
tic, 14-23 X 8-10 cm, the base obtuse to truncate,
briefly decurrent,the apex obtuse, acuminateor cus-
pidatefor up to 1.2 cm, the marginsminutelyrecurved
near base or throughout;uppersurface light green to
olive-brown,waxy, the primaryto fourth-orderveins
raised, their prominencedecreasingwith rank;lower
surface densely golden to yellow pannose-sericeous,
the hairs ascending to appressed, reduced on main
veins, all vein orders raised, their prominence de-
creasing with rank;secondaryveins 3-5 per side, ?
evenly spaced, slightly more distantaroundmidlam-
ina, ascendingat 50-60? (moreacutelytowardsapex),
arcuate,the lowermostpair asymmetric(one or both
merging with margin at base), distal pairs loop-
connected; tertiaries roughly horizontal to midrib,
closely spaced, between secondaries straight to
forked. Staminateinflorescencesevenly spacedalong
currentflush in the axils of foliage leaves, to 25 cm
long with 12 lateralbranches,branchorders4-5, the
highest order dichasial, lax, the flowers distant, the
axes golden to yellow velutinous;bracts and bracte-
oles persistent at anthesis, eventually falling, ovate,
sericeous, the hairs as on axes; pedicels terete, to 4
mm long, those supportingsecondaryflowersslightly
shorter.Flowers urceolate, 3 mm diam., densely sil-
very to yellow tomentellose outside; receptacle cy-
athiform,2 X 3 mm, slightly constrictedbelow tepals,
grey-velutinous inside. Tepals chartaceous,broadly
ovate, 0.5 X 1 cm (the innerwhorl slightly narrower),
erect, surroundingandroeciumat anthesis, the outer
surface silvery tomentellose, the inner surface gla-
brous, the marginspapillose. Stamensof whorlsI and
II 0.5 mm tall, sessile, the anthersovate, 0.3 X 0.4
mm, glabrous, the apex truncate, the connectives
broadabove the 2 locelli, these ellipsoid, introrse,the
filaments ligulate, broaderthan anthers,densely sil-
very tomentose; whorl III stamens sessile, the indu-
ment and shape as in outer whorls, slightly narrower,
locelli 2, introrse-latrorse,the basal glands absent;
whorl IV wanting;pistillode filiform. Pistillate inflo-
rescence with indument and color as in staminate
plants,but shorterand with fewer lateralbranches,the
flowers slightly deeper; stamens sterile, smaller;
FLORANEOTROPICA
ovary glabrous; style slender, distinct from ovary;
stigma tri-lobed,0.5 mm diam. Fruitsborne on stout
claviformpedicels of up to 3 X 0.7 cm; cupules thick
walled, hemispherical,to 2 X 3.5 cm, glabrousinside
and outside, the marginsentire;drupesovoid, to 3.5
X 2 cm.
Distribution (Fig. 8) and ecology. Medium-sized
trees known only from lowland (ca. 120 m) white
sand forests aroundIquitos, Peru. Flowers have been
collected in June, July, and October, and the single
fruitingcollection was made in June.
Additional specimens examined. PERU. LORETO:
Maynas,PuertoAlmendras, 122m, 20 Oct1982(fl 9) Vds-
quez3319 (F, MO,NY); Iquitos,carretera
Iquitos-Nauta,
km 45, 120 m, 12 Jun 1987 (fl d) Vdsquez& Jaramillo
9180 (F,MO),(fr),Vdsquez& Jaramillo9183 (MO).
Endlicheriachrysovelutinadiffers from E. metal-
lica by its velutinousbranchletsanddenser,morefelt-
like, lower leaf surface indument.Both species most
likely belong to a species group in Rhodostemono-
daphne centeredaroundR. grandis (see discussion in
Relationships, above) wherein an undescribed spe-
cies, representedby Steyermark59001 (F, MO) from
Mount Roraima,also has the velutinousindumentof
Endlicheria chrysovelutina. Flowers in Steyermark
59001 are campanulate,as in E. chrysovelutina,but
stamens are spathulateand anthersare four-locellate.
Wereit not for this specimen,E. chrysovelutinawould
be recognizablein this transgenericallianceby its in-
dument alone, while flowers, or remnantstamens in
fruiting material, are necessary to assign all other
membersto genus.
10. Endlicheria browniana Mez, Jahrb.Konigl.Bot.
Gart. Berlin 5: 115. 1889. New name for Ayden-
dron macrophyllumMeisn., DC. Prodr.15(1): 92.
1864. Type. Panama. Darien: Cape Corrientes,
withoutdate (fl 9, fr juv), Seemann1094 bis (ho-
lotype: K).
Trees to 15 m. Branchletsmoderatelystout, mid-
way along flush 4-5 mm diam., distally weakly an-
gular,soon terete,rusty or silvery strigillose, the sur-
face clearly exposed to concealed by the indument
cover, the hairs very short, to 0.1 mm, straight,ap-
pressed;terminalbuds plump, 3 X 2 mm, sericeous.
Leaves alternate,widely andevenly spacedalong cur-
rent flush;petioles slenderor robust,to 2.5 X 0.4 cm,
semi-terete, the indumentas on branchlets;laminae
coriaceous to membranaceous, plane, obovate to
ovate, 15-35 X 7-15 cm, the base bluntly rounded,
or acuteandshortlyattenuate,the apexbroadlyobtuse
SYSTEMATICTREATMENT
then abruptlyacuminatefor up to 2.5 cm, the margins
minutely recurved throughout;upper surface dark
brown to light or olive green, waxy, the primary to
fourth-orderveins raised, their prominencedecreas-
ing with rank;lower surface sparsely strigillose, the
hairs uniformly distributed, all vein orders raised,
their prominence decreasing with rank; secondary
veins 6-9 per side, ? evenly spacedor closer towards
leaf base, ascendingat 50-60? (moreobtuselytowards
apex), arcuate,distal pairs loop-connected;tertiaries
oblique to midrib, between secondaries once-forked
to straight. Staminate inflorescences evenly spaced
along currentflush in the axils of foliage leaves, to
15 cm long with 8 lateralbranches,branchorders3-4,
the highest order dichasial, lax, the flowers distant,
the axes strigillose;bractsandbracteolescaducousby
anthesis, lanceolate, sericeous; pedicels graduallyin-
creasingin diameterapicallyand mergingwith recep-
tacle, to 2 mm long, those supportingsecondaryflow-
ers slightly shorter.Flowers hypocrateriform,3 mm
diam., densely rusty or silvery strigillose outside;re-
ceptacle globose, 0.6 X 1 mm, constrictedbelow te-
pals, densely rustyor silvery tomentoseinside. Tepals
chartaceous, ovate, 1 X 0.6 mm (the inner whorl
slightly broader),spreadingat anthesis,the inner sur-
face densely pubescent, the hairs yellowish, tightly
crinkledto papillose. Stamensof whorls I and II ses-
sile, 0.6 mm tall, the anthersovate, 0.4 X 0.5 mm,
minutely papillose, the apex apiculate, the connec-
tives prolongedbetween the 2 locelli, these obliquely
hemispherical, introrse-latrorse,the filaments ligu-
late, as broadas anthers,densely yellowish papillose;
whorl III stamens sessile, 0.6 mm tall, the anthers
depressed-ovate, 0.3 X 0.5 mm, erect, locelli 2,
extrorse-latrorse,the filaments as broad as anthers,
ligulate, the indument as in outer whorls, the basal
glands sessile, minute, globose; whorl IV wanting;
pistillode wanting. Pistillate inflorescencewith indu-
ment, color, and branchingas in staminateplants,the
flowers slightly deeper; stamens sterile, smaller;
ovary sparselytomentellose or glabrous,ovoid; style
stout, weakly distinguished from ovary; stigma tri-
lobed, papillose, 0.5 mm diam. Fruitsbome on stout
terete pedicels of up to 2 X 0.5 cm; cupules hemi-
spherical, to 1 X 2 cm, glabrousinside and outside,
the marginsundulate;drupesovoid, to 3 X 1.5 cm.
Distribution (Fig. 8) and ecology. Small to
medium-sized trees from lowlands and lower mon-
tane slopes (100-800 m) west of the Andes from Ec-
uador to Panama. Flowering material has been
collected from September to April, and fruits from
Octoberto June.
37
Additional specimens examined. PANAMA. BOCAS
DEL TORO: Chiriquf Lagoon, Fish Creek Mtns., 18 Apr
1941 (fl 6), von Wedel2257 (A). COCLE:Rd. to Coclesito,
12 mi from Llano Grande,600 m, 8 Dec 1983 (fr), Churchill
et al. 3974 (MO), 9 Dec 1983 (fr), Churchillet al. 3998
(MO). COL6N:Hills just N of Rio Guanche, 100-200 m,
16 Nov 1975 (fl 6), Davidse & D'Arcy 10097 (MO); Rio
Boquer6n, 100 m, ca. 14 mi from hwy., 13 Apr 1985 (fr),
Hammel13518 (MO);Rio Guanche,ca. 2.5 km upriverfrom
bridge on rd. to Portobelo,3 Jun 1975 (fr), Mori et al. 6476
(MO). DARIEN: Cape Corrientes, s.d. (fr), Seemann 1094
(K). PANAMA: El Llano-Carti Rd., 300 m, 6 Sep 1980 (fl
d), Sytsma 994 (MO), 21 Mar 1996 (fl 6), Guerra 646b
(MO); GorgasMemorial labs, ca. 25 km NE of CerroAzul
on Rfo Piedras,550 m, 25 Nov 1974 (fl 2), Mori & Kallunki
3472 (MO). SANBLAS: El Llano-CartiRd., km 16.7, trail
W to Rio Carti Grande, 250-350 m, 4 Nov 1984 (fl 6),
Nevers & Herrera4186 (MO);El Llano-CartiRd., km 26.5,
along Rio Carti Chico, 200 m, 12 Apr 1985 (fr), Nevers et
al. 5339 (MO);Play6n Chico, Rio Grande,50-100 m, 2 Nov
1991 (fl 2), Herreraet al. 1057 (HBG, MO).
COLOMBIA. CHOC6:Ca. 50 km W of Las Animas,
ca. 4 km E of Rfo Pato on Pan American Hwy., 250 m, 11
Jan 1979 (fl 6), Gentry& Renterfa23983 (HBG, MO), (fr),
Gentry & Renteroia 23982 (MO). VALLE: Buenaventura,
Quebrada"LaTrojita,"30-100 m, 20 Mar 1992 (2 fl & fr),
Cogollo et al. 5119 (MO); Rio Calina,0-5 m, 11 Mar 1944
(fl 6), Cuatrecasas16855 (F, US).
ECUADOR. CARCHI: San Marcos de los Coaqueres,
Chical-Tobar Donoso Trail, 800 m, 8 Feb 1985 (fl 6),
0llgaard et al. 57630 (MO, NY), (fr), 0llgaard et al.
57623A (MO).ESMERALDAS:SanLorenzo,ReservaEtnica
Awa, 250 m, 22 Mar 1993 (fl 2), Aulestia & Aulestia 1289
(MO, NY); Eloy Alfaro, Reserva Ecol6gica Cotacachi-
Cayapas,ParroquiaLuis VargasTorres,250 m, 23-27 Oct
1993 (fr), Tirado et al. 586 (MO); Muisne, Sitio San Sal-
vador,orillas del Rfo Sucio, 100-150 m, Mar 1995 (fl 6),
Palacios 13750 (MO).
Endlicheria browniana is unmistakable in fruit on
account of its hemispherical cupules with undulate
marginsand stout pedicels. Its hypocrateriformflow-
ers with distally constrictedreceptaclesbelow hori-
zontally spreadingtepals are much like those found
in E. jefensis, E. mishuyacensis, and E. pyriformis,
but the laxly reticulatetertiariesof these threespecies
are quite unlike the percurrentpattern formed by
those of E. browniana.
The concept of Endlicheria browniana adopted
here is broaderthan that suggested by the type ma-
terial and most of the other collections from the Dar-
ien-Choc6 region. These have large obovate coria-
ceous leaves with roundedbases and stout,seemingly
swollen, petioles. Moreover, pistillate flowers have
densely pubescent ovaries, a feature otherwise un-
known in Endlicheria. These charactersseem suffi-
cient to circumscribea highly distinctivespecies, but
38 FLORANEOTROPICA

ovaries are glabrousin a vegetativelyindistinguisha-

Leaves alternate,widely andevenly spacedalong cur-
ble collection from quite distantSan Blas in Panamarentflush;petioles slender,to 2 X 0.2 cm, semi-terete,
(Herreraet al. 1057). Furthermore,in Gentry& Ren- sparsely grey-strigillose above, otherwise glabrous;
terfa 23982 from the Choco, young drupes are gla- laminae coriaceous, plane, ovate to elliptic, 9-14 X
brous and leaves are much smallerwith ratherslender3-5 cm, the base acute, attenuate,the apex acute,acu-
petioles compared to other Darien-Choco material. minatefor up to 1 cm, the marginsminutelyrecurved
Given this variation in ovary pubescence and leaf throughout;upper surface deep olive-brown, waxy,
morphology in the Darien-Choco region, and since the primaryto fourth-orderveins raised, their prom-
staminate flowers from there (Cuatrecasas 16855, inence decreasing with rank;lower surface sparsely
Gentry & Renteria13983) are indistinguishablefrom strigillose, the hairs as on branchlets,denseron main
those collected elsewhere, an expandedconcept of E.veins, soon lost, all vein orders raised, their promi-
browniana is unavoidable. Unfortunately,this re- nence decreasingwith rank;secondaryveins 4-7 per
coursealso increasesthe rangeof vegetativevariation.
side, ? evenly spaced, slightly more distant around
Variationin leaf size, shape, and textureappearscon-
midlamina, ascending at 50-60 (more obtusely to-
tinuous and can be ignored as a possible source of wardsapex), arcuate,distalpairsloop-connected;ter-
specific characters,but variationin indumentis lesstiaries laxly reticulate. Staminate inflorescences
readily dismissed. In most collections, including the
evenly spaced along currentflush in the axils of fo-
Darien-Choc6 material discussed above, the indu- liage leaves, to 7 cm long with 12 lateral branches,
ment of branchlets,leaves, inflorescences,andflowers
branchorders2-3, the highest orderdichasial,lax, the
consists of dull rust-coloredhairs, but all Ecuadorian
flowers distant, the axes sparsely grey-strigillose;
material have a shiny silvery-grey indument. How- bracts and bracteoles caducous by anthesis (none
ever, any apparent correlation between indument seen); pedicels graduallyincreasingin diameterapi-
color and geographic location is disrupted by two cally, to 3 mm long, those supportingsecondaryflow-
Panamaniancollections with silvery-grey indument ers slightly shorter.Flowershypocrateriform,to 4 mm
(von Wedel2257 and Churchillet al. 3974). Further- diam., sparsely grey-strigillose, the hairs scattered,
more, fruits of silvery-grey forms from Ecuador as soon lost; receptacledeeply infundibuliform,the apex
well as Panamashow the distinctive cupules of Dar- slightly constricted below tepals, 1.8 X 2 mm,
ien-Choc6 material. Within both color morphs, the densely tawny papillose inside. Tepals chartaceous,
indumentdensity on branchletsrangesfrom sparseto broadly ovate, 1.1 X 0.9 mm, spreading,the inner
surface rusty papillose. Stamens of whorls I and II
subsericeous,and in the lattercase providesan aspect
reminiscentof the E. sericea group (species 46-55).sessile, 0.6 mm tall, the anthersbroadlyovate, 0.3 X
Separationon the basis of indumentcolor or density 0.6 mm, glabrous,the apex apiculate,the connectives
alone or in combinationwould produceeithertwo or prolongedbetween the 2 locelli, these obliquelyhem-
ispherical, introrse-latrorse,the filaments broader
four entities, neitherof which could be recognizedon
the basis of other characters. than anthers,densely rusty papillose; whorl III sta-
mens sessile, 0.6 mm tall, the anthersovate, 0.4 X
0.4 mm, erect, locelli 2, extrorse-latrorse,the fila-
ments broaderthan anthers,shortly columnar,rusty
11. Endlicheria jefensis van der Werffex Chander-
yellowish papillose, the basal glands sessile, globose;
bali, sp. nov. Type. Panama.Panama':Cerro Jefe
whorl IV wanting;pistillode wanting.Pistillateplants
region, along road to Rio Crist6bal in Chagras
unknown.
drainage,600 m, 25 Feb 1986 (fl d'), McPherson
8493 (holotype:MO; isotypes: F, HBG, MO, NY, Distribution (Fig. 11) and ecology. Known only
PMA, US). Fig. 10 from the type material taken from a staminatetree

Endlicheriaebrownianae et affinibussimiliset nullodu- found floweringin Februaryon the forestedslopes of

bio his speciebusproxima,sed floribussub-glabris et foliis CerroJefe (Panama).
ovatisdiffert.
In the Endlicheria browniana species group, the
Trees to 8 m. Branchlets slender, midway along narrowlyovate to elliptic coriaceous leaves and rel-
flush 2-3 mm diam., distally weakly angular,soon atively large subglabrousflowers of E. jefensis are
terete, sparsely strigillose, the surface clearly ex- without equal. In Panama,E. browniana has much
posed, dark brown, the hairs short, to 0.1 mm, larger leaves and densely pubescent flowers, and E.
straight,appressed,greenishgrey;terminalbuds slen- formosa has depressed-globoseflowers. In adjacent
der, 2 X 0.6 mm, densely greenish white sericeous. Colombia, E. colombiana has similar indument,but
SYSTEMATICTREATMENT 39

FIG. 10. Endlicheriajefensis (McPherson8493). A. Habit. B. Leaf base below. C. Flower. D. Flower ls. E. Whorl
Ill stamen seen from without. F. Whorl I stamen seen from within.
40 FLORA NEOTROPICA

7i

-- . ----- - - ----

*~~~~~~~~~~~~~~~~~~~~~~

A ...~~~~~~~~~~~~~J

y %~~~~~~tf4

--- 11 Ditibto
FIG of Enlcei eessEclmin n ihycn

broader leaves with canaliculate petioles. Western 0.3 cm, canaliculate,the indumentas on branchlets;
AmazonianE. mishuyacensishas similar-sizeddark- laminae chartaceous,plane, elliptic, 11-25 X 5-12
drying leaves, but these are obovate to elliptic, and cm, the base obtuse, briefly decurrent,the apex ob-
rust brown rather than greyish hairs appear on the tuse, acuminatefor up to 1 cm, the marginsminutely
branchlets. recurved throughout; upper surface light greyish
green to olive-brown, waxy, the primary to fourth-
order veins strongly raised; lower surface sparsely
12. Endlicheria colombiana (Meisn.) Mez, Jahrb. strigillose, the hairs soon restricted to main veins,
Konigl. Bot. Gart. Berlin 5: 117. 1889. Oreoda- eventually lost, the primary to fourth-orderveins
phne colombiana Meisn., DC. Prodr. 15(1): 137. raised, their prominence decreasing with rank; sec-
1864. Type. Colombia. Antioquia:Withoutlocal- ondary veins 5-9 per side, ? evenly spaced, slightly
ity and date (fl 9), Jervise s.n. (lectotype, desig- more distant aroundmidlamina,ascending at 50 60?
nated by Kostermans,1937: K). (more obtusely towardsapex), arcuate,or archingaf-
Trees to 8 m. Branchlets slender, midway along ter midcourse, distal pairs loop-connected;tertiaries
flush 3-5 mm diam., distally weakly angular and oblique to midrib, between secondaries straight to
sparsely strigillose, soon terete and glabrous,the sur- once-forked. Staminateinflorescences evenly spaced
face always clearly exposed, grey to darkbrown, the along current flush in the axils of foliage leaves, to
hairs short, to 0.15 mm, straight,appressed,greenish 10 cm long with 9 lateralbranches,branchorders2-3,
grey; terminal buds plump, 3 X 2.5 mm, densely the highest order dichasial, lax, the flowers distant,
grey-sericeous. Leaves alternate,widely and evenly the axes sparsely grey-strigillose,distal branchesand
spaced along currentflush; petioles slender,to 1.5 X pedicels more densely so; bracts and bracteoles ca-
SYSTEMATICTREATMENT 41

ducous by anthesis, none seen; pedicels terete, to 7 flowers are infested with fungal mycelia, and thereis
mm long, those supportingsecondaryflowersslightly no sign of pollen in often shallow locelli.
shorter. Flowers hypocrateriform, 5 mm diam.,
sparselygrey-strigilloseoutside;receptacleinfundib-
uliform, 2 X 3 mm, merging with tepals, densely 13. Endlicheria mishuyacensis A. C. Sm., Bull.
rusty tomentellose inside. Tepals chartaceous,ovate, TorreyBot. Club 58: 102. 1931. Type. Peru.Lor-
2 X 1.3 mm, ascending to spreadingat anthesis, the eto: Mishuyacu, near Iquitos, 100 m, Oct-Nov
inner surface rusty tomentellose-papillose.Stamens 1929 (fl d), Klug 204 (holotype:NY; isotypes: F,
of whorls I and II sessile, 0.7 mm tall, the anthers US).
broadlyovate, 0.6 X 0.7 mm, minutelypapillose, the
apex apiculate,the connectivesbluntlyprolongedbe- Trees, 3-20 m. Branchletsslender,midway along
tween the 2 locelli, these obliquely hemispherical, flush 2-3 mm diam., angular,sparselystrigillose, the
introrse-latrorse,the filamentsligulate,as broadas an- surfaceclearly exposed, darkbrown,the hairsreddish
thers, rusty tomentellose;whorl III stamenssessile, 1 brown, ca. 0.1 mm, appressed;terminalbuds slender,
mm tall, the anthersovate, 0.6 X 0.5 mm, erect,locelli 1 X 0.3 mm, tan sericeous. Leaves alternate,widely
2, extrorse-latrorse,the filaments broader than an- and evenly spaced along currentflush; petioles slen-
thers, columnar,rusty tomentellose, the basal glands der, to 1 X 0.2 cm, canaliculate,the indumentas on
sessile, globose; whorl IV wanting; pistillode fusi- branchlets;laminaechartaceous,plane, elliptic to ob-
form. Pistillateinflorescencewith indumentandcolor ovate, 7-20 X 3-8 cm, the base acute, attenuate,the
as in staminateplants,but shorterand with fewer lat- apex acute, acuminatefor up to 1.5 cm, the margins
eral branches, the flowers slightly smaller; stamens flat; upper surface dark green to blackish brown,
sterile, smaller;ovary glabrous;style slender,distinct waxy, the primaryto fourth-orderveins raised, their
from ovary; stigma tri-lobed.Fruitsunknown. prominence decreasing with rank; lower surface
sparselyrusty strigillose, the hairs slightly denseron
Distribution (Fig. 11) and ecology. Small trees main veins, all vein orders raised, their prominence
from the Andes of Antioquia(Colombia)at ca. 1500 decreasing with rank;secondary veins 5-7 per side,
m. The two collections with calendar information ? evenly spaced, slightly more distant aroundmid-
were flowering in May and November. lamina, patent, diverging at 70-85? (more obtusely
aroundmidlamina),arcuateto abruptlyascendingaf-
Additionalspecimensexamined.COLOMBIA.AN- ter midcourse, distal pairs loop-connected;tertiaries
TIOQUIA: Medellfn, May 1852 (fl 2), Triana2040-2 (K);
laxly reticulatingbetween secondaries.Staminatein-
Frontino,Murri,Nutibara-La BlanquitaRd.,km 22, 1450
florescences evenly spaced along currentflush in the
m, 4 Nov 1988 (fl 6), Zarucchi et al. 7120 (INPA, MO).
WITHOUT LOCALITY AND DATE: (fl 2), Linden429 (K);(fl
axils of foliage leaves, to 5 cm long with 3 lateral
2), Triana 1033 (P). branches,branchorders2-3, the highest orderdicha-
sial, lax, the flowersdistant,the axes rustystrigillose,
Endlicheria colombiana is a poorly known spe- distal branchesand pedicels more densely so; bracts
cies. The 19th century collections seen by Meissner and bracteolescaducousby anthesis,ovate, rusty se-
(1864), Mez (1889), and Kostermans(1937) are all riceous; pedicels graduallyincreasingin diameterap-
pistillate plants with unfertilizedflowers. They have ically, to 2 mm long, those supporting secondary
spreadingtepals and bear sessile stamens with ovate flowers slightly shorter.Flowers hypocrateriform,ca.
antherstypical of the E. brownianaspecies group,but 3 mm diam., densely rusty strigillose outside;recep-
their smooth receptacle-tepal transition stands out tacle infundibuliform,0.3 X 0.6 mm, slightly con-
among the distally constricted receptacle elsewhere stricted below tepals, rusty papillose inside. Tepals
in this species group. Vegetatively, these pistillate chartaceous,ovate to triangular,1 X 0.6 mm (the in-
plants are almost identicaland undoubtedlyrepresent ner whorl slightly narrower),spreadingto reflexedat
the same species, but a staminateplant from nearthe anthesis, the inner surface minutely rusty papillose,
type locality in Antioquia, Zarucchi et al. 7120, is the indumentreduced to a basal triangularpatch in
placed here with hesitation.Flowers of that specimen the outer whorl but covering surface of inner whorl.
show the smooth receptacle-tepaltransitionnoted as Stamens of whorls I and II 0.5 mm tall, sessile, the
unique to E. colombianain the E. brownianaspecies anthersovate, 0.3 X 0.3 mm, minutelyrustypapillose
group, but they are much more robust and show a nearbase on the abaxialside, otherwiseglabrous,the
sharperpedicel-receptacle transitionthan the pistil- apex apiculate, the connectives prolonged between
late flowers.Yet this may be a consequenceof disease. the 2 locelli, these obliquely hemispherical,introrse-
Almost all leaves show severe insect damage,several latrorse,the filamentsfleshy, slightly broaderthanan-
42
thers, densely rusty papillose; whorl III stamens ses-
sile, 0.6 mm tall, the anthersovate, equal to those of
the outerwhorls, erect, locelli 2, extrorse-latrorse,the
filamentsas broad as anthers,fleshy, columnar,rusty
papillose, the basal glands absent;whorl IV wanting;
pistillode filiform. Pistillate inflorescencewith indu-
ment, color, and branchingas in staminateplants,the
flowers similar in size and shape; stamens sterile,
smaller; ovary glabrous, ovoid; style stout, weakly
distinguished from ovary; stigma tri-lobed, ca. 0.4
mm diam., minutely papillose. Fruitsborne on clav-
iform pedicels of up to 1.5 X 1 cm; cupules shallowly
hemispherical,thick walled, to 1 X 2 cm, glabrous
inside and outside, the marginsentire;drupesovoid,
to 3 X 2 cm.
Distribution (Fig. 11) and ecology. Medium-
sized trees already fertile at 3 m occurring in non-
flooded lowland(100-400 m) forestsin westernAma-
zonia. Flowers and fruits have both been collected in
all months but January,February,and March.
Representativespecimensexamined.COLOMBIA.
AMAZONAS: Rio Apaporis,entre el RfoPacoy el Rfo Kan-
anari, Soratama,250 m, 25 Sep 1951 (fl 5d), Schultes &
Cabrera 14125 (COL, NY); Rio Miritiparana,CainoGua-
caya, 250 m, 24 Apr 1952 (fr), Schultes & Cabrera 16230
(GH,HBG,NY).
ECUADOR.PASTAZA: Pozo petrolero'Golondrina'de
PETRO-CANADA,25 km al NW del pueblo de Curaray,
400 m, 23 Jun 1989 (fl Y), Rubio & Gudifio199 (G, MO);
Auca, 115 km al S de Coca, Rfo Tigiiino, 320 m, 29 Apr
1989 (fr), Rubio 115 (G, MO).
PERU. LORETO: Maynas,Mishuyacu,nearIquitos, 100
m, Dec 1929 (fl Y), Klug 703 (F, NY, US); Iquitos, Quis-
tococha, 22 Nov 1940 (fl d), Asplund14682 (G, K, NY, R);
Mishana, Estaci6n Biologica Callicebus, 15 Aug 1980 (fr),
Foster 4302 (HBG, MO, NY, U).
BRAZIL.AMAZONAS:Basin of Rio Jurua,near mouth
of Rio Embira(tributaryof Rio Tarauaca),21 Jun 1933 (fl
d5),Krukoff4959 (A, F, G, K, MO, NY, U); Lago do Cas-
tanho, 25 Jun 1973 (fl ), Albuquerqueet al. 850 (INPA).
ACRE:Cruzeirodo Sul, Rio Jurua& Rio Moa, Serrade Moa
village, 28 Apr 1971 (fl d), Prance et al. 12633 (HBG, K,
MO, NY, US).
Local names. Colombia: ko-mwa-ko-ree (canoe
tree). Peru:muena.
Endlicheriamishuyacensisalone in the E. brown-
iana species group combines rusty pubescence and
darkbrown leaves with patent secondariesand retic-
ulate tertiaries. SympatricE. dysodanthais superfi-
cially similar with dark leaves of similar size, shape,
and color, but has barbellatetufts of hairs in the axils
of the secondary veins below and scalariformterti-
aries.
FLORA NEOTROPICA
14. Endlicheria pyriformis (Nees) Mez, Jahrb.Kon-
igl. Bot. Gart.Berlin. 5: 116. 1889. Cryptocarya
pyriformis Nees, Syst. laur. 220. 1836. Mespilo-
daphne pyriformis (Nees) Meisn., DC. Prodr.
15(1): 108. 1864. Type. French Guiana.Without
locality and date (fr), Poiteau s.n. (holotype: B-
n.v.; isotypes: G, LE, NY, P).
Endlicheriaglaberrima Mez, Bull. Herb. Boiss. 5 (ser.
2): 236. 1905.Type.Peru.Loreto:Yurimaguas, Aug
1902 (fl Y), Ule 6296 (holotype:B-n.v.; isotype:
HBG).
AnibaflexuosaA.C. Sm., Phytologia1: 117. 1935.
Type.Brazil.Amazonas: Nearmouthof RfoEmbira
(tributaryof RioTarauaca),
21 Jun1933(fl ), Kru-
koff5030 (holotype:NY).
Trees to 10 m. Branchletsslender,midway along
flush 2-3 mm diam., angular,glabrous, light green;
terminal buds slender, 2 X 0.6 mm, sparsely grey-
strigillose. Leaves alternate, widely and evenly
spaced along currentflush; petioles usually slender,
usually 1 X 0.1 cm (rarelyto 4 X 0.3 cm), canalic-
ulate, glabrous;laminae chartaceous,plane, obovate
to elliptic, 10-25 X 3-17 cm, the base cuneate,atten-
uate, the apex obtuse, acuminatefor up to 1.5 cm, the
marginsflat;uppersurfacelight green, waxy, the pri-
mary to fourth-orderveins raised, their prominence
decreasing with rank, the midrib and secondaries
creamish yellow, conspicuous against the greenish
lamina;lower surfaceglabrous,all vein ordersraised,
their prominence decreasing with rank; secondary
veins 6-8 per side, ? evenly spaced, slightly more
distantaroundmidlamina,patent,divergingat 70-85?
(more obtusely aroundmidlamina),abruptlyascend-
ing after midcourse,distal pairs loop-connected;ter-
tiaries laxly reticulating between secondaries. Sta-
minate inflorescences evenly spaced along current
flush in the axils of foliage leaves, to 15 cm long with
10 lateral branches, branch orders 3-4, the highest
order dichasial, lax, the flowers distant, the axes
sparselystrigillose, glabrescent;bractsandbracteoles
persistentat anthesis, triangular,sparsely strigillose;
pedicels graduallyincreasingin diameterapically,to
5 mm long, those supporting secondary flowers
slightly shorter. Flowers hypocrateriform, 3 mm
diam., sparselygrey-strigilloseoutside;receptaclecy-
athiform,2 X 1.3 mm, densely pubescentinside, the
hairs papillose, crisped, or straight. Tepals charta-
ceous, ovate, 1 X 0.6 mm, both whorls equal, spread-
ing at anthesis, the inner surface minutely papillose
throughout,or the hairs crisped to straightnearbase.
Stamens of whorls I and II broadly stipitate, 1 mm
tall, the anthersnarrowlyovate, 0.5 x 0.3 mm, gla-
brous, the apex apiculate, the connectives strongly
SYSTEMATICTREATMENT 43

prolongedbetween the 2 locelli, these obliquely hem- Flowering from April to December. Fruits collected
ispherical, introrse-latrorse,the filamentsligulate, al- throughoutthe year.
most as broadas anthers,sparselypubescent,the hairs
crisped to papillose; whorl III stamens sessile, 1.2 Representative specimens examined. COLOMBIA.
mm tall, the anthersnarrowly ovate 0.6 x 0.3 mm, META: Sierra de La Macarena,630 m, 24 May 1973 (fr),
Garcfa et aL 380 (COL); Villavicencio, Bosques de Viena,
erect, locelli 2, extrorse-latrorse,the filaments as
28 Apr 1988 (fr), Quiflones 1419 (MO). PLTTuMAYO:Mo-
broad as anthers,columnar,the indumentas in outer
coa, 28 Jul 1990 (fl d), Garcia et al. 103 (MO); Mocoa,
whorls, the basal glands sessile, minute, globose- 1350-1420 m, 20 Apr-I May 1994 (fr), Ferndndezet al.
apiculate;whorl IV wanting; pistillode filiform. Pis- 11051 (COL, GH).
tillate inflorescencewith indument,color, andbranch- VENEZUELA. ARAGUA: Henri Pittier National Park,
ing as in staminateplants, the flowers slightly deeper; 770 m, 6 Apr 1990 (fl d), Edwards & Roe 423 (MO).
stamens sterile, smaller;ovary glabrous,ovoid; style GUYANA. EssEQuiBo: Rupununi Region, Acarai
slender,distinctfrom ovary; stigmabroadlytri-lobed, Mtns., 500 m, 5 Mar 1994 (fr), Henkel et al. 4946 (MO),
0.6 mm diam. Fruits borne on claviform pedicels of 300-600 m, 2 Nov 1996 (fl 6), Clarke et al. 2865 (MO).
up to 2.5 x 0.5 cm; cupules infundibuliform,to 0.8 SURINAME. Tafelberg,East Ridge Creek Gorge, 750
X 1.7 cm, glabrous inside and outside, the margins m, 29 Aug 1944(fl d), Maguire 24538 (A, NY, U); Wih-
entire;drupesellipsoid, to 3 x 1.5 cm. elmmina Mtns., JulianaTop, 1236 m, 13 Aug 1963 (fl 6),
Irwin et al. 54744 (A, HBG, MO, U).
Distribution (Fig. 12) and ecology. Small un- FRENCH GUIANA. Montagne Bellevue de 1' Inini,
derstory trees of noninundated forests from the 600 m, 1 Sep 1985 (fl 6), de Granville et al. 7949 (MO,
Guianas to western Amazonia and along lower An- NY, US); Eau Claire,near Saul, 200 m, 11Aug 1993(fl 6),
dean slopes from Peru to Venezuela,at 100-1600 m. van der Werf/et al. 12944 (MO, NY).

4* > .
--.*

'' ", ';"q.'> \' .. ' ' ' 1; ' '; ii... W 4N ;'\.
4 e ' '',~~ ~~~~~~~~~~~~~~~~~~~
.7' >. , v
- t T._v..

f -* t t s- 9 / . g . ;. gS . S. ;\rffi * r
fi

FIG 12 D istribution of
t Endl hena pynfo. . *

-~~~
.. - - - -- .tz-r
f- - ,
:-
. 7Az-

FIG. 12. Distribution of Endlicheria pyriformis.

44
ECUADOR. MORONA SANTIAGO: El CentroShuar 15. Endlicheria formosa A. C. Sm., Phytologia 1:
Kankaim, Rfo Kankaim, 500 m, 18 Dec 1985 (fr), Shiki 118. 1935. Type. Brazil. Amazonas:Basin of Rio
RBAE359 (MO). NAPO: Estaci6n Biol6gica Jatun Sacha, Jurua,nearmouthof Rio Embira(tributaryof Rio
450 m, 17-21 Nov 1988 (fl 6), Ceron & Iguago 5562 (MO); Tarauaca),4 Jul 1933 (fl d), Krukoff5156 (holo-
Reserva Floristica "El Chuncho,"250 m, 7 Oct 1987 (fr),
type: NY; isotypes: A, F, G, K, S, U).
Ceron 2397 (HBG, MO).
PERU. HUANUCO: Rio Cuchara, Villa Ysabel, 21 Sep
1961 (fr), Schunke5679 (F, K). LORETO:Maynas, Rio Pu-
tumayo, Florida, mouth of Rfo Zubineta, 180 m, Oct-Nov
1931 (fl d), Klug 2313 (A, MO); Alto Amazonas, Balsa-
puerto, 220 m, Mar 1933 (fl 6), Klug 2958 (GH, K, MO).
PUNO:Rio Tavara,400 m, 19 May 1992 (fl Y), Gentry et
al. 76844 (MO); Sandia,between Rio Azata-Colorado,1100
m, 26 Jun 1986 (fl 6), Niiunez& Munioz5324 (MO). SAN
MARTiN: Lamas, Convento,trail to Tioyacu and Nuevo La-
mas, km 68 on Tarapota-Yurimaguasrd., 270 m, 23 Jun
1984 (fl 6), Knapp & Mallet 6531 (F, MO, NY, US); Mar-
iscal Caceres,Campanilla,SE of Caseriode Si6n, Rfo Si6n,
3 Oct 1969 (fr), Schunke3474 (COL, G, GH, NY, US).
BRAZIL. AMAPA:Rio Jari,400 m, 17 Aug 1993 (fl 9),
de Granvilleet al. 12329 (MO, US); Colonia do Torrao,29
Aug 1962 (fl 9), Pires & Cavalcante52676 (MG, NY, US).
AMAZONAS: Manaus, Reserva Florestal Ducke, Manaus-
ItacoatiaraRd., km 26, 31 Jul 1997 (fl 6), AssunCdo& Silva
583 (MO); Rio Ituxi, B6ca do Rio Curuquete,8 Jul 1971 (fl
6), Prance et al. 13987 (HBG, INPA, K, MG, MO, NY).
PARA:Altamira-ItaitubaRd., near EMBRAPA station, 30
Oct 1977 (fr), Berg 770 (HBG, MO, NY, U); Oriximind,Rio
Trombetas, 17 Jul 1980 (fl 9), Cid Ferreira et al. 1563
(HBG, INPA, MG, MO, NY). RONDONIA: Porto Velho,
UHE de Samuel, Rio Jamari,8 Jun 1986 (fl 6), Cid Ferreira
7387 (F, K, MO, NY).
Endlicheriapyriformisis conspicuous for its lack
of indument.Except for a fringe of short appressed
hairs on terminalbud scales, vegetativestructuresare
completely glabrous.Sparsehairs are found on inflo-
rescence bracts,bracteolesand floralpedicels, but the
inner surface of flowers bears the only noteworthy,
and variable, indument. In all material from the
Guianas and adjacentBrazil, the tepals and recepta-
cles inside arepapillose, as is typical of the E. brown-
iana species group. Such flowers are also found in
western Amazonia, e.g., Grandez & Jaramillo 817,
but most specimens from western Amazonia and the
eastern Andean slopes have a pilose indument of
straight erect hairs, e.g., Krukoff5030 (the type of
Anibafiexuosa) andPalacios 3260. Another,conceiv-
ably intermediate,indument form is found in Ule
6296 (type of Endlicheriaglaberrima),Prance et al.
13987, and Edwards & Roe 423, from Peru, Brazil,
and NE Venezuelarespectively.In these the hairs are
erect, as in material from western Amazonia, but
crisped ratherthan straight.As none of these three
forms can be recognized on the basis of other char-
acters, all are here kept togetherin E. pyriformis.
FLORA NEOTROPICA
Trees to 30 m. Branchlets stout, midway along
flush 4-8 mm diam., sharply angular,sparsely rusty
strigillose, the surface clearly exposed, dark brown,
the hairs very short, to 0.1 mm, straight,appressed;
terminalbuds slender,6 X 2 mm, greyish sericeous.
Leaves alternate,widely andevenly spacedalong cur-
rent flush; petioles slender,to 4 X 0.3 cm, canalicu-
late, the indumentas on branchlets;laminae charta-
ceous to coriaceous, plane, obovate, 12-30 X 3-15
cm, the base acute to cuneate,attenuate,the apex ob-
tuse, acuminatefor up to 2 cm, the marginsminutely
recurved throughout; upper surface dark greyish
green to olive-brown,waxy, the midrib prominulous
throughoutor sunken before midcourse, the higher-
order venation raised; lower surface sparsely pubes-
cent, the hairsas on branchlets,uniformlydistributed,
all vein orders raised, their prominence decreasing
with rank;secondary veins 8-10 per side, + evenly
spaced, slightly more distant aroundmidlamina,as-
cending at 50-60? (moreobtuselyaroundmidlamina),
arcuate,distalpairsloop-connected;tertiariesoblique
to midrib,between secondariesstraightto forked.Sta-
minate inflorescences evenly spaced along current
flush in the axils of foliage leaves, to 10 cm long with
16 lateral branches, branch orders 3-4, the highest
orderdichasial,lax, the flowersdistant,the axes rusty
to grey-strigillose, distalmostbranches and pedicels
more densely so; bracts and bracteoles caducous by
anthesis, ovate, grey-sericeous;pedicels terete, to 1
mm long, those supportingsecondaryflowersslightly
shorter. Flowers depressed-globose, 2 mm diam.,
sparsely to densely rusty strigillose outside; recepta-
cle patelliform,0.3 X 2 mm, densely rusty papillose
inside. Tepals chartaceous,broadlytriangular,0.3 X
0.6 mm (the inner whorl slightly narrower),erect to
inflexed at anthesis, the androeciumincluded except
for a narrowapical pore throughwhich anthervalves
protrude,the inner surface densely rusty papillose.
Stamens of whorls I and II sessile, unequal, whorl I
0.6 mm long, whorl II half as tall, the anthersovate,
0.4 X 0.2 in whorl I, 0.3 X 0.3 in whorl II, sparsely
papillose, the apex apiculate, the connectives pro-
longed between the 2 locelli, these obliquely hemi-
spherical, introrse-latrorse, the filaments fleshy,
broaderthan anthers,densely rusty papillose; whorl
III stamens sessile, 0.8 mm tall, the anthersnarrowly
ovate, 0.4 x 0.2 mm, erect,locelli 2, extrorse-latrorse,
the filaments broader than anthers, fleshy, densely
SYSTEMATICTREATMENT
rusty papillose, the basal glands globose, sessile;
whorl IV wanting;pistillode wanting.Pistillateinflo-
rescence with indument, color, and branchingas in
staminateplants,the flowerssimilarin size andshape;
stamens sterile, smaller;ovary glabrous,ovoid; style
slender,distinctfrom ovary;stigmatri-lobed,0.5 mm
diam., papillose. Fruitsbormeon teretepedicels of up
to 1 x 0.3 cm; cupules shallowly hemisphericalto
patelliform,to 2 x 0.3 cm, glabrousinside and out-
side, the marginsentire;drupesellipsoid to obovoid,
to 4 x 2 cm.
Distribution (Fig. 13) and ecology. Medium-
sized to tall trees from Amazonianlowlands as well
as uplandforests from Peruto Costa Rica, at ca. 100-
1700 m. Flowering and fruitingspecimens were col-
lected in almost every month of the year.
Representativespecimensexamined.COSTARICA.
PUNTARENAS: BuenosAires,Vallede Diquis,300 m, 24
Nov 1993 (fl 6), Morales et al. 2083 (F, MO, NY); Osa,
trail from PalmarNorte to Jalisco, on divide between Que-
bradaBatambaland QuebradaBenjamin,SW slope of Fila
Retinto, 300-400 m, 9 Dec 1988 (fl ), Grayumet al. 9153
(F, MO, NY).
PANAMA. PANAMA: Cerro Jefe, ca. 1.5 mi along Rio
Pacorard. fromjct. with CerroJefe rd., 750 m, 23 Jan 1986
(fl 6), McPherson & Merello 8131 (HBG, MO).
COLOMBIA.AMAZONAS: Leticia, Tarapaca,Parque
Nacional NaturalAmacayacu, 100 m, 25 Jun 1991 (fr), Ru-
das et al. 2504 (INPA, MO); Rio Caqueta, frente a Villa
Azul, 14 Sep 1989 (fl 6), van Andel et al. 293 (K, MO).
HUILA: ParqueNacional NaturalCueva de los Guacharos,
Rio Suaza, 1710 m, 21 Jun 1979 (fl d), Henao 152 (B).
META: Sierrade la Macarena,Rio Guapaya,500 m, 21 Jan
1950 (fl Y, fr juv), Philipson et al. 2192 (COL, US, NY).
PUTUMAYO: Mocoa, 1500-1670 m, 20 Apr-I May 1994
(fr), Bentancuret al. 5404 (COL, HUA, MO); Rfo Caucaya,
Parque Nacional La Paya, 240 m, 10-14 May 1993 (fr),
Bernal et al. 2039a (MO).
ECUADOR.CARCHI: Maldonado, Parroquia Tobar
Donoso, ReservaEtnicaAwa, Sabalera,900 m, 22 Nov 1992
(fr), Aulestia et al. 819 (MO). ESMERALDAS:Eloy Alfaro,
ReservaEcol6gica Cotacachi-Cayapas,CharcoVicente,Rfo
San Miguel, 130 m, 15 Jan 1993 (fl 6), Tipaz2516 (MO),
150 m, 6-9 Sep 1993 (fr), Palacios & Tirado11165 (MO).
NAPO:El Chaco,Rio Quijos,frentea la Estaci6nde Bombeo
del Rio Salado, 1400 m, 11 Sep 1990 (fl d), Palacios 5482
(F, MO); Estaci6n ExperimentalINIAP-Napo, Payamino,
Reserva Floristica "El Chuncho,"250 m, 29 Nov 1986 (fr
juv), Neill 7514 (HBG, MO, NY). SUCUMBiOS:Lago Agrio,
Reserva FaunisticaCuyabeno, 230 m, 15 Nov 1991 (fl 6),
Palacios et al. 8963 (F, HBG, MO), 250 m, 30 Jul 1996 (fr),
Palacios 13906 (MO). ZAMORA-CHINCHIPE:Nangaritza,
Miazi, Rio Nangaritza,930 m, 26 Oct 1991 (fl 6), Palacios
et al. 8644 (F, MO, NY), hill ca. 1 km upstream from
Shaime, 900-1100 m, 16 Feb 1994 (fl 6), van der Werf et
al. 13147 (F, HBG, MO).
45
PERU. AMAZONAS: Bagua, Imaza, ComunidadAgu-
arunade Kusd-Listra,CerroApag, 600-700 m, 15 Sep 1996
(fl d ), Diaz et al. 8144 (MO). Cuzco: Cuzco, Camisea,
CampamentoSan Martin-C,Camisea ProductionUnit, 467
m, 12 Jan 1997 (fl Y), Acevedo et al. 8646 (MO); Quispi-
canchis, hills aroundRfo Arazabetween Pan de Azucarand
Quince Mil Airport,643 m, 10 Aug 1991 (fr), Nuiniez13986
(MO). LORETO:Maynas, Iquitos, Nina Rumi (Rio Nanay),
122 m, Jul 1984 (fl d), Vasquez & Jaramillo 5277 (MO,
NY); Requena,ReservaNacionalPacaya,CochaYarina,125
m, 16 Jul 1985 (fl d), Vdsquezet al. 6665 (MO, NY). MA-
DRE DE DIos: Tambopata,Cusco Amaz6nico, 200-220 m,
19 Jun 1989 (fl d), Phillips et al. 468 (F, MO). SAN
MARTiN:Lamas, Alonso de Alvarado,Fundo las "Flores,"
800-900 m, 11 May 1973 (fl d), Schunke6231 (HBG).
BRAZIL. ACRE:Basin of Rio Purus,nearmouthof Rio
Macauhan(tributaryof Rio Yaco), Aug 1933 (fr), Krukoff
5281 (A, B, F, G, MO, NY, U); Manoel Urbano,Rio Purus,
Nova Olinda, 24 Nov 1996 (fr), Silveira et al. 1555 (MO).
AMAZONAS:Basin of Rio Jurua,near mouth of Rio Embira
(tributaryof Rio Tarauaca),8 Jun 1933 (fl ), Krukoff4714
(A, F, G, K, MO, NY, U, US); Limoeiro, Est. Ecol6gica do
Juami-Japurd, 26 Apr 1986 (fr), Cid Ferreiraet al. 7227 (F,
MO, NY, US). MATOGROSSO:Angustura,MachadoRiver
region, source of JatuaranaRiver, Dec 1931 (st), Krukoff
1565 (A, F, G, MO, NY, P, U, US). PARA:Itaituba,estrada
Santar6m-Cuiabd,BR 163, km 1221, 21 May 1983 (fl 6,
juv), Amaralet al. 1375 (GH, INPA,MO, NY). ROND6NIA:
Ji-Parand,May 1987 (fl ), Cid Ferreira9037 (INPA);Porto
Velho, 23-24 Apr 1987 (fl d), Cid Ferreira8900 (F, K, MO,
NY).
Local names. Costa Rica: aguacat6n.Colombia:
yiyiwa-ageyi (Mui). Ecuador:temachi. Peru: cunshi
moena, casha moena, moena blanca,moena de altura.
Brazil: louro abacate.
Endlicheriaformosa differs from most congeners
by its depressed-globose flowers with incurved to
erect tepals, but belongs in the E. brownianaspecies
group on the basis of dense papillosityinside flowers
and sessile stamens with ovate anthers.Also, in the
E. brownianaspecies group, very similarflowers are
found in E. paradoxa. Indeed,withoutthe scalariform
to percurrenttertiariesof E. formosa to contrastthe
reticulatetertiaryvenation of E. paradoxa, it would
be questionablewhetherlargerflowersize in the latter
is sufficient to separatethe two species. Outside of
the E. browniana species group, only E. robusta of
the E. sericea species group has similar depressed-
globose flowers.
Flowers from the Andean uplands and Central
America tend to be larger,more strongly depressed,
and more densely pubescentthanthose from lowland
Amazonia. Leaf size and textureis also variable,but
as with flowers, variationappearsto be continuous.
46 FLORA NEOTROPICA

7----
-~

__ , .-. . . .- . . . .

AV!

,. ':es y .
... ....d .......-t. xm,

FIG. 13. Distributionof Endlicheriaformosa and E. paradoxa.

16. Endlicheria paradoxa Mez, Jahrb.Konigl. Bot. soon lost, all vein ordersraised, theirprominencede-
Gart.Berlin. 5: 114. 1889. Type. Peru.Cajamarca: creasing with rank;secondary veins 6-8 per side, +
SantaCruz,Jun 1865 (fl d), Pearce s.n. (holotype: evenly spaced, slightly more distant aroundmidlam-
K 59; isotype: K 60). ina, ascending at 50-60?, arcuate, distal pairs loop-
connected; tertiaries reticulating between secondar-
Trees to 18 m. Branchlets stout, midway along
ies. Staminate inflorescences evenly spaced along
flush 5-7 mm diam., angular, sparsely strigillose,
to cm
soon glabrous, the surface clearly exposed, dark currentflush in the axils of foliage leaves, 10
brown, the hairs short, to 0.15 mm, straight, ap- long with 6 lateral branches,the highest order dicha-
pressed, greyish white; terminalbuds plump, 0.8 X sial, lax, the flowers distant,the axes sparsely strigil-
0.5 cm, grey-sericeous. Leaves alternate,widely and lose, distal branches and pedicels more densely so;
evenly spaced along currentflush; petioles slenderor bracts and bracteoles caducous by anthesis, lanceo-
robust, to 4 cm long and 0.3-0.6 cm diam., semi- late, grey-sericeous; pedicels terete, to 7 mm long,
terete to canaliculate,the indumentas on branchlets; those supportingsecondary flowers slightly shorter.
laminae coriaceous, plane, elliptic to obovate, 10-35 Flowers depressed-globose,to 7 mm diam., sparsely
X 6-15 cm, the base acute to cuneate, briefly decur- grey-strigilloseoutside;receptacleinfundibuliform,1
rent, the apex roundedor obtuse, acumen lacking or x 3 mm, rusty papillose inside. Tepals chartaceous,
mucronate for up to 0.3 cm, the margins minutely ovate to triangular,0.6 X 0.6 mm, inflexedat anthesis,
recurved throughout; upper surface olive-brown, the androeciumincluded. Stamens of whorls I and II
waxy, shining, the primary to fourth-orderveins sessile, 0.6 mm tall, the anthersovate, 0.5 X 0.5 mm,
prominent,conspicuousagainstthe lamina;lower sur- glabrous, the apex apiculate,the connectives sharply
face sparsely strigillose, the hairs as on branchlets, prolongedbetween the 2 locelli, these obliquely hem-
SYSTEMATICTREATMENT 47

ispherical, introrse-latrorse,the filamentsligulate, as posed, dark brown to black, the hairs very short, to
broad as anthers,rusty papillose; whorl III stamens 0.1 mm, appressed;terminalbuds slender, 3 X 0.6
sessile, 0.6 mm tall, the anthersovate, 0.5 X 0.5 mm, mm, densely chalky-white sericeous. Leaves alter-
erect,locelli 2, extrorse-latrorse,the filamentsbroader nate, widely and evenly spaced along currentflush;
than anthers,rusty papillose, the basal glands absent; petioles slender, to 1.5 X 0.2 cm, canaliculate,the
whorl IV wanting;pistillode wanting.Pistillateinflo- indumentas on branchlets;laminae membranaceous
rescence with indument, color, and branchingas in or chartaceous,plane,elliptic-ovate,10-17 X 3-8 cm,
staminateplants, the flowers smaller,slightly deeper; the base obtuse to acute, attenuate,the apex acute,
stamens sterile, smaller;ovary glabrous;style stout, acuminatefor up to 2.5 cm, the marginsminutelyre-
weakly distinguishedfrom ovary; stigma discoid, 0.2 curved throughout;upper surface blackish green to
mm diam. Maturefruitsunknown.Youngfruitsbome darkor olive-brown,minutelypunctulate,the primary
on claviform pedicels of 1 X 0.4 cm; cupules hemi- to fourth-orderveins raised, their prominence de-
spherical,0.5 X 1.5 cm, glabrousinside and outside, creasing with rank;lower surfaceglabrousexcept for
the marginsundulate;drupes ellipsoid, to 1.5 X 0.7 barbellatetufts in the axils of the sparsely strigillose
cm. midrib and secondary veins, all vein orders raised,
Distribution (Fig. 13) and ecology. Medium- their prominence decreasing with rank; secondary
sized trees known only from lower montane forest veins 4-6 per side, + evenly spaced, slightly more
around 1900-2000 m in Cajamarca,Peru. Flowering distantaroundmidlamina,patent,divergingat 70-85?
material collected in July and with young fruits in (more obtusely aroundmidlamina),abruptlyascend-
October. ing after midcourse, brochidodromous; tertiaries
roughly horizontal, between secondaries straightto
Additional specimens examined. PERU. CAJA- once-forked.Staminateinflorescencesevenly spaced
MARCA: San Ignacio, Chirinos, vic. of Pacasmayo, 1900-
along currentflush in the axils of foliage leaves, to 5
2000 m, 2 Oct 1997 (fl Y, frjuv), Camposet al. 4480 (MO),
cm long with 5 lateral branches,branchorders 2-3,
(fl Y), Camposet al. 4490 (MO).
the highest order dichasial, lax, the flowers distant,
Endlicheria paradoxa has been rediscoveredal- the axes sparselystrigillose;bractsandbracteolesca-
most 150 years after the mid-19th century type ma- ducous by anthesis,narrowlyovate to lanceolate,the
terial was collected. The additionalcollections, Cam- indumentas on axes; pedicels terete, to 5 mm long,
pos et al. 4480 and Camposet al. 4490, have smaller those supportingsecondary flowers slightly shorter.
leaves than the type, but depressed-globoseflowers Flowers hypocrateriform,3.5 mm diam., sparsely
and reticulate tertiaries refer them to E. paradoxa. grey-strigilloseoutside, the indumentdensertowards
Widespreadand frequentlycollected E. formosa has tepals; receptacle infundibuliform, 1 X 1 mm,
similardepressed-globoseflowers,but these aremuch sparsely grey-pilose inside, the hairs straight,erect.
smaller, and its tertiary veins are straight to once- Tepalsmembranaceous,ovate, 1.2 X 0.8 mm, spread-
forked ratherthan reticulate.Mez's (1889) isolation ing to recurvedat anthesis,the innersurfaceglabrous
of E. paradoxa in monotypic subgen. Hemiajoueais except for the minutely papillose tips. Stamens of
based on supposedsterilityof whorl III stamens.Sta- whorls I and II broadlystipitate,0.6 mm tall, the an-
minateflowersareknownonly fromthe type material, thersbroadlyovate, 0.3 X 0.5 mm, glabrous,the apex
wherein,as Kostermans(1937) also noted,all stamens apiculate, the connectives prolonged between the 2
appearto be fertile with well-developedlocelli. locelli, these obliquely hemispherical, introrse-
latrorse,the filamentslaminar,slightly narrowerthan
anthers, glabrous; whorl III stamens columnar,0.6
17. Endlicheria dysodantha (Ruiz & Pav.) Mez, mm tall, the anthersovate,0.3 X 0.5 mm, erect,locelli
Jahrb.Konigl. Bot. Gart.Berlin 5: 118. 1889. Lau- 2, extrorse-latrorse,the filamentsas broadas anthers,
rus dysodantha Ruiz & Pav., Fl. Peruv. 4: plate laminar,glabrous, the basal glands sessile, globose,
355. 1802. Goeppertiadysodantha(Ruiz & Pav.) relatively large, filling the space between filaments;
Nees, Linnaea 21: 514. 1848. Type. Peru. Huan- whorl IV wanting;pistillode filiform. Pistillate inflo-
uco: Macora,without date (fr), Ruiz & Pav6n s.n.
rescence with indument,color, and branchingas in
(lectotype: MA-n.v., designated by Kostermans, staminate
plants,the flowerssimilarin size andshape;
1937; isolectotypes:B-n.v., BM-n.v.)
stamens sterile, smaller;ovary glabrous,ovoid; style
Trees to 10 m. Branchletsslender,midway along slender,distinct from ovary; stigma discoid, 0.3 mm
flush 2-3 mm diam., distally weakly angular,soon diam. Fruitsborne on narrowlyclaviformpedicels of
terete,sparselygrey-strigillose,the surfaceclearlyex- up to 1.5 X 0.4 cm; cupules infundibuliform,to I x
48 FLORA NEOTROPICA

1.5 cm, glabrousinside and outside, the marginsen- (MO, NY, U), 370-450 m, 17 May 1993 (fr), Vargaset al.
tire; drupesellipsoid, to 2 X 1.4 cm. 2478 (MO,NY).

Distribution (Fig. 14) and ecology. Small trees Local names. Peru: moenito amarillo, palometa
or shrubs from inundatedand well-drained soils in micuna.
SW Amazoniaand adjacenteasternAndeanfoothills.
Endlicheria dysodantha is immediately recog-
Floweringfrom Juneto February,fruitsavailableyear
nized by its dark-dryingsubglabrousleaves with bar-
round.
bellate tufts of hairs in the axils of secondary veins
Representative specimens examined. ECUADOR. below. Close relationshipwith E. paniculata and al-
NAPO:Aguarico, SamonaYuturi,Rio Napo, 200 m, 11 Nov lies is suggested by stipitatewhorl I and II stamens
1991 (fl 6), Neill & Rojas 9950 (HBG, MO, NY); Aniangu, and pilose indument inside flowers. This has weak
ParqueNacionalYasuni, 14 Jul 1982 (fr), Luteynet al. 8680 molecular support (Fig. 2), but I prefer to place E.
(F, MO, NY). PASTAZA:Ceilan, Path from Ceilan to Rfo
dysodanthaclose to the E. brownianaspecies group.
Conoaco on S side of Rio Curaray,200 m, 7 Jun 1980 (fr),
Like other species therein, in E. dysodanthaall an-
Brandbyge& Asanza 31785 (MO); Pozo Villana2 de Arco,
2 km del pueblo de Villano,400 m, 3 Dec 1991 (fl 9), Tipaz
thers are ovate with obliquely hemisphericallocelli,
et al. 581 (F, HBG, MO, NY). and slender claviform pedicels support shallowly
PERU. Cuzco: Paucartambo,Pilcopata,Villa Carmen, hemisphericalcupules.Its vestitureof extremelyshort
720 m, 9 Feb 1975 (fr), Plowman & Davis 5098 (F); Rfo (0.1 mm) appressedhairs is also characteristicof the
Mapituriani,La Convenci6n,CordilleraVilcabamba,670 m, E. brownianaspecies group.
31 Jul 1968 (fl 6), Dudley 11493 (F, MO, NY). HuANuco:
Codo de Pozuzo, 300 m, 22 Oct 1982 (fl 9, fr juv), Foster
9394 (MO); Pachitea,Pucallpa, Sira Mtns., 260-400 m, 31
May 1988 (fr), Wallnofer17-31588 (MO). JUNiN: Rio Ne- 18. Endlicheria tomentosa Chanderbali, sp. nov.
gro to Satipo, 400 m, 17 Aug 1960 (fl 9, fr), Woytkowski Type. Peru. Cajamarca, Huarango, El Triunfo
5829 (G, MO). LORETO: Previsto,420 m, Oct 1962 (fl 6), (propertyof EdilbertoDelgado), 1500-1800 m, 13
Woytkowski7593 (GH, K, MO, US); Rio Santiago, s.d. (fl Jul 1996 (fl d), Campos et al. 2938 (holotype:
6), Tessmann3999 (F, G, NY). MADRE DE DIos: Manui, MO; isotypes: F, G, HBG, MO, NY, U). Fig. 15
Parque Nacional Manui,Cocha Cashu Station, Rio Manu,
300-400 m, 17-24 Aug 1974 (fl 6), Foster 3371 (F, HBG, A Endlicheriaedysodanthaeramuliset foliis subtuspilis
MO, NY); Tambopata,Explorer'sInn Tourist camp at jct. erectisstatimdiagnoscenda.
of Rios La Torreand Tambopata,SW of PuertoMaldonado,
CapironaTrail, 600 m, 25 Jan 1989 (fr), Smith et al. 1599 Trees to 6 m. Branchletsslender, midway along
(F, G, K, MO, NY, US). PUNO:Ridge betweenRio Candamo flush 2-3 mm diam., distally weakly angular,soon
and Rio Guacamayo,40-600 m, 22 May 1992 (fr), Gentry terete, densely tomentose, the surface barely visible,
et al. 76951 (MO). SANMARTiN: Mariscal Caceres, Toca- the hairs short, to 0.3 mm, straightto crooked, erect
che Nuevo, Quebradade Huaquisha,Rio Huallaga, Puerto to ascending,greenishyellow; terminalbuds slender,
Pisana, 15 Mar 1971 (fr), Schunke4766 (F, G, INPA, MO, 3 X 1 mm, densely pubescent, the hairs as on bran-
NY), 400-450 m, 29 Jun 1974 (fl 9, fr), Schunke 7053 chlets, ascending.Leaves alternate,widely andevenly
(HBG, MO). UCAYALI: Purns, Rio Curanja,circa la co- spaced along currentflush; petioles slender, to 1 X
munidadnativa de Colombiana,250 m, 24 Feb 2000 (fr),
0.1 cm, terete,the indumentas on branchlets;laminae
Graham& Schunke1099 (MO).
membranaceousor chartaceous,plane, narrowellip-
BRAZIL. AcRE: CaramariAmazonas,Rio Jurua,Lago
da Cigana, 150 m, 22 Aug 1986 (fr), Croat 62517 (HBG, tic, 6-10 X 1.5-3 cm, the base acute, briefly decur-
MO); Tarauaca,Rio Tarauaca',Seringal Tamandare,Colo- rent, the apex acute, acuminatefor up to 1 cm, the
caqao SantaMaria, 25 Dec 1995 (fr), Ehringhauset al. 396 marginsminutelyrecurvedthroughout;uppersurface
(MO). AMAZONAS: Basin of Rio Jurua,near mouth of Rio greyish green to olive-brown, waxy, the primary to
Embira(tributaryof Rio Tarauaca),12 Jun 1933 (fl 6), Kru- fourth-orderveins raised, their prominencedecreas-
koff4767 (A, F, G, K, MO, NY, U); Rio Purus,PontoAlegre, ing with rank; lower surface moderatelytomentose,
8 Apr 1904 (fr), Huber s.n. (NY). the hairs 0.3 mm long, crookedly erect; denser and
BOLIVIA. COCHABAMBA:Prov.Carrasco,Vallede Sa- formingbarbellatetufts in the axils of the midriband
jta, Estacion Piscicola Pirahiba,215 m, 21 Mar 1995 (fr),
secondaries, all vein ordersraised, their prominence
Ritter 1706 (MO). LA PAZ: San Antonio, Dec 1907 (fl 6),
decreasing with rank; secondary veins 4-6 per side,
Buchtien1986 (US); Prov.Larecaja,Tuiri (nearMapiri,Rio
? evenly spaced, slightly more distant aroundmid-
Mapiri), 490-750 m, 12-30 Sep 1939 (st), Krukoff10787
(A, F, MO); Mapiri, May 1886 (fr), Rusby 2671 (F, NY). lamina, ascending at 50-600 (more obtusely around
SANTA CRUZ: Prov. Ichilo, ParqueNacional Ambor6, 0-2 midlamina),arcuate,distal pairs loop-connected;ter-
km SW of El Carmen,360 m, 9 Nov 1990 (fl 6), Nee 39834 tiaries laxly reticulating between secondaries. Sta-
SYST'EMATICTREATMENT
'LI ..14. ...arnilr,ad..aahooof
.yoata
.nlcei
Ditibto
minate inflorescences evenly spaced along current
flush in the axils of foliage leaves, to 5 cm long with
3 lateralbranches,branchorders 2-3, the highest or-
der dichasial, lax, the flowers distant, the axes
sparsely grey-strigillose; bracts and bracteoles cadu-
cous by anthesis,lanceolate,the indumentas on axes;
pedicels terete, to 3 mm long, those supportingsec-
ondary flowers slightly shorter. Flowers hypo-
crateriform, 3.5 mm diam., sparsely grey-strigil-
lose outside; receptacle infundibuliform,1 X 1 mm,
densely pubescent inside, the hairs silvery, straight,
rigidly erect. Tepals chartaceous,ovate, 1.2 X 1 mm,
spreading to recurved at anthesis, the inner surface
sparsely silvery strigillose, the margins and apex
inside minutely papillose. Stamens of whorls I and
II broadly stipitate, 0.6 mm tall, the anthers
broadlyovate, 0.3 X 0.5 mm, glabrous,the apex apic-
ulate,the connectivesprolongedbetween the 2 locelli,
these obliquely hemispherical, introrse-latrorse,the
filaments laminar, narrowerthan anthers, glabrous;
whorl III stamens sessile, 0.6 mm tall, the anthers
49
Si-*kr .r 0s |f 0nt o 9s,v jT 0 X ; 7 bb!; X , -
e
E. to etoa
ovate, 0.3 X 0.5 mm, erect, locelli 2, extrorse-latrorse,
the filamentsbroaderthan anthers,ligulate, glabrous,
the basal glands sessile, relatively large, filling the
space between filaments;whorl IV wanting;pistillode
fusiform. Pistillate plants unknown.
Distribution (Fig. 14) and ecology. Known only
from the type material,taken from a small tree found
flowering in July in the lower montaneforests of Ca-
jamarca,Peru.
Flowers of Endlicheriatomentosaare identical to
those of E. dysodantha,and in both species the axils
of secondary veins on the underside of dark-drying
leaves are barbellate.However,in E. tomentosathese
barbellatetufts are all but lost amongst a dense erect
indumentum,whereas in E. dysodanthathey are con-
spicuous against the backgroundof sparse appressed
hairs elsewhere on the lower leaf surface.Laxly retic-
ulate tertiariesin E. tomentosa also contrastwith the
scalariformpatternformed by the tertiariesof E. dy-
sodantha. If E. tomentosais correctlyassigned to the
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SYSTEMATICTREATMENT 51

E. brownianaspecies group, it alone has a vestiture to recurved at anthesis, the inner surface papillose
of erect hairs. However,this may not be as uniqueas neartips, otherwiseglabrous.Stamensof whorlsI and
it seems since the allies of the E. brownianaspecies II stipitate,0.6 mm tall, the antherstransverselyob-
groupin Rhodostemonodaphne(e.g., R. laxa) all have long, 0.3 X 0.4 mm, glabrous, the apex truncateor
erect hairs. emarginate,the connectiveslevel with or reducedbe-
tween the 2 locelli, these suborbicular,introrse,the
filaments laminar, much narrower than anthers,
19. Endlicheria arunciflora (Meisn.) Mez, Jahrb. sparsely grey-villose; whorl III stamens stipitate,0.6
Konigl. Bot. Gart. Berlin 5: 131. 1889. Ampelo- mm tall, the anthersequal to outer whorls, erect, lo-
daphne aruncifioraMeisn., DC. Prodr.15(1): 81. celli 2, extrorse-latrorse,the filamentsnarrowerthan
1864. Type. Venezuela. Amazonas: Rio Negro, anthers, laminar, sparsely grey-villose, the basal
San Carlos, 1853-1854 (fl d), Spruce 3061 (lec- glands sessile, globose; whorl IV wanting;pistillode
totype, designated by Kostermans,1937: K; iso- filiform. Pistillate inflorescenceunknown,old pistil-
lectotypes: B-n.v., BM-n.v., K, NY-n.v., P-n.v.). late flowers glabrous,stamens sterile, smaller;ovary
glabrous, ovoid; style stout, weakly distinguished
Treesto 10 m. Branchletsslenderto stout,midway from ovary; stigma broadly tri-lobed, 0.6 mm diam.
along flush 3-6 mm diam., terete,densely pubescent, Fruits borne on terete pedicels of up to 5 X 3 mm;
the surface barely visible to concealed by the indu- cupules hemispherical,to 0.7 X 1.5 cm, glabrousin-
ment cover, the hairs relatively long, to 0.6 mm, side and outside, the marginsentire;drupesellipsoid,
straightto crooked, erect, reddish or a few grey; ter- to 2 x 1 cm.
minal buds plump, 5 X 3 mm, densely rusty pubes-
cent, the hairs as on branchlets,straight,appressedto Distribution (Fig. 14) and ecology. Medium-
ascending. Leaves closely spiraled at tips of current sized trees in flooded forests of the upperRio Negro
flush; petioles robust,to 2 X 0.3 cm, semi-terete,the area in Brazil and adjacentVenezuelaat 100-200 m.
indumentas on branchlets;laminae coriaceous, sub- Flowering July to October,fruits availablefrom No-
bullate,lanceolateto oblong, or elliptic, 10-25 X 2-8 vemberto March.
cm, the base obtuse to rounded, the apex obtuse to
acute, acuminatefor up to 1.5 cm, the marginsmin- Additional specimens examined. VENEZUELA.
utely recurved throughout;upper surface green to AMAZONAS: Rio Negro,CerroCucuy,2 Mar 1944 (fr),
Baldwin3211 (A, MO, US); Rio Casiquiare,CafnoPimichin,
olive-brown,minutely punctulate,the midrib promi-
128 m, 6-19 Jul 1969 (fl 6), Buntinget al. 4079 (MO, U);
nent, the secondaries impressed, the higher-order lower Rio Baria, 80 m, 22-23 Jul 1984 (fl 6), Davidse
venationimmersed,inconspicuousagainstthe lamina; 27595 (MO, NY, U); San Carlos de Rio Negro, 120 m, 16
lower surface sparsely pubescent, the hairs 0.6 mm May 1979 (fr), Liesner 7466 (MO, VEN); Atabapo, Nov
long, sprawlingto weakly ascending,rusty,denseron 1989 (fr), Marin 438 (MO); Rio Yatua,2 Dec 1984 (fl Y, fr
main veins, soon lost, all vein orders raised, their juv), Stergios & Aymard 7524 (MO, NY); Rio Orinoco,
prominence decreasing with rank; secondary veins Cano Tama-Tama,125-150 m, 23 Jun 1959 (fr), Wurdack
14-20 per side, ? evenly spaced, slightly more dis- & Adderley43153 (NY); Rio Pacimoni-RioYatua,along Rio
tant around midlamina,patent, diverging at 70-85? Yatuabetween mouth and Laja Catipan,110-120 m, 13 Jul
(more obtusely aroundmidlamina),abruptlyascend- 1959 (fl d), Wurdack& Adderley43435 (F, K, NY, U, US).
BRAZIL.AMAZONAS: Rio Negro, Rio Curicuriary,7
ing after midcourse, brochidodromous,or strongly
Sep 1979 (fl d), Kubitzkiet al. 79-172 (G, HBG, INPA),
loop-connected by midlamina; tertiaries oblique to
18 May 1973 (fr), Silva et al. 1707 (HBG, INPA);Caman-
midrib, between secondaries straightto forked. Sta- aus, Rio Miua, 30 Oct 1978 (fl 6), Nascimento826 (HBG,
minateinflorescencesdistally clusteredin the axils of NY); Rio Curicuriari,13 Jul 1979 (fl d), Poole 1972 (HBG,
cataphylls, to 20 cm long with 15 lateral branches, K, MG, MO, NY).
branchorders 3-4, the highest orderbotryoid, or ir-
regular, lax, the flowers loosely crowded, the axes The subglabrous inflorescences of Endlicheria
sparsely rusty tomentose, the hairs restrictedto first- aruncifloraoccur in only two other species of Endli-
and second-orderbranches,soon lost; bractsandbrac- cheria, both also of the Ampelodaphnespecies group.
teoles persistentat anthesis,lanceolate,sparselyrusty Of the two, Endlicheriaverticillatahas prominentter-
tomentose,glabrescent;pedicels terete,to 2 mm long, tiaries above, i.e., it belongs to the E. bracteata sub-
those supportingsecondary flowers slightly shorter. group, while E. arachnocomehas unique arachnoid
Flowers campanulate,2 mm diam., glabrousoutside; indumentand nevershows the brochidodromytypical
receptaclecyathiform,0.6 x 0.6 mm, glabrousinside. of E. arunciflora.Brochidodromyappearsto be un-
Tepalschartaceous,ligulate, 1.3 X 0.6 mm, spreading stable in E. arunciflorasince the basalmost second-
52 FLORANEOTROPICA

aries sometimes lack loop connections (e.g., Poole apex. Stamens of whorls I and II stipitate, 0.6 mm
1972). Even so, E. arunciflorais easily distinguished tall, the antherstransverselyoblong, 0.3 X 0.4 mm,
from E. arachnocome by its rusty tomentose bran- glabrous, the apex truncateor emarginate,the con-
chlets. nectives level with or reducedbetween the 2 locelli,
these suborbicular,introrse, the filaments laminar,
much narrowerthan anthers, sparsely grey-villose;
whorl III stamens stipitate,0.6 mm tall, the anthers
20. Endlicheria arachnocome Chanderbali,sp. nov.
equal to outer whorls, erect, locelli 2, extrorse-
Type. Peru. Loreto: Maynas, Rio Nanay, Pun-
latrorse,the filamentsnarrowerthananthers,laminar,
chana, 90 m, 21 Apr 1993 (fl d), Rimachi 10532
sparsely grey-villose, the basal glands sessile, glo-
(holotype:MO; isotype: NY) Fig. 16
bose; whorl IV wanting;pistillode filiform.Pistillate
Species habitu cum Endlicheriae arunciflorae optime inflorescencewith indument,color, and branchingas
congruens,sed differtramuliiset foliis subtuspilis arach- in staminateplants, the flowers slightly deeper; sta-
noidespraeditis. mens sterile, smaller; ovary glabrous, ovoid; style
slender,distinctfrom ovary;stigmabroadlytri-lobed,
Treesto 12 m. Branchletsslenderto stout,midway reniform,0.6 mm diam. Fruitsborne on terete pedi-
along flush 3-6 mm diam., terete,densely pubescent, cels of up to 6 X 4 mm; cupules hemispherical,to
the surface barely visible to concealed by the indu- 0.5 X 1.3 cm, glabrousinside and outside, the mar-
ment cover, the hairs relatively long, to 0.6 mm, gins entire;drupesellipsoid, to 2 X 1 cm.
crooked, sprawling, greyish white; terminal buds
plump, 5 X 3 mm, densely pubescent, the hairs Distribution (Fig. 14) and ecology. Small trees
straight, appressed to ascending, white. Leaves from seasonally flooded (tahuampa)lowland forests
closely spiraled at tips of currentflush; petioles ro- near Iquitos, Peru, at ca. 100 m. Flowering February
bust, to 3 X 0.4 cm, semi-terete,the indumentas on to October,fruitingNovemberto March.
branchlets;laminaechartaceous,plane, broadlyellip- Additionalspecimensexamined.PERU.Loreto:May-
tic, 15-30 X 5-11 cm, the base obtuse to acute, the nas,Mishana,SantaMariade Nanay,90 m, 2 Oct 1990(fl
apex acute, acuminate for up to 2 cm, the margins 6), Pipoly et al. 12727 (MO); Rfo Nanay, 130 m, 26 Jul
minutely recurved throughout;upper surface light 1984 (fl 6), Vasquezet al. 5434 (F, MO, NY), 140 m, 18
greyishgreento olive-brown,minutelypunctulate,the Aug 1988 (fl Y), van der Werifet al. 10207 (HBG, MO,
midrib prominent,the secondaries prominulous,the NY), MoronaCocha,120 m, 21 Mar1977(fr),Gentryet
higher-order venation immersed, inconspicuous al. 18525(F,MO);Iquitos-Nauta, km 10,90 m, 3 Apr1989
against the lamina;lower surfacesparselypubescent, (fl 6), Rimachi 9113 (MO); Mishuyacu, nearIquitos,100
the hairs 0.6 mm long, sprawlingto weakly ascend- m, Apr1930(fl 6), Klug1264(F,NY,US),May-Jun1930
1403 (F, NY, US); Iquitos,RIoItaya,Buena
ing, greyish white, denser on main veins, all vein or- (fl 6), Klug
Suerte, 122 m, 16 Nov 1986 (fr), Vasquez& Jaramillo8414
ders raised, their prominence decreasing with rank; (F, MO, NY); Rfo Momon, 95 m, 7 Feb 1985 (fr), Rimachi
secondary veins 9-11 per side, ? evenly spaced, 7739 (US); Puerto Almendras, 130 m, 22 Feb 1979 (st),
slightly more distantaroundmidlamina,ascendingat Gentryet al. 24873 (HBG, MO), 122 m, 16 Jul 1984 (fl 6),
50-60? (more obtusely aroundmidlamina),arcuateto Vdsquez& Jaramillo5253 (MO, NY), 14 Aug 1984 (fl 6),
archingafter midcourse,distal pairs loop-connected; Vasquez& Jaramillo5457 (MO), 2 Jun 1986 (fl 6), Vdsquez
tertiaries oblique to midrib, between secondaries & Jaramillo7620(MO),4 Jun1986(fl 6), Vdsquez &Jar-
straight to arched. Staminateinflorescences distally amillo 7647 (MO), 20 Jul 1982 (fl 6), Vdsquez& Jaramillo
clusteredin the axils of cataphylls,to 35 cm long with 3157 (MO); Punchana,90 m, 25 Feb 1993 (fl 6), Rimachi
10 lateral branches, branch orders 3-4, the highest 10402 (MO); Alto Nanay,5 Mar 1968 (fr), Simpson782 (G,
US); Requena, Sapuena,Bagazan-RfoUcayali, 130 m, 13
order botryoid, or irregular, the flowers loosely
Jan 1987 (fr), Vdsquez& Jaramillo8766 (MO).
crowded,the axes moderatelygrey-tomentose,the in-
dumentthinningdistally,lost by third-orderbranches; Material of Endlicheriaarachnocomewas previ-
bractsandbracteolespersistentat anthesis,lanceolate, ously distributedas E. arunciflora.The two species
the indument as on axes; pedicels terete, to 2 mm share subglabrousinflorescences and immersed ter-
long, those supporting secondary flowers slightly tiary venation above, and without noting the authen-
shorter.Flowers campanulate,2 mm diam., glabrous ticity of the arachnoidindumentof E. arachnocome
outside; receptacle cyathiform,0.6 X 0.6 mm, gla- its representativescould be thoughtdiseasedE. arun-
brous inside. Tepals chartaceous,ligulate, 1.3 X 0.6 ciflora. However, sprawling greyish white hairs and
mm, spreadingto recurvedat anthesis,the inner sur- eucamptodromous secondaries consistently distin-
face glabrous, the margins minutely papillose near guish this species from rusty tomentose and brochi-
SYSTEMATICTREATMENT 53

3.~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~

0.5 mm

0.3 -m
FIG. 16. Endlicheria arachnocome (Rimachi 10532). A. Habit. B. Leaf base below. C. Flower. D. Flower l.s. E.
Whorl I stamen seen from within. F. Whorl HI stamen seen from without.
54 FLORA NEOTROPICA

dodromousE. arunciflora.Further,with E. arachno- form,0.7 X 0.6 mm, slightlyconstrictedbelow tepals,

come knownonly fromPeruviantributariesof the Rio glabrousinside. Tepalsmembranaceous,ovate, 0.6 X
AmazonasandE. aruncifloraonly from the upperRio 0.5 mm, spreadingto recurvedat anthesis, the inner
Negro and Rio Orinoco aroundthe Brazil-Venezuela surfaceglabrousexcept for sparselypapillosetips and
border,these two riparianspecies appearto be allo- distal margins. Stamens of whorls I and II stipitate,
patric in differentdrainagesystems. 0.5 mm tall, the antherstransverselyoblong, 0.25 x
0.3 mm, glabrous, the apex truncateto emarginate,
the connectives level with or reduced between the 2
locelli, these suborbicular,introrse,the filamentslam-
21. Endlicheria arenosa Chanderbali,sp. nov. Type.
inar, narrowerthan anthers,glabrous;whorl III sta-
Brazil. Amazonas: Mun. Manaus, Estrada Ma-
mens stipitate, 0.5 mm tall, the anthers depressed-
naus-Caracaraf,km 130, IgarapeLages, 11 May
oblong, 0.2 X 0.3 mm, bowed towards the outer
1974 (fl d), Nelson & Lima P21108 (holotype:
whorls, locelli 2, extrorse-latrorse,the filamentsnar-
MO; isotypes: F, GH, HBG, INPA, K, NY, US).
rower than anthers,laminar, silvery strigose on the
Fig. 17
adaxial side, the basal glands stipitate,globose, rela-
Endlicheriaemacrophyllaeaccedens, ob pilis perbrevi- tively large, filling the space between filaments of
bus ab eo removendum. outer and inner whorls; whorl IV wanting;pistillode
fusiform. Pistillate inflorescence with indumentand
Trees to 5m. Branchlets usually stout, midway color as in staminate plants, but shorter and with
along flush (3-)5-8 mm diam., terete,densely tomen- fewer lateralbranches,the flowers similarin size and
tose, the surfaceconcealedby the indumentcover,the shape; stamens sterile, smaller;ovary ellipsoid, gla-
hairs relatively short, to 0.2 mm, erect, rust brownto brous;style slender,distinctfromovary,0.3 mm long;
dark red; terminal buds plump, 1 X 0.6 mm, rusty stigma tri-lobed, papillose, 0.25 mm diam. Fruits
tomentose, the hairs as on branchlets,erect, crisped bome on short terete pedicels of up to 2 mm, rarely
to crooked. Leaves closely spiraledat tips of current claviform and 1 cm long; cupules hemispherical,to
flush; petioles robust,to 3 X 0.4 cm, semi-terete,the 0.6 x 1 cm, glabrousinside and outside, the margins
indumentas on branchlets;laminae stiff chartaceous, entire;drupesovoid, to 1 X 0.8 cm.
plane to subbullate,obovate to elliptic, 10-27 X 3-
10 cm, the base acute to obtuse, the apex acute, acu- Distribution (Fig. 18) and ecology. Small trees
minatefor up to 2 cm, the marginsminutelyrecurved or shrubsfrom white sand (campina)forests in low-
throughout;upper surface olive to reddish brown, land Amazonia and savanna formations of north-
waxy, the midrib and secondaries prominulous,the western South America at 100-350 m. Flowering
higher-orderveins immersed;lower surface sparsely specimens have been collected throughoutthe year,
pubescent,the hairson the uppersurfaceof the midrib and fruitingmaterialfrom Septemberto April.
and secondaries crooked to crisped, dark red, soon
falling, those on the sides of the main veins and lam- Additionalspecimensexamined.COLOMBIA. AMA-
ZONAS: Corregimiento Puerto Rio
Santander, Caqueta,100-
ina light brown to translucent,0.2 mm long, straight,
350 m, 13-21 Feb 1997 (fl d ), Arbeldtez& Sueroke711 (U),
erect, persistent,denseron main veins, all vein orders
(st), Arbelaez & Sueroke 715 (U). VAUPES:Rio Kuduyari,
raised, their prominence decreasing with rank; sec- CerroYapoboda,450 m, 5-6 Oct 1951 (fl d), Schultes &
ondaryveins 8-11 per side, ? evenly spaced,slightly Cabrera 14356 (US), quarzite savannahnear headwaters,
more distant aroundmidlamina,patent, diverging at Apr 1953 (fr), Schultes & Cabrera20003 (US).
70-85? (more acutely towards apex), arching after VENEZUELA. AMAZONAS: Atabapo, 120 m, 8 Dec
midcourse, distal pairs loop-connected; tertiaries 1978 (fl i, fr), Huber & Tillett2975 (MO, NY); Rio Ori-
oblique to midrib, between secondaries once-forked noco, SerraYapacanabase, 100 m, 1 Jan 1951 (fr), Maguire
to straight.Staminateinflorescencesdistallyclustered et al. 30601 (NY); Sabanade Basil, Cano Chimoni,22 Feb
in the axils of cataphylls,to 25 cm long with 12 lateral 1989 (fr), Stergioset al. 13295 (HBG, NY); sabanade arena
branches, branch orders 2-4, the highest order bot- blanca al N de San Juan de Ucata, 70-80 m, 20 Oct 1989
(fl 5), Romero& Melgueiro2129 (GH, MO).
ryoid, lax, the flowers distant,the axes rusty tomen-
BRAZIL. AMAZONAS: ParqueNacional do Jad, Cam-
tose, the indument moderately dense, thinning dis- pina do Pataud,30 Aug 1998 (fl 5), Vicentiniet al. 1317A
tally; bracts and bracteoles persistent at anthesis, (INPA,MO), 2 Sep 1998 (fl Y), Vicentiniet al. 1359 (INPA,
lanceolate,the indumentas on axes; pedicels slender, MO), 6 Sep 1998 (fl d), Vicentiniet al. 1394 (INPA,MO);
terete, to 1 mm long, reduced to lacking below sec- Itapiranga,Rio Uatuma, 13 Aug 1979 (fl d), Cid Ferreira
ondary flowers. Flowers campanulate, to 1.5 mm et al. 266 (HBG, INPA, MO, NY, US), Igarap6Catitu, 20
diam., sparsely pubescent outside; receptaclecyathi- Aug 1979 (fl d), Cid Ferreira582 (HBG, MO, NY); Estrada
SYSTEMATICTREATMENT 55

FIG. 17. Endlicheriaarenosa (Nelson & Lima P21108). A. Habit. B. Inflorescence.C. Leaf base below. D. Flower
E. Flower ls. F. Pistillode. G. Whorl I stamen seen from within. H. Whorl III stamen seen from without.
56
Mr
A~~~~~~~~~~~..x
4 .......... ...............~
-IA1 DstibtinofEnliheiamarphll
aen,
da BR 17, km 20, 27 Jul 1961 (fl T), Rodrigues & Coe'lho
3033 (NY); Estradada PraiaDourada,Tarumizinho,29 Jun
1976 (fl Y), Monteiro 1246 (INPA); Estradado Ponta Ne-
gra-Tarumi,24 Oct 1966 (fr), Allen et al. 323 (HBG, INPA,
NY); EstradaManaus-Caracaraf,km 11, nearIgarapeLeao,
25 Aug 1959 (fl 6), Rodrigues & Chagas 1259 (NY), 10
May 1961 (fl 6), Rodrigues & Coilho 2564 (INPA, NY),
17 Mar 1967 (fr), Prance et al. 4678 (HBG, MG, NY, R),
km 130, 15 Feb 1974 (fr), Loureiroet al. s.n. INPA47996
(INPA), 18 Feb 1974 (fr), StewardetaaLP20357(GH, HBG,
INPA, K, MO, NY, R, S, US), km 125, 19 Feb 1979 (fr),
Coelho et al. 896 (INPA); EstradaManaus-Itacoatiara,kmn
181, 21 Dec 1966 (fr), Prance et al. 3817 (HBG, MG, NY);
Igarap6da Cachoeira Alta do Tarum&,6 Jun 1961 (fl 6),
Rodrigues & Chagas 2743 (INPA,NY), 12 Jun 1961 (fl d),
Rodrigues & Chagas 2783 (INPA, NY), 30 Jul 1962
(fl d), Rodrigues & Chagas 4558A (INPA, NY); Igarapdda
CachoeiraGrande,28 Aug 1962 (fl d), Rodrigues& Chagas
4611 (NY); Igarap6de Bolivia, 28 Dec 1955 (fr), Cotlho
s.n. INPA3206 (INPA,MG, NY); Igarapede RiachoGrande,
14 Aug 1957 (fl 6), Rodrigues 508 (INPA); Parque 10, 5
Jun 1956 (fl,2), Coilho s.n. INPA 3895 (INPA, MO, NY);
FLORA NEOTROPICA
Al~~~~~~~~~~~
17~~~~~~~~~:yk
ad
PontaNegra, 12 May 1961 (fl 3), Rodrigues& Coelho 2588
(INPA, NY), 24 Jan 1962 (fr), Rodrigues & Lima 4124 (G,
INPA, NY), 2 Dec 1960 (fr), Rodrigues et al. 1980 (INPA,
MO, NY), 24 May 1972 (fr), Coilho 160 (INPA);Reserva
FlorestalDucke, campinaranaAcari, 21 Aug 1979 (st), Ku-
bitzki & Poppendieck79-29 (HBG, NY); Rio Negro, May
1910 (fl 3), Ule 8848 (G, L, NY, MG, US); Nova Prainha,
Rio Aripuana,ProjetoRADAM/BRAZIL,SB-20-ZB, Ponto
02, 10 Aug 1976 (fl 6), Mota & Monteiros.n. INPA61281
(INPA), Ponto 10, 9 Jul 1976 (fl 6), Mota s.n. INPA60608
(INPA), Ponto 15, 15 Sep 1976 (fl 6), Ramosetal. s.n. INPA
62154 (INPA); Presidente Figueiredo, Rio Uatuma, UHE
Balbina, 26 Sep 1988 (fl 6), Lopes et al. 124 (INPA); Rio
Aripuana,53 km W along Trans-AmazonHwy., 27 Jun 1979
(fl 6), Calderon et al. 2704 (MPA, NY, MO, US); Serra
Araca, Rio Jauari, 29 Feb 1984 (fl juv), Rodrigues et al.
10494 (INPA, K, MO, NY). PARA:Campos do E de Faro,
Aug 1907 (fl 6), Ducke 8429 (MG); Camposdo Mariapiry,
Jul 1912 (fl 6), Ducke s.n. R 19970 (HBG, U); Campos do
Tigre, Dec 1919 (fl 6), Ducke s.n. R 11373 (U); Estradade
T. Santa, km 65, 16 Sep 1988 (fl 6), Soares 462 (INPA);
Oriximinu,Rio Mapuera,Campinadas Tres Ilhas, 19 Aug
SYSTEMATICTREATMENT
1986(fl d), CidFerreiraet al. 7803(F,GH,NY,MO,US),
25 Sep 1987(fr),Farneyet al. 2012 (MO),25 Nov 1987
(fr),CidFerreira9676 (INPA,MO).
Local name. Brazil: louro.
Endlicheriaarenosa is conspicuous in the Ampe-
lodaphne species group for its indument of stiffly
erect straighthairs only ca. 0.2 mm in length. Such
shorthairs are occasionally found in E. directonervia
but E. arenosa is best compared with species with
which it sharesimmersedtertiaryvenationabove and
rusty tomentose branchlets and inflorescences. Of
these, E. macrophyllahas similarflowers,but hairsat
least four times as long providea soft pilose indument
on the leaves below. Further,E. macrophyllaappears
to be a species of flooded forests while E. arenosa
has been found only on well-drainedsands.
Specimens from open savannas and scrub vege-
tation in Brazil (Rodrigues et al. 10494), Colombia
(Arbeldez& Sueroke711, 715) andVenezuela(Huber
& Tillett2975, Romero & Melgueiro 2129, Stergios
et al. 13295) departfrom the typical campinaforest
materialby their denser, darkerred indument.They
also maintaincrooked hairs on the veins below be-
yond leaf maturity,butthe shorterecthairsthatpersist
on older leaves are typical of E. arenosa.
22. Endlicheria macrophylla (Meisn.) Mez, Jahrb.
Konigl. Bot. Gart.Berlin. 5: 128. Ampelodaphne
macrophyllaMeisn., DC. Prodr. 15(1): 81. 1864.
Type. Brazil. Amazonas: Manaus, Apr 1851 (fl
d ), Spruce 1453 (lectotype, designated by Kos-
termans,1937: K; isolectotypes:B-n.v., BM-n.v.,
C-n.v., E, G, GH-n.v., L.-n.v., NY-n.v., P, W-
n.v.).
Treesto 15 m. Branchletsslenderto stout,midway
along flush 3-6 mm diam., terete, densely rusty to-
mentose, the surface barely visible to concealed by
the indumentcover, the hairs relatively short, to 0.5
mm, erect, crooked, matted;terminalbuds plump, 3
X 2 mm, the indument as on branchlets. Leaves
closely spiraled at tips of currentflush; petioles ro-
bust, to 3.5 X 0.5 cm, semi-terete, the indumentas
on branchlets;laminae stiff chartaceous,plane, ovate
to ovate-elliptic, 15-25 X 5-10 cm, the base rounded
to acute, the apex acute, acuminatefor up to 1.5 cm,
the marginsminutelyrecurvedthroughout;uppersur-
face greyish green to olive-brown, minutely punctu-
late, the midrib and secondaries prominulous, the
higher-order venation immersed, inconspicuous
against the lamina;lower surfacedensely pubescent,
57
the hairs 1 mm long, erect, straightor curvednearthe
tip, uniformlydistributed,all vein ordersraised,their
prominencedecreasingwith rank;secondaryveins 8-
11 per side, ? evenly spaced, slightly more distant
around midlamina, ascending at 50 60? (more ob-
tusely aroundmidlamina),arcuate,the lowermostpair
occasionally asymmetric(one or both merging with
marginat base), distal pairsloop-connected;tertiaries
oblique to midrib, between secondaries straight to
forked. Staminateinflorescencesdistally clusteredin
the axils of cataphylls,to 30 cm long with 12 lateral
branches, branch orders 3-4, the highest order bot-
ryoid, or irregular,the flowers loosely crowded, the
axes densely rusty villose, the hairs 1 mm long,
crooked;bracts and bracteolespersistentat anthesis,
lanceolate, greyish white villose; pedicels terete,to 1
mm long, those supportingsecondaryflowersslightly
shorter.Flowers campanulate,2 mm diam., sparsely
greyish white villose outside;receptaclenarrowlycy-
athiform, 1 X 0.3 mm, slightly constrictedbelow te-
pals, glabrous inside. Tepals membranaceous,ligu-
late, 1 X 0.6 mm (the inner whorl slightly broader),
ascending to recurved at anthesis, the inner surface
lightly papillose near the apex, otherwise glabrous.
Stamens of whorls I and II stipitate,0.6 mm tall, the
antherstransverselyoblong, 0.3 X 0.4 mm, glabrous,
the apex truncateto emarginate,the connectiveslevel
with or reducedbetween the 2 locelli, these suborbi-
cular, introrse,the filamentslaminar,much narrower
than anthers,sparselygrey-villose; whorl III stamens
stipitate, 0.6 mm tall, the anthers equal to outer
whorls, erect,locelli 2, extrorse-latrorse,the filaments
narrowerthan anthers,laminar,sparselygrey-villose,
the basal glands sessile, globose; whorl IV wanting;
pistillode filiform. Pistillate inflorescencewith indu-
ment and color as in staminateplants,but shorterand
with fewer lateralbranches,the flowers similarin size
and shape; stamens sterile, smaller; ovary glabrous,
ovoid; style slender, distinct from ovary; stigma tri-
lobed, 0.4 mm diam. Fruitsborneon shorttereteped-
icels of up to 1.5 X 1 mm, often subsessile and
clustered; cupules hemispherical, to 1 x 1.5 cm,
glabrous inside and outside, the margins entire;
drupesellipsoid, to 2 X 1.2 cm.
Distribution (Fig. 18) and ecology. Medium-
sized ripariantrees of centralandwesternAmazonian
lowlands ca. 100 m. Flowering from Septemberto
June,fruitsavailableby Decemberuntilthe following
August.
Representative specimens examined. COLOMBIA.
9 kmNEde
Vegadel CafnoPaufil,
CAQUETA: Rio Caqueta,
Araracuara,1 Dec 1988 (fr), Sanchez et al. 1870 (MO).
58
BRAZIL. AMAZONAS: Manaus, Igarapeda Cachoeira
Grande,28 May 1936 (fl d), Ducke 441 (A, F, K, MO, NY,
R, U, US); Rio Cuieiras,marginof Rfo Branchinho,12 Sep
1973 (fl d), Prance et al. 17785 (HBG, INPA, K, MO, NY,
R, US). PARA: Oriximina,Rio Trombetas,17 Jul 1980 (fl
d ), Cid Ferreira et al. 1601 (HBG, INPA, MO, NY); San-
tarem,Rio Curuauna,Prainha,19 Jan 1979 (fr), Santos 563
(INPA, MG). ROND6NIA: Bananeiras, Madeira-Mamor6
Railway, 13 Sep 1963 (fl d), Maguire et al. 56624 (HBG,
MO);PortoVelho,UHE de Samuel,Rio Jamari,14 Jun 1986
(fl d), Thomaset al. 5089 (F, INPA, K, MO, NY).
Local name. Brazil: louro.
Endlicheria macrophyllais the only member of
the Ampelodaphnespecies group with ovate leaves.
Such leaves are prevalentin available material, but
occasional ovate-ellipticforms reduce the diagnostic
value of leaf shapefor this species. Of close relatives,
E. multiflorashows similar soft pilose indumenton
the leaves below, and also has rusty tomentose
branchlets,but its minute and stipitate basal glands
are inconspicuous compared to the sessile globose
glands that surroundthe whorl III filamentsof E. ma-
crophylla. Further,E. multifiora appears to be re-
stricted to the Guianas while central AmazonianE.
macrophyllacan best be comparedto sympatricE.
arenosa. The latter bears a much shorterindument
and occurs in well-drainedwhite sand soils, whereas
E. macrophyllais a species of flooded forests.
Flowersfrom Rondoniaareless denselypubescent
and have more reflexedtepals than those from north-
em Amazonia.
23. Endlicheria multiflora (Miq.) Mez, Jahrb.K6n-
igl. Bot. Gart. Berlin 5: 130. 1889. Goeppertia
multifioraMiq., Stirp.Surin.Sel. 203. 1851. Type.
Suriname.Withoutlocality and date (fl d), Host-
mann & Kappler1163 (holotype:U; isotypes: B-
n.v., BM-n.v., G, K-n.v., MO, OXF-n.v., P, S, W-
n.v.)
Endlicheriavillosa Mez, Jahrb.Konigl. Bot. Gart.Berlin
5: 129. 1889. Type. Jamaica?Without date (fl d),
March s.n.? (syntypes: GOET-2 sheets).
Treesto 15 m. Branchletsslenderto stout,midway
along flush 3-6 mm diam., terete, densely rusty to-
mentose, the surface barely visible to concealed by
the indumentcover, the hairs relatively short, to 0.3
mm, crooked, erect, matted;terminalbuds plump, 3
X 2 mm, the indument as on branchlets. Leaves
closely spiraled at tips of currentflush; petioles ro-
bust, to 2 X 0.4 cm, semi-terete,the indumentas on
branchlets;laminae coriaceous to stiff chartaceous,
FLORA NEOTROPICA
plane to subbullate,obovate to narrowlyelliptic, 12-
20 X 4-10 cm, the base acute to rounded,the apex
obtuse to acute, acuminatefor up to 1.5 cm, the mar-
gins minutely recurved throughout;upper surface
dark green to olive-brown, minutely punctulate,the
midribandsecondariesprominulous,the higher-order
venationimmersed,inconspicuousagainstthe lamina;
lower surfacedensely pubescent,the hairsof the mid-
rib andsecondariesca. 0.6 mm long, crooked,densely
matted,rustyred, those on the laminagreyishto trans-
lucent, 1 mm long, erect, straightor curved near tip,
uniformly distributed, all vein orders raised, their
prominencedecreasingwith rank;secondaryveins 7-
9 per side, ? evenly spaced, slightly more distant
aroundmidlamina,patent,divergingat 70-85O(more
obtusely aroundmidlamina),archingaftermidcourse,
distal pairs loop-connected;tertiariesoblique to mid-
rib, between secondariesstraightto forked.Staminate
inflorescences distally clustered in the axils of cata-
phylls, to 25 cm long with 14 lateralbranches,branch
orders 3-4, the highest order botryoid, or irregular,
the flowers clustered,the axes densely rusty villose,
the hairs 1 mm long, crooked;bracts and bracteoles
persistent at anthesis, lanceolate, greyish white vil-
lose; pedicels terete, to 1 mm long, those supporting
secondary flowers slightly shorter.Flowers campan-
ulate, 2 mm diam., densely greyish white villose out-
side; receptacle narrowlycyathiform, 1 X 0.3 mm,
slightly constrictedbelow tepals, glabrousinside. Te-
pals membranaceous,ligulate, 1 X 0.6 mm (the inner
whorl slightly broader),spreadingto recurvedat an-
thesis, the inner surface lightly papillose near the
apex, otherwise glabrous.Stamensof whorls I and II
stipitate,0.6 mm tall, the antherstransverselyoblong,
0.3 X 0.4 mm, glabrous,the apex truncateto emar-
ginate, the connectiveslevel with or reducedbetween
the 2 locelli, these suborbicular,introrse,the filaments
laminar,much narrowerthan anthers,sparsely grey-
villose; whorl III stamens stipitate, 0.6 mm tall, the
anthers equal to outer whorls, erect, locelli 2,
extrorse-latrorse,the filamentsnarrowerthananthers,
laminar,sparsely grey-villose, the basal glands min-
utely stipitate,globose; whorl IV staminodialor ab-
sent, columnar;pistillode filiform. Pistillate inflores-
cence with indumentand color as in staminateplants,
but shorterand with fewer lateralbranches,the flow-
ers similarin size and shape;stamenssterile, smaller;
ovary glabrous, ovoid; style slender, distinct from
ovary; stigma tri-lobed, 0.4 mm diam. Fruits borue
on short, terete pedicels of up to 5 X 3 mm, often
subsessile and clustered;cupules hemispherical,to 1
X 1.5 cm, glabrous inside and outside, the margins
entire;drupesellipsoid, to 1.7 X 1.2 cm.
SYSTEMATICTREATMENT 59

Distribution (Fig. 19) and ecology. Medium- SURINAME. Nickerie R., Blanche Marie Falls, 20 Jun
sized trees of lowland riparianforests (5-100 m) in 1965 (fl d), Maas & Tawjoerans.n. LLB. 10928 (U); Re-
Guyana,Suriname,and borderingareasof Brazil and chter CoppenameR., 14 May 1963 (fr), WesselsBoer 1381
Venezuela.Flowering specimens collected from June (F, NY, U).
BRAZIL. PARU:Oriximind, Rio Trombetas, 29 Nov
throughNovember,fruits collected in all months but
1907 (fl 9), Ducke 8920 (U); Rio Mapuera, 12 Aug 1986
Septemberand January.
(fl d), Cid Ferreiraet al. 7652 (F, INPA, K, MO, NY).
Representativespecimensexamined.VENEZUELA.
BOLiVAR: Rio Venamo, 90 m, 31 Jul 1981 (fl d), Aymard Local names. Guyana: bastard silverballi. Suri-
et al. 372 (MO, NY); Rfo Uiri-yuk, 16 Aug 1962 (fl 6), name: siroewaballioenilebobandikoro(Arawak)
Maguire et al. 46713 (NY).
GUYANA. BERBICE:Berbice River KuruduniCk., 50 In the Ampelodaphnespecies group, Endlicheria
m, 30 Apr 1995 (fr), Mutchnick1243 (MO), Melissa Falls multiflorashares densely rusty tomentose branchlets
to the Gate, 120 m, 3 May 1995 (fr), Mutchnick1303 (MO). and inflorescence axes with E. arenosa, E. macro-
DEMERARA: DemeraraRiver,Sep 1889 (fl 6), Jenman4127 phylla, E. melinonii, and E. reflectens,but the minute
(K, US), 19 Mar 1923 (fl 9, fr), Persaud 59 (F, INPA, K, basal glands associatedwith its whorl IHstamenssug-
MO,NY, U). ESSEQUIBO: Cuyuni-MazaruniRegion, Ma-
gest affinitywith E. dictifarinosaand E. levelii. Grey-
zaruni River, Jun 1889 (fl 6), Jenman 5321 (K, NY, US);
Potaro-SiparuniRegion, IwokramaRainforestReserve, Es-
ish tomentose branchletsand inflorescence axes, and
sequibo River, 80 m, 13-27 Jul 1996 (fl 6), Chanderbaliet subsessile leaves with subcordatebases, distinguish
al. 153 (BRG, MO, US); RupununiRegion, TorobaroeCk., E. dictifarinosa from E. multiflora,but delimitation
145 m, 21 Nov 1987 (fr juv), Jansen-Jacobset al. 1118 (B, from E. levelii is not as straightforward.In the Suri-
K, NY, U). namese type of E. multiflora,most collections from

R EU

4 A ~~~~~~~~~~~
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~~~~~~~~~~~~~~~~~~~~~~~~~~~~W gr-;Wie

_v foK ~~~~~~~~~~~~~~~~~~~~~~~~~~
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; ~~~~~~~~~~~~~~~~~~~~~~~~
A . .'w.'.~:^l
-,. ....! ;-'i.

.,..~ ~
ofl. 9 Ditrbuio
EMlcei
FI. muf.r
E- lS, ; { . evein an E ;
-cSfnn

S c$--*. 1---
..;1 ,+ r +, , tS
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60
Guyana,and two collections from adjacentBolivarin
Venezuela, branchletsand inflorescence axes bear a
dense rusty tomentosecover of ca. 0.6 mm long hairs
whilst material from the upper Rfo Negro-Rio Ori-
noco type locality of E. levelii have a dense greyish
setose-villose indumentumof 1.2 mm long hairs. On
the basis thereforeof indumentcolor andlength,most
specimens can be assigned to either E. multifloraor
E. levelii without hesitation. Conceivablyintermedi-
ate conditions are found only in Cid Ferreira et al.
7652 and Ducke 8920 (both from northernPara in
Brazil),Maas & Tawjoerans.n. (fromSuriname),and
Maguire & Fanshawe 23481 and Hoffinann et al.
2029 (both from Guyana). In these specimens, new
leaves have sparse sprawlingrusty hairs resembling
those in the several sparselypubescent specimens of
E. levelii. Yet these 0.6 mm long rust-redhairs are a
magnitudeof order shorterthan the 1.2 mm greyish
hairs of the latter.Since these specimens differ from
E. levelii in both indumentlength and color, but from
typical E. multifloraonly in indumentdensity, they
are more parsimoniouslyassigned to the latter spe-
cies. With this recourseadopted,E. multifloraand E.
levelii appearallopatric with the former centered in
the Guianasand the latterin centralAmazonia.
AmpelodaphnedasyanthaMeisn., listed as a syn-
onym of Endlicheria multifloraby Mez (1889) and
Kostermans(1937), is an illegitimatename based on,
among others, Hostmann & Kappler 1163, the type
of Goeppertiamultiflora.The only synonym of En-
dlicheria multiflora accepted here, E. villosa, was
based on two unlabeled collections found by Grise-
bach among W. T. March'scollections from Jamaica.
As Endlicheriadoes not occurin JamaicaandtheAm-
pelodaphnespecies groupis entirelySouthAmerican,
Grisebach'sannotationsandMez's (1889) protologue
justifiablyexpress uncertaintyaboutcollector and lo-
cality. These sheets were certainlymisplaced among
March'sJamaicancollections but their source is still
unknown.If strongestresemblancewith Cid Ferreira
et al. 7652 andDucke 8920 is any clue, northernPara
in Brazil seems a likely locality.
24. Endlicheria levelii C. K. Allen, Mem. New York
Bot. Gard. 10 (5): 62. 1964. Type. Venezuela.
Amazonas: Rio Orinoco 10 km above mouth of
Rio Atabapo,CanioYagual, 150 m, 30 May 1954
(fl 1), Level 127 (holotype:NY).
Treesto 20 m. Branchletsslenderto stout,midway
along flush 3-6 mm diam., terete,densely pubescent,
the surfacebarely visible, the hairsrelativelylong, to
1.2 mm, straight,or crooked,erect,greyishwhite;ter-
FLORANEOTROPICA
minal buds plump, 3 X 3 mm, densely pubescent,the
hairs as on branchiets, straight, ascending. Leaves
closely spiraledat tips of currentflush;petioles slen-
der, to 3 X 0.3 cm, semi-terete,the indumentas on
branchlets;laminaecoriaceous to chartaceous,plane,
obovate to elliptic, 10-25 X 3-9 cm, the base acute,
the apex obtuse to acute, acuminatefor up to 1.5 cm,
the marginsminutelyrecurvedthroughout;uppersur-
face greyish green to olive-brown,minutely punctu-
late, the midrib and secondaries prominulous, the
higher-order venation immersed, inconspicuous
against the lamina;lower surface densely pubescent,
the hairs 1.2 mm long, rigid, ascending,greyish,soon
lost on lamina, persistentand denser on main veins,
all vein orders raised, their prominence decreasing
with rank;secondaryveins 7-11 per side, + evenly
spaced, slightly more distantaroundmidlamina,pat-
ent, divergingat 70-850 (more obtusely aroundmid-
lamina), arcuate,distal pairs loop-connected;tertiar-
ies oblique to midrib, between secondaries
once-forkedto straight.Staminateinflorescencesdis-
tally clusteredin the axils of cataphylls,to 20 cm long
with 18 lateralbranches,branchorders3-4, the high-
est order botryoid, or irregular,the flowers loosely
crowded, the axes densely greyish white villose, the
hairs 1 mm long, straightto crooked;bractsand brac-
teoles persistentat anthesis,lanceolate,the indument
as on axes; pedicels terete, to 2 mm long, those sup-
porting secondary flowers slightly shorter.Flowers
campanulate,2 mm diam., densely greyish white vil-
lose outside; receptacle cyathiform,0.6 X 0.4 mm,
slightly constrictedbelow tepals, glabrousinside. Te-
pals membranaceous,ligulate, 0.6 X 0.5 mm (the in-
ner whorl slightly broader),spreadingto recurvedat
anthesis, the inner surface lightly papillose near the
apex, otherwise glabrous.Stamensof whorls I and II
stipitate,0.6 mm tall, the antherstransverselyoblong,
0.3 X 0.4 mm, glabrous,the apex truncateto emar-
ginate, the connectiveslevel with or reducedbetween
the 2 locelli, these suborbicular,introrse,the filaments
laminar,much narrowerthan anthers,sparsely grey-
villose; whorl III stamens stipitate,0.6 mm tall, the
anthers equal to outer whorls, erect, locelli 2,
extrorse-latrorse,the filamentsnarrowerthananthers,
laminar,sparsely grey-villose, the basal glands min-
utely stipitate,globose; whorl IV staminodialor ab-
sent, columnar;pistillode filiform. Pistillate inflores-
cence with indumentand color as in staminateplants,
but shorterand with fewer lateralbranches,the flow-
ers similarin size and shape;stamenssterile, smaller;
ovary glabrous, ovoid; style slender, distinct from
ovary; stigma tri-lobed, 0.4 mm diam. Fruits borne
on short, terete pedicels of up to 5 X 3 mm, often
SYSTEMATICTREATMENT 61

subsessile and clustered;cupules hemispherical,to 1 Trees, 5-20 m. Branchletsslender to stout, mid-

X 1.5 cm, glabrous inside and outside, the margins way along flush 3-6 mm diam., terete, densely to-
entire;drupesellipsoid, to 1.7 X 1.2 cm. mentose, the surface barely visible to concealed by
the indumentcover, the hairs relatively short, to 0.3
Distribution (Fig. 19) and ecology. Medium-
mm, crooked, erect, matted, rusty to greyish white;
sized riparian trees from central and western Ama-
terminal buds plump, 3 X 2 mm, the hairs as on
zonia at 80-250 m. Floweringfrom April to October,
branchlets,ascending.Leaves closely spiraledat tips
fruits availableyear round.
of currentflush;petioles robust,to 1 X 0.4 cm, semi-
Representativespecimensexamined.COLOMBIA. terete, the indumentas on branchlets;laminae coria-
VAUPES:Rio Apaporis,Soratama, 250 m, 1 Sep 1951 (fl ceous to stiff chartaceous,plane, obovate, 11-25 X
6), Schultes & Cabrera 13849 (GH, U, US). 5-13 cm, the base abruptlyroundedto cordate, the
VENEZUELA.AMAZONAS: Atabapo,Rio Cunucun-
apex obtuse, acuminatefor up to 1.5 cm, the margins
uma,180-210m, 28-30 Jan& 6-8 Feb 1982(fr),Steyer-
minutely recurved throughout;upper surface light
market al. 126162(HBG,MO,NY);Atures,43 kmNEde
SantaBarbara delOrinoco,220 m, Apr 1990 Marin
(fr), 981 greyish green to reddishbrown,minutelypunctulate,
(MO);RioNegro,RfoBaria,80 m, 21 Jul1984(fl 6), Dav- the midrib and secondariesprominulous,the higher-
idse 27575 (MO,NY,US).BARINAS: RfoZulia,SantaBar- order venation immersed;lower surface moderately
baradeBarinas,21 Sep 1988(fl 6), Valverde &Penia(MO). pubescent,the hairs0.7 mm long, crookedon the mid-
BRAZIL.AMAZONAS:RioCuricuriari, IgarapeCariua, rib and secondaries,on the lamina straightand erect,
12 Jul 1979(fl 6), Maiaet al. 562 (HBG,INPA,K, MO, uniformly distributed, all vein orders raised, their
NY); Rio Negro,betweenManausand Sao Gabriel,Ilha prominencedecreasingwith rank;secondaryveins 7-
Acaburu,4 Apr 1979(fl 6), Maiaet al. 430 (HBG,MO, 11 per side, ? evenly spaced, slightly more distant
NY,US); Rio Solimoes,Sao Paulode Olivenqa,nearPal-
divergingat 70-85? (more
mares,11 Sep-26Oct 1936(fr),Krukoff 8096 (A, F, G, K, aroundmidlamina,patent,
MO,NY,U); RioTea,affluentedo RioNegro,12Jun1976 obtusely around midlamina), archingaftermidcourse,
(fl Y), Coelho460 (INPA);SerraAraga,Rio Jauari,28 Jul distal pairs loop-connected; tertiariesoblique to mid-
1985 (fr), Silva 181 (GH, K, MO, NY). MATOGROSSO: rib, between secondariesstraightto forked.Staminate
NucleoPioneirode Humboldt,28 Oct 1973 (fr),Berg & inflorescences distally clustered in the axils of cata-
StewardP19932(HBG,INPA,K, MO,NY);Serrado Ron- phylls, to 25 cm long with 12 lateralbranches,branch
cador,RioVau,11Oct1964(fr),Prance&Silva59385(GH, orders3-4, the highest orderbotryoidto irregular,the
MO,NY,US). flowers loosely crowded,the axes densely villose, the
BOLIVIA.SANTA CRUZ: Prov.Velasco,ReservaBiol- hairs 1 mm long, crooked, greyish white throughout,
6gicaEl Refugio,210 m, 23 May 1995(fl 6), Guilln &
a few reddish,or distally rusty;bractsand bracteoles
Chore3814 (MO), ibid., 27 Jan 1997 (fr), Carrionet al. 531
persistent at anthesis, lanceolate, densely greyish
(MO).
whitevillose;pedicels terete,to 1 mm long, those sup-
Local name. Venezuela:laurel de babilla. porting secondary flowers slightly shorter.Flowers
Endlicheria levelii is distinguishedfrom E. mul- campanulate,2 mm diam., densely greyish white or
tiflora primarilyby the greyish white indumentof its
rusty villose outside;receptaclenarrowlycyathiform,
branchletsand inflorescences (see discussion of the 1 X 0.3 mm, slightly constrictedbelow tepals, gla-
latter), and from E. dictifarinosa by its acute rather brous inside. Tepals membranaceous,ligulate, 1 X
than subcordateleaf bases. The type specimen and 0.6 mm (the inner whorl slightly broader),spreading
material from Bolivia and adjacentMato Grosso in to recurvedat anthesis,the inner surfacelightly pap-
Brazil have a dense indumentumon the leaves below, illose near the apex, otherwise glabrous.Stamens of
but hairs are restrictedto midribs below in all other whorlsI and II stipitate,0.6 mm tall, the antherstrans-
specimens. These less pubescent specimens are rem- versely oblong, 0.3 X 0.4 mm, glabrous, the apex
iniscent of E. bracteata whereinhairs can be equally truncateto emarginate,the connectives level with or
long, or longer, but leaves tend to dry blackish green reducedbetween the 2 locelli, these suborbicular,in-
and always show prominenttertiariesabove. trorse,the filamentslaminar,much narrowerthanan-
thers, sparsely grey-villose; whorl III stamens stipi-
tate, 0.6 mm tall, the anthersequal to outer whorls,
25. Endlicheria dictifarinosa C. K. Allen, Mem. erect, locelli 2, extrorse-latrodrse, the filaments nar-
New YorkBot. Gard.12 (3): 108. 1965. Type.Ven- rowerthananthers,laminar,sparselygrey-villose,the
ezuela. Bolivar:MaihiaRapidsalong Rio Paragua, basal glands minutely stipitate, globose; whorl IV
500-510 m, 1 Jan 1962 (fl 6), Steyermark 90535 staminodialor absent, columnar;pistillode filiform.
(holotype: NY; isotypes: G, U). Pistillate inflorescence unknown. Fruits borne on
62
short, terete pedicels of up to 5 X 3 mm, often sub-
sessile and clustered;cupules hemispherical,to 1 X
1.5 cm, glabrousinside and outside, the marginsen-
tire; drupesellipsoid, to 1.7 X 1.2 cm.
Distribution (Fig. 19) and ecology. Small to
medium-sizedripariantrees from SE Venezuelaand
the adjacentstateof Roraimain Brazil at ca. 100-800
m. Floweringmaterialcollected in January,May, and
from July to October, and fruiting specimens from
Novemberto May.
Representativespecimensexamined.VENEZUELA.
AMAZONAS: Rio Orinoco, Tama-Tama,125-150 m, 28 Jul
1959 (fl d), Wurdack& Adderley43647 (B, COL,F, G, NY,
W); Sierra Parima,vecindades de Simarawochi,Rio Mata-
cuni, 795-830 m, 18 Apr-23 May 1973 (fr), Steyernark
107391 (G, HBG, NY). BOLiVAR: Cedenlo,Icutd, cuenca
del Rio Nachare, 150 m, 1 Aug 1995 (fl d), Knab-Vispo&
Rodriguez479 (MO, VEN); Uriman,Rio Garoni,480 m, 31
Aug 1954 (fl d), Bernardi1626 (COL, G, NY).
BRAZIL. RORAIMA: Boa Vista, EstaqaoEcol6gica de
Maraca,19 May 1987 (fr), Lima 783 (E, INPA,K, MO, NY);
Rio Mucajai, river island, 17 Mar 1971 (fr), Prance et al.
11069 (HBG, K, MO, NY, US).
Local names and uses. Venezuela:laurel hormi-
guero, peroua-yek (Arecuna), wakamei (Ye'kwana),
Brazil:moroman(Uaica-Mucajai).Fruitsedible.
Endlicheria dictifarinosa shares greyish tomen-
tose branchletsand inflorescenceswith E. levelii but
is distinguishedby its subsessile leaves with subcor-
date bases. The hairs of the two also differ in length,
with those of E. dictifarinosa half as long as the ca.
1.2 mm hairs in E. levelii. Occasionally,distal inflo-
rescence branches,pedicels, and exterualflowerparts
are rustytomentose(e.g., Bernardi1626), or rustcol-
ored hairs are interspersedthroughoutthe dominant
greyish indumentum (e.g., Knab-Vispo & Rodriguez
479). These specimens appear marginalbetween E.
dictifarinosa and E. multiflora on the basis of indu-
ment color but do not show the soft pilose indument
that typically persists on lower leaf surfaces in the
latter. Their peculiar subsessile leaves are also un-
matched in E. multiflora. Leaves of similar shape
combine with adaxially prominentthird- and fourth-
order veins in E. bracteata, E. cocuirey, and E. ver-
ticillata.
26. Endlicheria melinonii Benoist, Bull. Soc. Bot.
France 75: 977. 1928 (nomen nudum), validated
in Arch. Botan.V: 63. 1931. Type.FrenchGuiana.
FLORANEOTROPICA
Cayenne, without date (fl d), Melinon s.n. (holo-
type: P; isotype: P).
Treesto 30 m. Branchletsslenderto stout,midway
along flush 3-6 mm diam., terete, densely rusty to-
mentose, the surface barely visible to concealed by
the indumentcover, the hairs relatively short, to 0.3
mm, crooked, erect, matted;terminalbuds plump, 3
X 2 mm, densely rustytomentose.Leaves closely spi-
raled at tips of currentflush; petioles robust,to 1 X
0.3 cm, semi-terete, the indument as on branchlets;
laminaechartaceousto coriaceous,plane, obovate,8-
15 X 3-7 cm, the base acute, the apex obtuse, acu-
minatefor up to 1 cm, the marginsminutelyrecurved
throughout;uppersurfacedarkgreen to olive-brown,
minutely punctulate, the midrib and secondaries
prominulous,the higher-ordervenationimmersed,in-
conspicuousagainstthe lamina;lower surfacedensely
rusty pubescent, the hairs ca. 0.5 mm long, erect,
straight,uniformlydistributed,all vein ordersraised,
their prominence decreasing with rank; secondary
veins 5-8 per side, ? evenly spaced, slightly more
distantaroundmidlamina,patent,divergingat 70-85?
(more obtuse aroundmidlamina),archingafter mid-
course, distal pairs loop-connected;tertiariesoblique
to midrib,between secondariesstraightto forked.Sta-
minateinflorescencesdistally clusteredin the axils of
cataphylls, to 11 cm long with 8 lateral branches,
branchorders 2-3, the highest order condensed, the
flowers clustered, the axes densely rusty villose, the
hairs0.7 mm long, crooked;bractsandbracteolesper-
sistent at anthesis, ovate, the indument as on axes;
pedicels wanting.Flowers campanulate,2 mm diam.,
densely rusty villose outside, the hairs sparseron te-
pals;receptaclecyathiform,1 X 0.4 mm, slightly con-
stricted below tepals, sericeous inside. Tepals mem-
branaceous, ovate, 1 X 0.6 mm (the inner whorl
slightly broader),erect at anthesis, the inner surface
glabrous,the marginsminutelypapillose. Stamensof
whorlsI andII stipitate,0.6 mm tall, the antherstrans-
versely oblong, 0.3 X 0.4 mm, glabrous, the apex
truncateto emarginate,the connectives level with or
reducedbetween the 2 locelli, these suborbicular,in-
trorse,the filamentslaminar,much narrowerthanan-
thers, sparsely grey-villose; whorl III stamens stipi-
tate, 0.6 mm tall, the anthersequal to outer whorls,
erect, locelli 2, extrorse-latrorse,the filaments nar-
rowerthananthers,laminar,sparselygrey-villose,the
basal glands sessile, globose; whorl IV wanting;pis-
tillode fusiform. Pistillate inflorescence with indu-
ment and color as in staminateplants,but shorterand
with fewer lateralbranches,the flowerssimilarin size
and shape; stamens sterile, smaller; ovary glabrous,
SYSTEMATICTREATMENT
ovoid; style slender, distinct from ovary; stigma tri-
lobed, 0.3 mm diam. Cupules hemispherical,sessile,
to 8 x 5 mm, glabrousinside andoutside, the margins
undulate,drupesellipsoid, to 1.5 X 1 cm.
Distribution (Fig. 20) and ecology. Tall trees
from flooded and nonflooded moist forests of NE
South America at ca. 200-450 m. Floweringmaterial
collected in August, September, and January,and
fruits in Februaryand September.
Additional specimens examined. VENEZUELA.
DELTAAMACURO: El Palmar,Rio Grande, 19 Aug-7 Sep
1964 (fl 6), Marcano-Berti361 (F, G, HBG, SPA, MO,
NY).
FRENCH GUIANA. Cayenne, s.d. (fl d), Patris s.n.
(G), Saul, La Fum6e Mtn. Trail, 200-300 m, 16 Sep 1984
(fr juv), Mori et aL 20927 (MO, NY), 220 m, 21 Jun 1988
(st), Gentry et al. 62990 (NY); Bassin du Sinnary,22 Feb
1989 (fr), Sabatier 2365 (MO, NY); Crique Passoura, 26
Sep 1992 (fr juv), Sabatier & Prevost 4073 (B, MO, NY);
Mont Atachi Bacca, Region de l'Inini, 420 m, 10 Jan 1989
.-^tsx55rl X*ir;tY; f EL
M\-
F.
S - Ro
I ~~~~~~~~~~~~~~~~~~~~~~~7
FIG. 20. Distributionof Endlicheriamelinonii E. reflectens,and E. bracteata.
63
(fr juv), Granville 10538 (B, MO, NY, U, US); Pistie de
Saint-Elie, 26 Sep 1991 (fl 9), Sabatier & Prevost 3819
(MO, NY, U).
SURINAME. Brokopondo, 21 Jan 1965 (st), van don-
selaar 2037 (MO); Rikanauprope Moengo, 2 Jun 1954 (st),
Lindeman 5943 (U), (st), Lindeman 6056 (U); Suriname
River, Wilhelminagebergte-MapaneCk., 1 Oct 1953 (st),
Lindeman4787 (NY, U); s.loc., 2 Mar 1957 (st), Heyligers
485 (U).
BRAZIL. AMAPA: Reserva INCR, Rio Falsino, 22-26
Aug 1983 (fl dc), Campbellet al. 14650 (NY).
Local names. Venezuela:laurel negro. Suriname:
harige pisie, pisie.
Kostermans (1937) placed Endlicheria melinonii
in synonymy underE. multiflora.The two sharerusty
tomentose indumentand obovate leaves but E. meli-
nonii is easily distinguishedby its sessile flowerswith
erect tepals. Further,stiff straight ca. 0.5 mm long
hairs on the lower leaf surface permitrecognitionon
vegetative grounds alone. Sterile specimens thus as-
; - Ux~~~~~~~~~~~~~~~~~~~~~~~~~
XesX----f - %
64 FLORANEOTROPICA

signed establish its presence in Surinameand reduce porting secondary flowers slightly shorter.Flowers
the severity of an otherwise striking disjunctionbe- campanulate,3 mm diam., sparselytawnypilose out-
tween French Guiana-NE Brazil and NE Venezuela. side, the hairs ascending;receptacle cyathiform,0.6
Although the lack of materialfrom interveningGuy- X 0.5 mm, slightly constrictedbelow tepals,glabrous
ana may be a collecting artifact,similar disjunctions inside. Tepals membranaceous,ovate, 1 X 0.6 mm
between Suriname and Venezuela are found in the (the inner whorl slightly broader), spreading to re-
rangeof widespreadE. bracteolataandE. canescens. curved at anthesis, the inner surface glabrous, the
As both have been found in Guyanaonly aroundthe marginsminutelypapillose. Stamensof whorls I and
upper Cuyuni-MazaruniRegion that borders Vene- II narrowly stipitate,0.6 mm tall, the antherstrans-
zuela, perhapshere too is where E. melinonii occurs versely oblong, 0.3 X 0.5 mm, glabrous, the apex
in Guyana.Nevertheless,if real, the absence of these truncateto emarginate,the connectives level with or
three species from relatively well-collected central reducedbetween the 2 locelli, these suborbicular,in-
Guyanais enigmatic. trorse,the filamentslaminar,much narrowerthanan-
thers, glabrous; whorl III stamens broadly stipitate,
0.6 mm tall, the anthersdepressed-elliptic,0.3 X 0.5
27. Endlicheria reflectens (Nees) Mez, Jahrb.Kon- mm, slightly bowed towardsthe outer whorls, locelli
igl. Gart. Berlin 5: 126. 1889. Goeppertiareflec- 2, extrorse-latrorse,the filaments slightly narrower
tens Nees, Linnaea21: 514. 1848. Type. Guyana. than anthers, laminar;opaque pilose near base, the
Withoutlocality and date (fl d), Schomburgk475/ basal glands sessile, globose, relatively large, filling
801 (holotype:B-n.v.; isotypes: E, K-n.v.). the spacebetweenfilamentsof innerandouterwhorls;
whorl IV wanting;pistillode wanting.Pistillateinflo-
Shrubsto 4 m. Branchletsslender,midway along rescence with indument, color, and branchingas in
flush 2-3 mm diam., distally weakly angular,soon staminateplants,the flowerssimilarin size andshape;
terete,densely rustytomentose,the surfaceconcealed stamens sterile, smaller;ovary glabrous,ovoid; style
by the indument cover or barely visible, the hairs stout, weakly distinguished from ovary; stigma
moderatelylong, to 0.5 mm, straightto crooked,erect, broadlytri-lobed,0.5 mm diam. Fruitsborneon slen-
loosely matted; terminal buds slender, 3 X 1 mm, der teretepedicels of up to 1 X 0.3 cm; cupules hem-
densely pubescent,the hairs as on branchlets,ascend- ispherical,to 1 X 1.5 cm, glabrousinside andoutside,
ing. Leaves altermate,widely and evenly spacedalong the marginsentire;drupesellipsoid, to 2 X 1 cm.
currentflush or loosely spiraledat tips of branchlets;
petioles slender,to 1 X 0.2 cm, terete, the indument Distribution (Fig. 20) and ecology. Shrubsfrom
as on branchlets;laminae chartaceousto coriaceous, savannaformationsof the Guiana shield at ca. 100-
plane, obovate to elliptic, 5-12 X 2-5 cm, the base 500 m. Flowering from March to November, and
acute,briefly decurrent,the apex obtuseto acute,acu- fruits availablefrom Januaryto June.
minate for up to 1 cm, the marginsminutelyrecurved
Representative specimens examined. COLOMBIA.
throughout;upper surface dull green to olive-brown,
VICHADA: Puerto Ayacucho, laja outcropsbehind Casuar-
waxy, the primaryto fourth-orderveins raised, their
ito, 4 Apr 1984 (fr), Gentry & Stein 46329 (MO, NY).
prominence decreasing with rank; lower surface VENEZUELA. BOLfVAR: Igneous outcrops at Agua
densely pilose, the hairs translucent,0.5 mm long, Amena, 100 m, 6 Sep 1985 (fl 6), Steyermarket al. 131425
erect, straightor curved near tips, uniformlydistrib- (MO,NY).
uted, all vein ordersraised,theirprominencedecreas- GUYANA. ESSEQUIBO: Rupununi Region, Rupununi
ing with rank;secondaryveins 4-6 per side, ? evenly R., Karanambo,1 Sep 1988 (fl 6), Maas et al. 7136 (K,
spaced, slightly more distant aroundmidlamina,as- MO,NY,U); Dadanawa,120-150m, 25 Jun1989(fl 6),
cending at 50-60o (more obtuse aroundmidlamina), Gillespie et al. 1691 (MO, NY, U, US).
arcuate,distal pairs loop-connected;tertiariesreticu- BRAZIL. AMAZONAS: Rio Branco,Aug 1913 (fl 6),
Kuhlmann3376 (U). RORAIMA: Rio Branco, Surumu,Jul
lating between secondaries.Staminateinflorescences
1909(fl 6'), Ule 8125 (G, K); SEMA Ecological Reserve,
distally clusteredin the axils of cataphylls,to 12 cm Ilha de Maraca, 11 Jul 1952 (fl d), Milliken et al. 439 (E,
long with 8 lateral branches,branchorders 2-3, the MO, NY).
highest order irregularto botryoid, lax, the flowers
distant, the axes sparsely pubescent, the hairs light Endlicheriareflectensalone in the Ampelodaphne
brown, 0.5 mm long, soft, erect, straightto crooked; species group has reticulatetertiaries.Here too basal
bractsandbracteolespersistentat anthesis,lanceolate, glands of whorl III stamensextendbetween filaments
densely pubescent,the hairs as on axes, appressedto of whorls I and II to an extent unmatchedin this spe-
ascending;pedicels terete, to 2 mm long, those sup- cies group. Elsewhere in the genus, similarly large
SYSTEMATICTREATMENT 65

glands arefound in E. acuminataandE. gracilis, both side; receptacle globose, 0.8 X 0.6 mm, constricted
with triplinervedleaves. Its obovate to elliptic leaves below tepals, glabrous inside. Tepals chartaceous,
with pilose lower surfaces and rusty tomentose ovate, 1 X 0.8 mm, spreadingto recurvedat anthesis,
branchletsand inflorescence are reminiscentonly of the inner surface minutely papillose towards apex,
E. multiflora.Ranges of the two species overlap in otherwiseglabrous,the marginspapillose. Stamensof
SW Guyana where, as elsewhere, E. multifloraare whorls I and II stipitate,relatively tall, up to 1 mm,
single-stemmedtrees of riparianforests andE. reflec- the antherstransverselyoblong, 0.3 X 0.5 mm, gla-
tens are shrubs of savanna formations with several brous, the apex truncateto emarginate,the connec-
slender stems that sprawl across adjacentvegetation. tives level with or reducedbetween the 2 locelli, these
The habit and habitatof E. reflectensis sharedwith suborbicular,introrse,the filamentslaminar,narrower
E. arenosa, but nonreticulate tertiaries immersed than anthers, glabrous; whorl III stamens stipitate,
above and short stiffly erect hairsbelow leaves of the equal to outer whorls, the anthersdepressed-oblong,
latterprovide easy vegetative contrast. erect, locelli 2, extrorse-latrorse,the filaments nar-
rowerthananthers,laminar,glabrous,the basalglands
broadly stipitate,globose, apiculate;whorl IV stami-
28. Endlicheria bracteata Mez, Repert.Spec. Nov. nodial or absent, columnar;pistillode fusiform. Pis-
Regni Veg. 16: 306. 1920. Type. Peru.San Martin: tillate inflorescencewith indumentandcolor as in sta-
Moyabamba, without date (fl d), Weberbauer minate plants, but shorter and with fewer lateral
4680 (holotype: B-n.v.; fragment,F; photo [neg. branches,the flowers slightly deeper;stamenssterile,
3813 ex B], F, G, GH, NY). smaller;ovary glabrous,ovoid; style slender,distinct
from ovary; stigma broadly tri-lobed, 0.5 mm diam.
Treesto 10 m. Branchletsslenderto stout,midway Fruits borue on terete pedicels of up to 7 X 3 mm;
along flush 3-7 mm diam., terete,densely hirsute,the cupules hemispherical,to 1 X 2 cm, initially rustyor
surface barely visible, the hairs very long, to 2 mm, grey-velutinousoutside, the indumentthinning with
straight,erect, grey to rusty red; terminalbuds slen- age, glabrousinside, the marginsentire;drupesellip-
der, 4 X 2 mm, densely pubescent, the hairs as on soid, to 2 X 1.5 cm.
branchlets,ascending. Leaves closely spiraledat tips
of currentflush;petioles short,robust,to 0.5-1 X 0.4 Distribution (Fig. 20) and ecology. Small un-
cm, semi-terete,the indumentas on branchlets;lam- derstorytrees of nonfloodedlowlandAmazoniato ca.
inae chartaceous,plane, obovate (often spathulate), 1200 m on the adjacenteastem Andeanslopes. Flow-
15-40 X 6-15 cm, the base acute or abruptlycon- ering materialcollected from JanuarythroughOcto-
tracted into the petiole, the apex obtuse, acuminate ber, fruits in March,July, August, and December.
for up to 3 cm, the margins minutely recurved
throughout;uppersurface darkgreen, the primaryto Representative specimens examined. ECUADOR.
fourth-orderveins raised, their prominencedecreas- NAPO:Tena,Estaci6nBiol6gica JatunSacha,400 m, 26 Jun
Palacios 7500 (MO).
ing with rank, or primary and secondary veins 1991 (fl 3),
PERU. HuANUCO:Leoncio Prado, Rupa Rupa, Tingo
sunken;lower surfacemoderatelypubescent,the hairs
Marfa,672 m, 21 Oct 1971 (fr), Schunke5060 (F, G, GH,
as on branchlets,rigidly erect, greyish green, denser MO, US); Pachitea,Sira Mtns., 800 m, 12 Aug 1988 (fl d
on main veins, all vein ordersraised,theirprominence juv), Morawetz & Wallnofer 11-12888 (MO). LORETO:
decreasingwith rank;secondaryveins 13-20 per side, Maynas, Iquitos, PuertoAlmendras,122 m, 16 Jun 1988 (fl
graduallyfurtheraparttowards apex, patent, diverg- i), Vasquez & Soto 12350 (GH, HBG, MO); Pumayacu,
ing at 70-85? (more obtuse towardsapex), arcuateto between Balsapuertoand Moyobamba,600-1200 m, Aug-
archingafter midcourse,distal pairs loop-connected; Sep 1933 (fl d), Klug 3187 (F, G, GH, K, MO, NY, US).
tertiaries oblique to midrib, between secondaries MADRE DE DIos: Tambopata,Zone Reservade Tambopata,
once-forkedto straight.Staminateinflorescencesdis- Rio Tambopata,260 m, 23 Aug 1990 (fl d), Alexiades &
tally clusteredin the axils of cataphylls,to 20 cm long Nicole 1042 (K, MO), 280 m, 19 Aug 1990 (fl d ), Reynel
al. 5244 (MO). SAN MARTiN: Jepelacio, near Moyob-
with 10 lateralbranches,branchorders3-4, the high- et
amba, 1100 m, Jul 1934 (fl d), Klug 3745 (F, GH, MO, NY,
est orderbotryoidto irregular,lax, the flowersdistant,
US); MariscalCaceres,TocacheNuevo, 400 m, 29 Jun 1978
the axes densely greyish hirsute, the hairs 1.5 mm (fl Y), Schunke 10304 (F, G, HBG, MO, NY). UCAYALI:
long, straight,erect; bracts and bracteolespersistent Coronel Portillo, Pucallpa, Estaci6n ExperimentalAlexan-
at anthesis, lanceolate, the indumentas on axes, as- der von Humboldt, Arboreto "A. Solagar Cavero,"23 Jul
cending; pedicels terete, to 2 mm long, those sup- 1980 (fl d), Sousa 76 (INPA).
porting secondary flowers slightly shorter.Flowers BRAZIL. ACRE:Tarauaca,Vit6riaVelho, 28 Sep 1994
campanulate,3 mm diam., sparsely grey-villose out- (fl d), Silveira et al. 943 (MO). AMAZONAS:Manaus,Re-
66
serva Florestal Ducke, Manaus-ItacoatiaraRd., km 26, 10
Aug 1965 (fl d), Rodrigues7028(INPA, NY), 4 Dec 1997
(fr), Souza& AssunVdo491 (MO). PARA:Oriximina,Rio
Trombetas,Porto Trombetas,29 Aug 1980 (fl 6), CidFer-
reiraet al. 1884 (HBG, INPA, K, MO, NY). ROND6NIA:
Santa Barbara,Rod. BR 364, km 120, 29 May 1982 (fl 5),
Teixeiraet al. 852 (F, MO, NY, US).
Local names. Peru: moena, shisho moena, shisho
moenita. Brazil: louro.
Endlicheria bracteata is conspicuous amongst
species of the Ampelodaphne species group with
prominenttertiaryand fourth-orderveins above (i.e.,
the E. bracteata subgroup)for its dense indumentum
of stiff greyish 2 mm long hairs on its branchlets,
lower leaf surfaces,andinflorescenceaxes. A globose
receptacle with strongly constricted apex below
spreadingtepals and definitely stipitatebasal glands
associatedwith whorl III stamensare also distinctive.
Leaf bases usually tapergraduallyinto the petiole, but
occasional subcordateforms (e.g., Klug 3187) resem-
ble those of typical of E. cocuirey, a species with a
much shorterindumentumof ca. 0.6 mm long hairs.
The variationof leaf shape in E. bracteata is paral-
leled by E. verticillata. The two coincide in central
Amazonia where, accordingto label data,E. verticil-
lata is found in flooded and E. bracteata in well-
drainedhabitats.Further,the hairs tend to be reddish
ratherthan greyish in E. verticillata, distal inflores-
cence axes and externalflowerpartsare glabrous,re-
ceptacles are campanulatewithoutconstrictedapices,
and basal glands are definitely sessile.
As young cupules can be densely pubescent out-
side (e.g., Schunke7245 & 8069), fruitingspecimens
of Endlicheria bracteata can be confused with E.
chalisea, a vegetativelysimilarspecies wheredensely
pubescent cupules are typical. However, in E. brac-
teata these hairs soon fall and maturecupules are ei-
ther glabrous or only sparsely pubescent. Further-
more, cupules here are always glabrousinside, while
in E. chalisea, cupules remaindensely pubescentin-
side even if the hairs outside are sometimes lost.
29. Endlicheria verticillata Mez, Bull. Herb.Boiss.
5 (ser. 2): 235. 1905. Type.Brazil. Amazonas:Rio
Jurua,Jun 1901 (fl d), Ule 5584 (holotype: B-
n.v.; isotypes: G, HBG, K).
Trees, 3-20 m. Branchletsslender to stout, mid-
way along flush 4-8 mm diam., terete, densely rusty
hirsute,the surfacebarely visible, the hairslong, to 2
mm, straight,erect;terminalbuds slender,4 X 2 mm,
densely pubescent,the hairsas on branchlets,ascend-
ing. Leaves closely spiraled at tips of currentflush;
FLORANEOTROPICA
petioles relatively short, robust, to 0.5-1 X 0.4 cm,
semi-terete, the indumentas on branchlets;laminae
chartaceous,plane, obovate, 15-35 X 5-12 cm, the
base acute or abruptlycontractedinto the petiole, the
apex obtuse, acuminatefor up to 3 cm, the margins
minutely recurved throughout;upper surface olive-
brown,the primaryto fourth-orderveins raised,their
prominencedecreasingwith rank,or midriband sec-
ondaries impressed,the higher-ordervenationprom-
inent; lower surface moderatelypubescent,the hairs
as on branchlets, rigidly erect, reddish, denser on
main veins, all vein orders raised, their prominence
decreasingwith rank;secondaryveins 11-15 perside,
graduallyfurtheraparttowards apex, patent, diverg-
ing at 70-85? (more obtuse towardsapex), arcuateto
archingafter midcourse,distal pairs loop-connected;
tertiaries oblique to midrib, between secondaries
once-forkedto straight.Staminateinflorescencesdis-
tally clusteredin the axils of cataphylls,to 25 cm long
with 12 lateralbranches,branchorders3-4, the high-
est orderbotryoidto irregular,lax, the flowersdistant,
the axes glabrousexcept for the sparselyrustyhirsute
peduncle;bractsand bracteolespersistentat anthesis,
lanceolate, sparsely rusty hirsute, glabrous after
second-orderbranches;pedicels terete,to 2 mm long,
those supportingsecondary flowers slightly shorter.
Flowers campanulate,3 mm diam., glabrousoutside;
receptaclecyathiform,0.8 X 0.6 mm, glabrousinside.
Tepals chartaceous,ovate, 1 X 0.8 mm, spreadingat
anthesis,the inner surfaceglabrous,the marginspap-
illose. Stamens of whorls I and II stipitate,0.6 mm
tall, the antherstransverselyoblong, 0.4 X 0.5 mm,
glabrous, the apex truncateto emarginate,the con-
nectives level with or reducedbetween the 2 locelli,
these suborbicular,introrse, the filaments laminar,
narrower than anthers, sparsely grey-tomentose;
whorl III stamensstipitate,equal to outerwhorls, the
anthers depressed-oblong,erect, locelli 2, extrorse-
latrorse,the filamentsnarrowerthananthers,laminar,
sparselygrey-tomentose,the basal glands sessile, glo-
bose; whorl IV wanting;pistillode fusiform.Pistillate
inflorescencewith indumentand color as in staminate
plants,but shorterandwith fewerlateralbranches,the
flowers slightly deeper; stamens sterile, smaller;
ovary glabrous, ovoid; style slender, distinct from
ovary; stigma broadly tri-lobed, 0.6 mm diam., the
lobes minutely papillose. Fruitsborne on terete ped-
icels of up to 6 X 4 mm; cupules shallowly hemi-
sphericalto patelliform,to 0.3 X 1 cm, glabrousin-
side and outside, the marginsentire;drupesellipsoid,
to 3.5 X 1.5 cm.
Distribution (Fig. 21) and ecology. Small to
medium-sized trees of flooded lowland forests
throughoutcentralAmazonia at ca. 90-200 m. Flow-
 SYST'EMATICTREATMENT
~~~~~~~~~~~~~~~~~~~~~~~r.~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~J
. ..... .......
FIG. 21. Distributionof Endlicheria verticillata,E. cocuirey, and E. tessmannii
ering March to October, fruits available from August
to the following June.
Representative specimens examined. COLOMBIA.
AMAZONAS:Leticia, Tarapaca, Parque Nacional Natural
Amacayacu, 100 m, 21 Jun 1991 (fl 6), Rudas et al. 2290
(MO), 125 m, 12 Jun 1992 (fr), Rueda 445 (MO).
PERU. LORETO:Maynas, Rio Amazonas, Yanamono
Explorama Tourist Camp, halfway between Indiana and
mouth of Rio Napo, 120 m, 15 Jun 1986 (fl d), Gentry et
al. 54550 (F, MO, NY); Sapuena,ArboretoJenaroHerrera,
140 m, 10 Aug 1988 (fr), van der Werifet aL 10053 (F,
HBG, MO, NY). UCAYALI: Pucallpa,Pau-Cocha,200 m, 6
May 1961 (fl c), Woytkowski6304 (F, GH, K, MO, NY).
BRAZIL. AcRE: Sena Madureira,trailto Rio laco from
kn 7 along Sena Madureirato Rio Branco rd., 1 Oct 1968
(fr), Prance et al. 7744 (HBG, INPA, MG, MO, NY, R);
Traumaturgo,Rio Alto Jurua,Reserva Extractivistado Alto
Jurua,4 Apr 1993 (fr), Silveiraet al. 468 (NY). AMAZONAS:
Manaus, Igap6 no Paranado Careiro, 21 Oct 1947 (fl d),
Duckes.n. R 19966 (HBG, MO, U); Livramento,TresCasas,
bacia do Madeira, 19 Sep 1962 (fr), Duarte 7368 (HBG,
INPA,MO); Paranado Autaz Mirim, 18 Apr 1966 (fr), Rod-
rigues & Mello 7760 (INPA, NY); Rio Negro, 23 Sep 1975
(fl ; ma), Kubitzki75-34 (B, HBG, INPA). PARk: Rio Ja-
6
pure, 14 Oct 1904 (fl 6), Ducke 6764 (G); Rio Tapaj6s,
Missao Cururu,17 Jul 1959 (fl 6), Elger 914 (HBG). RoND-
6NA: Rio Abuna, 18 Jul 1968 (fl S), Pranceet al. 6182
(HBG, INPA, MO, NY, R).
Local names. Peru: moena, muena amarilla,pu-
cacuro caspi (ant tree). Brazil: louro abacate, louro
cedro, lourojambo.
Endlicheria verticillata shares glabrous flowers
with E. arachnocome and E. arunciflora,but on ac-
count of its long hirsute vestiture and subsessile
leaves with prominenttertiaryand fourth-orderveins
above is best comparedto E. bracteata,a species with
densely pubescent inflorescences and ratherdifferent
flowers (see above).
30. Endlicheria cocuirey Kosterm., Recueil Trav.
Bot. N6erl. 34: 522. 1937. Type. Peru. Loreto:
Florida, Rio Putumayoat mouth of Rio Zubineta,
180 m, May-Jul 1931 (fl d), Kiug2253 (holotype:
F; isotypes: A, B-n.v., G, GH, K, MO, NY, S, US).
68
Trees, 4-15 m. Branchlets stout, midway along
flush 4-6 mm diam., terete, densely rusty to grey-
tomentose, the surfacebarely visible to concealed by
the indumentcover, the hairs relatively short, to 0.6
mm, straightto crooked, erect; terminalbuds plump,
4 X 3 mm, densely pubescent, the hairs as on bran-
chlets, ascending. Leaves closely spiraled at tips of
current flush; petioles robust, to 8 X 4 mm, semi-
terete, the indumentas on branchlets;laminae char-
taceous, plane, obovate, 15-30 X 6-11 cm, the base
roundedto cordate,the apex obtuse, acuminatefor up
to 1 cm, the margins minutely recurvedthroughout;
upper surface dark green to olive-brown, waxy, the
primaryto fourth-orderveins raised,theirprominence
decreasing with rank; lower surface sparsely grey-
tomentose, the hairs as on branchlets,scattered,erect
to ascending, slightly denser on main veins, all vein
ordersraised, theirprominencedecreasingwith rank;
secondary veins 11-16 per side, ? evenly spaced,
slightly more distant around midlamina, patent, di-
verging at 70-85? (more obtuse aroundmidlamina),
arcuateto arching after midcourse,distal pairs loop-
connected;tertiariesoblique to midrib,between sec-
ondaries once-forkedto straight.Staminateinflores-
cences distally clusteredin the axils of cataphylls,to
20 cm long with 15 lateralbranches,branchorders3-
4, the highest orderbotryoid to irregular,the flowers
clustered, the axes densely rusty to grey-tomentose;
bractsand bracteolespersistentat anthesis,ovate, the
indumentas on branchlets;pedicels terete, to 1 mm
long, reduced to lacking below secondary flowers.
Flowers campanulate 2 mm diam., sparsely grey-
tomentose outside, the hairs thinningdistally, lost by
tepals;receptaclecyathiform,0.6 X 0.5 mm, glabrous
inside. Tepals membranaceous,elliptic, 0.8 X 0.6
mm, spreadingto recurvedat anthesis, the inner sur-
face glabrous, the margins minutely papillose near
apex. Stamens of whorls I and II stipitate, 0.6 mm
tall, the anthersdepressed-ovate,0.3 X 0.4 mm, gla-
brous, the apex truncateto emarginate,the connec-
tives level with or reducedbetween the 2 locelli, these
suborbicular,introrse,the filamentslaminar,narrower
than anthers,sparsely grey-tomentoseoutside; whorl
III stamens stipitate,0.6 mm tall, the anthersoblong,
0.3 X 0.2 mm, erect, locelli 2, extrorse-latrorse,the
filaments narrower than anthers, laminar, densely
grey-tomentoseinside, the basal glands sessile, glo-
bose; whorl IV staminodial,columnar;pistillode fil-
iform. Pistillate plants unknown.
Distribution (Fig. 21) and ecology. Small trees
of nonfloodedforest in westernAmazoniaat ca. 100-
200 m. Floweringspecimenscollected from Marchto
June. Fruitsunknown.
FLORANEOTROPICA
Additional specimens examined. PERU. LORETO:
Requena, Sapuena,ArboretoJenaroHerrera,125 m, 2 Mar
1989 (fl juv), Freitas 53 (MO).
BRAZIL. AMAZONAS: Rio Jutahy,Riosinho Juruema,
2 Jun 1945 (fl d), de Lemos Fr6es 21022 (F, NY).
Local names. Peru: hioma cocuir-ey (Huitoto),
puspo moena.
Endlicheria cocuirey is a poorly known species
that shares subsessile leaves with contractedor sub-
cordatebases and adaxiallyprominenttertiarieswith
E. bracteata and E. verticillata. The latter has gla-
brous flowers, and E. cocuirey is distinguishedfrom
E. bracteata by its short tomentose ratherthan stiff
hirsute indument. Similar indument and leaves, but
with immersed tertiariesabove, are otherwise found
only in E. dictifarinosa.
31. Endlicheria tessmannii 0. C. Schmidt,Notizbl.
Bot. Gart. Berlin-Dahlem 10: 227. 1928. Type.
Peru. Loreto: Iquitos, without date (fl d), Tess-
mann 5146 (holotype:B-n.v.; isotype: NY).
Trees to 22 m. Branchlets stout, midway along
flush 4-8 mm diam., angular,densely pubescent,the
surface concealed by a dense tomentellose cover of
grey to yellowish hairs (0.1 mm or less) interspersed
with straight to crookedly sprawling rusty hairs of
0.7-1 mm; terminalbuds plump, 5 X 4 mm, densely
pubescent, the hairs as branchlets,ascending to ap-
pressed. Leaves closely spiraled at tips of current
flush;petioles robust,to 7 X 0.5 cm, semi-terete,the
indument as on branchlets;laminae chartaceousto
membranaceous,plane, obovate to elliptic, 20-45 X
8-15 cm, the base acute, the apex obtuse, acuminate
for up to 4 cm, the margins minutely recurved
throughout;uppersurfacedull greyish green to olive-
brown,the primaryto fourth-orderveins raised,their
prominence decreasing with rank; lower surface
sparselypubescent,the hairs appressedto ascending,
0.1 mm long, denseron main veins, interspersedwith
sprawlingearly caducoushairsof 0.7 mm on the mid-
rib, all vein ordersraised, their prominencedecreas-
ing with rank; secondary veins 10-15 per side, ?
evenly spaced, slightly more distantaroundmidlam-
ina, patent,divergingat 70-850 (more obtuse around
midlamina),arcuateto archingaftermidcourse,distal
pairs loop-connected;tertiariesobliqueto midrib,be-
tween secondaries forked. Staminateinflorescences
distally clusteredin the axils of cataphylls,to 30 cm
long with 15 lateralbranches,branchorders3-4, the
highest orderbotryoidto irregular,the flowersloosely
crowded,the axes densely pubescent,the indumentas
SYSTEMATICTREATMENT 69

on branchlets;bracts and bracteolespersistentat an- al. 651 (F,G, MO,NY);Varadera de Mazan,27 Sep 1972
thesis, ovate, densely pubescent, the hairs appressed (fl d ), Croat20771(F,G, MO).
to ascending, silvery grey; pedicels of terminalflow- Local names. Peru:muena, moena amarilla.
ers terete,to 0.5 mm long, lacking below penultimate
flowers.Flowers campanulate,2.5 mm diam.,densely Endlicheriatessmanniihas uniqueindumentin the
pubescentoutside,the hairsyellowish to grey,0.3 mm Ampelodaphnespecies group. Vegetativesurfacesof
long, straight,ascending, sparsertowardstepals; re- no otherspecies thereinshow the extremelyshort(ca.
ceptacle cyathiform,0.6 X 0.6 mm, glabrousinside. 0.1 mm) hairsfound in E. tessmannii.Moreover,only
Tepals membranaceous, elliptic, 1.2 x 0.7 mm, here are hairs on the laminae and veins below ap-
spreading to recurved at anthesis, the inner surface pressed or weakly ascending. In all other species of
glabrous,the marginsand apex inside papillose. Sta- the Ampelodaphnespecies group, the hairs are erect
mens of whorls I and II stipitate, 0.6 mm tall, the on the lower leaf surface. Further,the flowers of E.
anthers transversely oblong, 0.25 X 0.3 mm, gla- tessmannii bear an indument of short (ca. 0.2 mm)
brous, the apex truncateto emarginate,the connec- straighthairs on the outer surfacethat is quite unlike
tives level with or reducedbetweenthe 2 locelli, these the villose indumentumtypicallyfoundin this species
suborbicular,introrse,the filamentslaminar,narrower group.These characterscircumscribea narrowlydis-
than anthers,the base sparsely grey-pilose;whorl III tributedspecies that is morphologicallyclose to the
stamens stipitate,ca. 0.6 mm tall, the anthersoblong, widespreadand highly variableE. directonervia.
0.3 X 0.2 mm, erect, locelli 2, extrorse-latrorse,the
filaments narrowerthan anthers, laminar, the base
sparselygrey-pilose,the basal glandssessile, globose; 32. Endlicheria directonervia C. K. Allen, Mem.
whorl IV wanting;pistillode filiform. Pistillate inflo- New YorkBot. Gard. 10(5): 60. 1964. Type.Ven-
rescence with indument and color as in staminate ezuela. Amazonas: Cerro Sipapo, 125 m, 17 Jan
plants,but shorterand with fewer lateralbranches,the 1949 (fl d), Maguire & Politi 28428 (holotype:
flowers slightly deeper; stamens sterile, smaller; NY; isotypes: GH, US).
ovary glabrous, ovoid; style slender, distinct from
Treesto 30 m. Branchletsslenderto stout,midway
ovary; stigma tri-lobed, 0.4 mm diam. Fruits borne
along flush 4-8 mm diam., terete, densely greyish or
on stout terete pedicels of up to 1 X 0.4 cm; cupules
rusty tomentose, the surface concealed by the indu-
hemispherical,to 1 x 2 cm, sparsely brown tomen- ment cover, the hairs relatively long, to 0.7 mm,
tellose outside, the indument thinning with age, straight, erect, sometimes interspersedwith longer
densely yellowish strigose inside, the marginsentire; crookedly erect hairs of 1 mm; terminalbuds plump,
drupesellipsoid, to 2 X 1.5 cm. 6 X 4 mm, densely pubescent,the hairsas on branch-
lets, erect to ascending.Leaves closely spiraledat tips
Distribution (Fig. 21) and ecology. Trees from
of currentflush;petioles robust,to 7 X 0.5 cm, semi-
nonfloodedforests aroundIquitos,Peru,at ca. 100 m.
terete, the indumentas on branchlets;laminae char-
Flowering Septemberto November,fruits Januaryto
taceous to membranaceous,plane, obovateto elliptic,
July.
15-45 X 6-18 cm, the base acute, the apex obtuse,
Representativespecimensexamined.PERU. LOR- acuminatefor up to 4 cm, the marginsminutely re-
ETO:Maynas,Iquitos,carretera km21.5,tro- curved throughout; upper surface green to olive-
Iquitos-Nauta,
cha de penetraci6nhasta el caserfo de Yarana,120 m, 1 Feb brown, sparsely tomentose, the primary to fourth-
1995 (fr), Rimachi11305 (MO); 7 km SW of Iquitos,31 Jul order veins raised, their prominencedecreasingwith
1972 (fr), Croat 18609 (MO); Estaci6n ExperimentalIIAP, rank; lower surface moderatelypubescent, the hairs
Allpahuayo, 106 m, 1 Jun 1990 (fr), Vdsquez& Jaramillo of mixed length, 0.3-1 mm, erect to ascending,
13945 (MO); Las Amazonas, ExplornapoCamp, 100-140 straightto crooked,longerhairsdenser,persisting,all
m, Feb 1991 (fr), Pipoly etal. 13554 (MO);Mishuyacu,near vein ordersraised, their prominencedecreasingwith
Iquitos, 100 m, Oct-Nov 1929 (fl d), Klug 161 (F, NY, US);
rank; secondary veins 12-18 per side, ? evenly
Punchana,trochade la Comunidadde San Antonio, 120 m,
spaced, slightly more distantaroundmidlamina,pat-
30 Jun 1998 (fr), Rimachi12263 (MO);Rio Amazonas,trail
from Caserfo Santa Maria de Ojeal, 30 Sep 1977 (fl d),
ent, diverging at 70-85o (in lower lamina often per-
Rimachi 3226 (MO); YanamonoExploramaTourist Camp, pendicular to midrib,more acutely towardsapex), ar-
halfway between Indianaand mouth of Rio Napo, 130 m, cuate, or arching after midcourse, distal pairs
23 Jan 1983 (fr), Gentry et al. 39734 (F, G, MO); Rio Mo- loop-connected;tertiariesoblique to midrib,between
m6n, 4 Sep 1972 (fl ; ma), Croat 19982 (MO); Estaci6n secondariesforked. Staminateinflorescencesdistally
Biologica Callicebus, 130, 26 Oct 1980 (fl d), Vasquezet clusteredin the axils of cataphylls,to 20 cm long with
70 FLORANEOTROPICA

10 lateral branches, branch orders 3-4, the highest ECUADOR.MORONA-SANTIAGO: Pumpuentza, 250
orderbotryoid to irregular,or condensed,the flowers m, 29 Jun 1980 (fr), Brandebyge & Asanza 32418 (MO);
clustered,the axes rustyto grey-tomentose;bractsand Tukupi,250 m, Jun 1980 (fr), Brandebyge& Asanza 32271
(MO). NAPO:Aguarico, Reserva Ethnica Huaorani,carre-
bracteoles persistent at anthesis, ovate, densely sil-
tera y oleoducto de Maxus, km 119-120, 235 m, Mar 1995
very grey-villose;pedicels terete,to 2 mm long, those
(fr), Aulestia et al. 3593 (MO);YasuniForest Reserve, E of
supportingsecondary flowers slightly shorter.Flow- Pontificia UniversidadCat6lica del EcuadorScientific Sta-
ers campanulate, 3 mm diam., sparsely grey or tion, 225 m, 20 Jun 1995 (fr), Acevedo & Cedenlo7416
densely yellowish pubescent outside, the hairs 0.7 (MO); Archidona,carreteraHollin-Loreto, Rfo Huataraco,
mm long, crooked;receptaclecyathiform,0.6 x 0.6 800-1000 m, 23-30 Aug 1989 (fr), Cer6n & Factos 7649
mm, glabrous inside. Tepals membranaceous,ovate, (MO);Loreto,faldasdel VolcanSumaco,690 m, 2 Mar 1996
1.2 X 0.7 mm, spreadingto recurvedat anthesis,the (fl Y, fr juv), Vargas & Grefa 764 (MO); Tena, Estaci6n
inner surface glabrous, the margins and apex inside Biol6gica JatunSacha,450 m, 12 Nov 1987 (fr), Cer6n2679
papillose. Stamensof whorls I and II stipitate,0.6 mm (HBG, MO). PASTAZA: PuertoSarayacu,3 Oct 1974 (fl 6),
Lugo 3906 (GB); Pozo petroleroVillan6n 2 de ARCO, 400
tall, the antherstransverselyoblong, 0.25 X 0.3 mm,
m, 1-18 Dec 1991 (fl 6), Hurtado2979 (MO). SANTIAGO-
glabrous, the apex truncateto emarginate,the con-
ZAMORA: Taisha, 1500 ft, 7 Feb 1962 (fr), Cazalet & Pen-
nectives level with or reducedbetween the 2 locelli, nington 7716 (NY). SucuMBios: Cascales, ParroquiaEl
these suborbicular,introrse, the filaments laminar, Dorado, 250 m, 5 May 1997 (fr), Freire et al. 2226 (MO).
narrowerthan anthers,the base sparsely grey-pilose; PERU. AMAZONAS: Bagua, Imaza, Comunidadde Ya-
whorlIII stamensstipitate,ca. 0.6 mm tall, the anthers mayakat,500 m, 17 Oct 1996 (fl d), Vdsquez& Jaramillo
oblong, 0.3 X 0.2 mm, erect, locelli 2, extrorse- 20300 (MO); Rio Cenepa, vic. of Huampami,200-250 m,
latrorse,the filamentsnarrowerthananthers,laminar, 8 Aug 1978 (fr), Ancuash 1343 (F, G, HBG). HUANUCO:
the base sparselygrey-pilose,the basal glands sessile, Pachitea,Pucallpa, 1080 m, 22 Dec 1987 (fl 6), Wallnofer
globose; whorl IV wanting;pistillode filiform.Pistil- 14-221287 (MO). LORETO:Requena, Sapuena, Arboreto
JenaroHerrera, 170 m, 15 Sep 1987 (fl juv), Vdsquez &
late inflorescencewith indumentand color as in sta-
Jaramillo 9606 (F, MO, NY), 3 Nov 1980 (fl Y), Castillo
minate plants, but shorter and with fewer lateral
65 (F). PAsco: Oxapampa,Iscozacfn, 16 Jun 1986 (fl 6),
branches,the flowers slightly deeper;stamenssterile, Pariona & Sebastian25 (F, MO, NY); Palcazu,300-600 m,
smaller;ovary glabrous,ovoid; style slender,distinct 4 Jun 1984 (fl 6), Hartshornet al. 2600 (MO, NY). PUNO:
from ovary; stigma tri-lobed, 0.4 mm diam. Fruits Carabaya,cabecerasdel Rfo Candamo,800-850 m, 14 Nov
borne on teretepedicels of up to 1 X 0.3 cm; cupules 1996 (fr), Cornejo & Balarezo 2701 (MO).
hemispherical,to 1 X 2 cm, sparsely brown tomen- BRAZIL. ACRE:MancioLima, MancioLimaRamaldo
tose outside, the indumentthinningwith age, densely Banho, 8 Nov 1991 (fr), Cid Ferreira et al. 10642 (MO):
yellowish strigose inside, the margins entire;drupes AMAZONAS: Humaita,on plateaubetween Rio Livramento
ellipsoid, to 2 X 1.5 cm. and Rio Ipixuna,Cipoal, 7-18 Nov 1934 (fr), Krukoff7206
(A, NY); EstradaManaus-Caracarai,km 130, 13 Nov 1973
Distribution (Fig. 22) and ecology. Small to large (frjuv), Berg et al. P19527 (HBG, INPA,MO, NY); Reserva
trees of both flooded and nonfloodedforests in Ama- FlorestalDucke, Manaus-ItacoatiaraRd., km 26, Bosque de
zonia and adjacent lower montane eastern Andean PalmeirasP-1068, Jul 1963 (fl 6), Rodrigues5389 (F, HBG,
slopes, to ca. 1500 m. FloweringJunethroughfollow- INPA, NY); PresidenteFigueiredo,Rio Uatuma,UHE Bal-
bina, 15 Aug 1986 (fl 6), Freitas et al. 239 (INPA); Rio
ing January,fruits availablethroughoutthe year.
Ituxi,Labreaairstrip,28 Jun 1971 (fl 9 ), Prance et al. 13888
Representativespecimensexamined.COLOMBIA. (HBG, INPA, MO, NY, US); Rio Negro, right side of Ilha
AMAZONAS: Leticia,nearEl Marco,200 m, 7 Sep 1963(fl Tamandua(locally Ilha Maraj6), 19 Oct 1987 (fr), Maas et
d), Soejarto 580 (GH, K, US); Rio Apaporis,Cachiverade al. 6798 (K, MO, NY, U, US); Sao Gabriel de Cachoeira,
Jirijirimoy alrededores, 250 m, 12 Jun1951(fr),Schultes Camanaus,19 Oct 1978 (fl 6), Nascimento685 (HBG, MG,
& Cabrera 12423 (GH, NY, U, US); Soratama,250 m, 21 NY).
Jun 1951 (fr), Schultes & Cabrera 12735 (GH, NY). CA-
QUETA: CordilleraOriental,vertienteoriental,bosquesentre Local names. Colombia: ha-ro, las brisas, laurel.
Sucre y La Portada,1200-1350 m, 5 Apr 1940 (fl d), Cua- Venezuela: laurel hediondo. Ecuador: ocatoe (Huaor-
trecasas 9123 (COL, US); Florencia, carreteraFlorencia- ani). Peru: muena, palta moena, roble anis amarillo,
Suaza, km 28, 1000-1400 m, 10 Nov 1993 (fl d), Ramirez tinci, tikis, uchitinchi, und yuwich (Huambisa), wau
et al. 4886 (MO). META: Sierra de la Macarena, 1500 m,
yuwich (Huambisa). Brazil: louro, louro cururti.
30 Dec 1949 (frjuv), Philipson & Idrobo2013 (COL, NY).
VENEZUELA. AMAZONAS:Yavita,Rio Guiania,Ma-
roa, 127 m, 14 Feb 1942 (fr), Williams14370 (A, G, F); Rio As circumscribed here Endlicheria directonervia
Negro, Cano 12 km NE of San Carlos, 120 m, 15 Apr 1979 is a highly variable species. Type material is conspic-
(fr), Liesner 6657 (MO). uous for rusty tomentose indumentum and secondary
SYSTEMATICTREATMENT
< -e;
; ~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~.
..
...-v....o;.
AMr.
...4 ~~~~~~~~~~~~
A"'k ~ ~ ~ ~ ~ ~ ~ ~ ~
FIG. 22. Distibution
FIG. 22. Distributionof Endlicheriadirectonervia.
veins directed ? perpendicularto midrib, but only
Rodrigues 5389, Berg et al. P19527 (both near Ma-
naus, Brazil), Soejarto et al. 580 (Leticia, Colombia),
and Wallnofer 14-221287 (Huanuco, Peru) match
these characters.All other specimenshere assignedto
E. directonerviadepartfrom typical materialby one
or more characters.
In Williams14370, from near the type locality in
Venezuela,and severalcollections fromEcuador(e.g.,
Lugo 3925 and Hurtado 2979), the perpendicularly
directedsecondariesof the type combine with sparse
sprawling greyish hairs. Provided that these can be
assigned to E. directonervia,specimens with similar
indument(e.g., Ceron & Factos 7649) or with a mix-
ture of tomentose and longer sprawlinggreyish hairs
(e.g., Vdsquez& Jaramillo 20300), but without per-
pendicularlydirectedsecondaries,become acceptable
as well. Similarly, another variant combines dense
greyish tomentose indumentwith (e.g., Freitas et al.
239 and Ram(rezet aL 4886) and without (e.g., Cua-
trecasas 9123) laterallydirectedsecondaries.The re-
71
~ ~ ~ A~-.-
of Endlicheria directonervia.~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
.. .. .. ..
~~ ~.,.~~~~~,
~
Wk AVjA*~R~
maining specimens, Acevedo & Cede*io7416, Aules-
tia et al. 3593 (Napo, Ecuador),Schultes & Cabrera
12423 & 12735 (Amazonas,Colombia),and Vdsquez
& Jaramillo 9606 and Vdsquezet aL 12391 (Loreto,
Peru) have non-perpendicularsecondaries and rela-
tively short erect hairs of ca. 0.3 mm on the leaves
below interspersed, on the midrib, with sprawling
0.5-0.7 mm hairs. These specimens approachE. tess-
mannii, but their hairs are still an orderof magnitude
longer (0.1 mm) and erect ratherthan appressed.
The variationin E. directonerviacan be arranged
so that specimens departfrom each otherby only one
character,either indumentor venation. Furthermore,
no geographicor altitutudinalcorrelationcan be dis-
cerned. Therefore, separation into distinct species
seems unwarranted.
33. Endlicheria chalisea Chanderbali, Novon 6:
329. 1996. Type. Guyana. Essequibo: Potaro-
SiparuniRegion, Kato and vicinity, dry forests N
72 FLORANEOTROPICA

of town, 750 m, 19 Mar 1989 (fl d), Hahn et al. smaller; ovary glabrous, ellipsoid, 0.5 X 0.3 mm;
5795 (holotype:MO; isotypes: F, NY, U, US). style slender, distinct from ovary; stigma tri-lobed,
0.3 mm diam. Fruits bome on short claviform pedi-
Trees to 25 m. Branchlets stout, midway along cels of up to 0.7 X 0.5 cm;
cupules hemispherical,to
flush 5-8 mm diam., terete, densely hirsute,the sur- 2 X 2 cm, densely velutinous outside, the indument
face concealed by the indumentcover, the hairslong, thinning with age, sericeous inside, the marginsen-
to 1.2 mm, straight,rigidly erect, yellowish; terminal tire; drupesellipsoid, to 4 X
1.5 cm.
budsplump,2 X 1.5 mm, densely pubescent,the hairs
as on branchlets,ascending. Leaves loosely spiraled Distribution (Fig. 23) and ecology. Medium-
at tips of currentflush;petioles robust,to 4 X 0.5 cm, sized trees of lowland Amazonia and upland forests
semi-terete, the indumentas on branchlets;laminae of the Guiana highlands and eastermAndean slopes
chartaceous to coriaceous, plane to subbullate, to ca. 2500 m. Flowering specimens collected from
broadly elliptic to obovate, 15-30 X 7-16 cm, the DecemberthroughAugust, fruitsavailablefromFeb-
base acute, the apex obtuse, acuminatefor up to 2.5 ruarythroughNovember.
cm, the marginsminutelyrecurvedthroughout;upper
surface green to olive-brown, waxy, the midrib and Representative specimens examined. VENEZUELA.
AMAZONAS: Atabapo, Alto Rfo Casiquiare,160 m, 6 Mar
secondariesimmersedto sunken,the higher-orderve-
1990 (fr), Aymard& Delgado 8483 (MO, NY); Rfo Mata-
nationraised;lower surfacedensely yellowish hirsute,
cuni, 1 Feb 1990 (fr), Stergios & Velasco 14495 (HBG,
the hairserect,up to 0.8-1.2 mm long, denseron main INPA, NY, US).
veins, all vein orders raised, their prominence de- GUYANA. EsSEQUIBO: Potaro-Siparuni, Iwokrama
creasingwith rank;secondaryveins 6-14 per side, + RainforestReserve, 13-27 Jul 1996 (fl d), Chanderbaliet
evenly spaced, slightly more distantaroundmidlam- al. 159 (MO); N Pakaraimas,Ciong Valley, Manawarrai
ina, patent,divergingat 70-85o (moreacutelytowards Mtn., 2000 m, 1 Jun 1995 (fr), Mutchnick1464 (MO); Mt.
apex), arching after midcourse, distal pairs loop- Ayangana,1140 m, 16 Aug 1960 (fr), Tillettet al. 45163 (K,
connected;tertiariesoblique to midrib,between sec- MO, NY, US); Maburaregion, Kurupukarimain, km 6, 23
ondaries once-forkedto straight.Staminateinflores- Mar 1994 (fr), Ek & Hammond1033 (MO); UpperTakatu-
UpperEssequibo,Rupununi,rd. from Lethemto 25 km past
cences distally clusteredin the axils of cataphyllsor
SuramaVillage entrance,90-110 m, 28 Feb 1990 (fr), Ac-
foliage leaves, to 7 cm long with 8 lateralbranches, evedo 3432 (MO, NY, US).
branch orders 2-3, the highest order condensed, the FRENCH GUIANA. Riviere Grand Inini-Bassin du
flowers clustered,the axes densely yellow to reddish Maroni, 11 Jul 1990 (st), Sabatier & Prevost 3195 (MO).
hirsute; bracts and bracteoles persistent at anthesis, PERU. LORETO: Requena, Sapuena, Arboreto Jenaro
ovate, the indumentas on axes; pedicels terete,short, Herrera,8 Jul 1986 (fl d), Valcdrcel& Chota 1/98 (MO), 9
to 1 mm long, reduced to lacking below secondary Dec 1980 (fl d), Castillo 87 (F). UCAYALI:CoronelPortillo,
flowers. Flowers campanulate,up to 3 mm diam., CordilleraAzul, 1200 m, 4 Jun 1981 (fr), Young& Sullivan
densely pubescent, the hairs yellowish, straight to 663 (MO).
BRAZIL. AMAZONAS: Manaus, Reserva Florestal
crooked, sparser towards tepals; receptacle cyathi-
Ducke, Manaus-ItacoatiaraRd., km 26, 27 Jun 1964 (fl d),
form, 1 X 1 mm, slightly constricted below tepals,
Rodrigues & Loureiro5926 (F, HBG, NY); PresidenteFi-
glabrous inside. Tepals membranaceous,elliptic, 1.3 gueiredo, Rio Uatuma, UHE Balbina, 15 Jul 1990 (fl d),
X 1 mm, the tips recurvedat anthesis,otherwiseerect, Cid Ferreira et al. 7582 (K, MO, NY). PARA: Oriximina,
the inner surface glabrous,the marginsand apex in- Rio Trombetas,terrafirme adjacentto lago Moura,25 Aug
side lightly papillose. Stamensof whorls I and II 0.5 1980 (fl d), Cid Ferreiraet al. 1832 (HBG, INPA, MO).
mm tall, stipitate, the anthers transversely oblong, BOLIVIA.LA PAZ: Prov. Sud Yungas,7 km de Huan-
0.25 X 0.5 mm, glabrous,the apex truncateto emar- cane en carreteraa San Isidro,2300 m, 13 Dec 1989 (fl 9),
ginate, the connectives level with or reducedbetween Smith et al. 13908 (MO); ibid., Huancane, 7.5 km hacia el
the 2 locelli, these suborbicular,introrse,the filaments sud sombreel camino nuevo, 2410 m, 9 Mar 1980 (fr), Beck
laminar,much narrowerthananthers,glabrous;whorl 3192 (MO).
III stamens stipitate, 0.6 mm tall, the anthers de- Local names. Venezuela: laurel carutillo, laurel
pressed-oblong, 0.3 X 0.4 mm, erect, locelli 2, ex- babosa. Guyana: yellow silverballi (locally usually
trorse-latrorse,the filaments narrowerthan anthers, applied to Aniba spp.), wild pear. Peru: moena de hoja
laminar,glabrous, the basal glands sessile, globose; grande. Brazil: louro de folha larga, louro imbauba.
whorl IV staminodial, stipitate, the apex elliptic-
ovate;pistillode fusiform.Pistillateinflorescencewith Endlicheria chalisea is immediately recognized in
indument,color, andbranchingas in staminateplants, fruit by its densely pubescent cupules. However, it
the flowers sessile, slightly deeper; stamens sterile, resembles E. glomerata in the long-hirsute yellowish
SYSTEMATICTREATMENT
>
. j > e - z ^ ;
i//A
F%IG.23. Distributionof Endlicheriachalisea and E. glomerata.
indumentumand (sub-)sessile flowers in dense clus-
ters. The two species are sometimes vegetatively in-
distinguishablebut the densely pubescent cupules of
E. chalisea have neverbeen found in the SE Brazilian
rangeof E. glomerata.There,cupules, whetheryoung
or mature,are glabrous.
34. Endlicheria glomerata Mez, Jahrb.Konigl. Bot.
Gart. Berlin 5: 127. 1889. Type. Brazil. Rio de
Janeiro:Maua, 15 November 1874 (fl 9), Glaziou
7781 (lectotype, designatedby Kostermans,1937:
P; isolectotypes: B-n.v., K).
Trees to 8 m. Branchletsslenderto stout, midway
along flush 4-8 mm diam., terete, densely pubescent,
the surfaceconcealed by the indumentcover, the hairs
yellowish, 0.4-1 mm long, straightto crooked, erect;
terminalbuds plump, 1 X 0.5 cm, densely pubescent,
the hairs as on branchlets,ascending.Leaves spiraled
at tips of currentflush; petioles robust, to 3.5 X 0.4
cm, semi-terete,the indumentas on branchlets;lam-
73
~* V .'Wr
inae chartaceous,bullate(rarelyplane), obovateto el-
liptic, 17-32 X 4-13 cm, the base obtuse, the apex
obtuse to acute, acuminatefor up to 2 cm, the margins
minutely recurved throughout; upper surface light
green to olive-brown,the midriband secondariescon-
vex between bullae (prominulous when laminae
plane), the higher-ordervenationraised;lower surface
moderatelyto densely pubescent,the hairsas on bran-
chlets, slightly denser on main veins, all vein orders
raised, their prominence decreasing with rank; sec-
ondary veins 10-14 per side, ? evenly spaced,
slightly more distant around midlamina, patent, di-
verging at 70-850 (more obtuse aroundmidlamina),
arching after midcourse, distal pairs loop-connected;
tertiaries oblique to midrib, between secondaries
straight to forked. Staminate inflorescences distally
clusteredin the axils of cataphylls,to 20 cm long with
12 lateral branches, branch orders 3-4, the highest
order condensed, the flowers clustered, the axes
densely yellowish to rusty pubescent, the hairs as on
branchlets;bracts and bracteoles persistentat anthe-
74 FLORA NEOTROPICA

sis, ovate, the indumentas on axes; pedicels terete,to US), 3 Oct 1930 (fl d), Mexia 5138 (F, G, GH, MO, NY, P,
0.5 mm long, reduced to lacking below secondary S, U, US). RiODE JANEIRO:Governador Portela,Monte
flowers. Flowers campanulate,2.5 mm diam., grey- Sinai, 1935 (fl 9, fr), MachadoNunes 318 (HBG, MO, U);
Maud,Nov 1890 (fl d), Schwackes.n. R 30973 (R), 26 Jun
villose to glabrousoutside;receptaclecyathiform,0.5
1901 (fr) Hemmendorif 457 (R, S); Morrode Viraqa6,Praia
x 0.6 mm, constrictedbelow tepals, silvery sericeous
Grande,3 Apr 1891 (fl 9, fr), Glaziou 18451 (F, L, K, NY,
inside. Tepals chartaceous,broadly ovate, 1 X 0.8 P); Capelinhade St. Antonio, 22 Aug 1894 (st), Glazious.n.
mm, erect to spreadingat anthesis, the inner surface (E, NY); Rio das Flores, FazendaSantaGenoveva,500-600
glabrous,the marginspapillose. Stamens of whorls I m, 7 Oct 1971 (fl 5), Sucre 7779 (HBG, MO); s.loc., 13
and II stipitate, 0.6 mm tall, the anthersdepressed- May 1878 (fr), Glaziou 8093 (K, P).
ovate, 0.3 x 0.4 mm, glabrous,the apex truncateto
Local names. Brazil:canela, canelao.
emarginate,the connectives level with or reducedbe-
tween the 2 locelli, these suborbicular,introrse,the Endlicheriaglomerataalone of theAmpelodaphne
filaments laminar,much narrowerthan anthers,gla- species group inhabitsthe Atlantic coastal forests of
brous; whorl III stamens stipitate, equal to outer SE Brazil. In most flowering specimens, a long hir-
whorls, the anthersdepressed-ovate,0.3 X 0.4 mm, sute vestiture of stiffly erect yellowish 1 mm hairs
erect, locelli 2, extrorse-latrorse,the filaments nar- combines with bullate laminae and densely clustered
rowerthananthers,laminar,glabrous,the basalglands subsessile pubescent flowers. This charactercombi-
sessile, globose; whorl IV wanting; pistillode fusi- nation is unique to E. glomerata but appearsto be
form. Pistillateinflorescencewith indumentandcolor unstable. Laminae range from strongly bullate in
as in staminateplants, but shorterand with fewer lat- Mexia 5138, throughmoderatelyso in the type ma-
eral branches, the flowers slightly deeper; stamens terial, to plane in Glaziou 18451 and Oliveira et al.
sterile, smaller;ovary glabrous,ovoid; style slender, 770 withoutaccompanyingvegetativeor floraldiffer-
distinct from ovary; stigma tri-lobed, 0.3 mm diam. ences.
Fruits borne on short terete pedicels of up to 5 X 3 Two other indumentvariantsshow parallelinsta-
mm, often sessile; cupules hemispherical,to 0.5 X 1 bility in bullaeformation.The specimensfromParque
cm, glabrous inside and outside, the margins entire; Estadualdo Rio Doce (Costa s.n. BHCB32676, Her-
drupesellipsoid, to 2.5 X 1.5 cm. inger 18530, 18580, and 16026, Heringer & Eiten
15083 and 15154, Lombardi 1987, and Martinelli et
Distribution (Fig. 23) and ecology. Small trees
al. 23), have a much shorterindumentumof 0.4 mm
from the lower montaneAtlanticcoastal forestsof SE
hairs, while in Hatschbachet al. 57930, Hatschbach
Brazil at 200-700 m. Flowering specimens collected
& Silva 52169, and Sucre 7799 vegetative indument
in May, July, September, October, and November.
is as in typical material but distal inflorescence
Fruitscollected in April, June,July,August, and No-
branchesand flowers are glabrous.Attachingspecific
vember.
importanceto the single characterthat distingishes
Additionalspecimensexamined.BRAZIL.GoIAs: both variants from typical material would result in
Niquelandia, FazendaEngenho,580 m, 27 Jun 1997 (fr), threevery similar,and sympatric,species thatcannot
Oliveira et al. 770 (MO).MINAsGERAIS:Caratinga,Esta- be confused with others in the genus, but probably
cvo Biol8gicade Caratinga, 6 Aug 1984 (fr), Strier992 with each other-in my opinion an unnecessaryin-
(MO,NY);Caratinga, FazendaMaced6nea/Cenibra, Ipaba flation of species diversity.
Trilhodo Triangulo (T2-37),4 Nov 1991(fr),Bragaet al.
s.n. BHCB27250(MO);Ilheu,Fazendada Tabunha, Capi-
chava, 220 m, 31 Aug 1930 (fr), Mexia 5025 (G, GH, NY,
US); Leopoldina, Rod. BR-116, Rio Pombos, 14 Jul 1988
35. Endlicheria anomala (Nees) Mez, Jahrb.Kon-
(fl 5), Hatschbach& Silva 52169 (HBG, US), 10 Oct 1992 igl. Bot. Gart.Berlin 5: 133. 1889. Goeppertiaan-
(fl 5), Hatschbachet al. 57930 (HBG, MO, NY, S); Mar- omala Nees, Syst. laur. 370. 1836. Type. Brazil.
lieria, ParqueEstadualdo Rio Doce, 400 m, 18 Sep 1975 (fl Amazonas:Rio Amazonas,nearEga, Sep 1831 (fl
d), Heringer & Eiten 15083 (HBG, US), 22 Sep 1975 (fl 5), Poeppig 2552 (lectotype, designatedby Kos-
5), Heringer & Eiten 15154 (MO, US), 23 Mar 1976 (fl Y), termans, 1937: W-n.v.; isolectotypes: B-n.v.,
Heringer 16026 (HBG, K, U), 200 m, 30 Aug 1973 (fr), BM-n.v., G-n.v., KIEL-n.v., LE-n.v., LZ-n.v.,
Martinelliet al. 23 (MO), 29 Jun 1993 (fr), Costa s.n. BHCB NY, OXF-n.v., P-n.v.).
32676 (MO), 13 Sep 1997 (fl 5), Lombardi 1987 (MO);
Tim6teo,ParqueEstadualdo RioDoce,20 Oct 1982(fl d), Goeppertia polyantha Meisn. DC. Prodr. 15(1): 175.
Heringer 18530 (HBG), 22 Oct 1982 (fl d), Heringer18580 1864. Syntypes. Brazil. Amazonas: Rio Negro,
(HBG); Vi,oca, oxcart rd. to Sao Miguel, ca. km 4, 685 m, Barra, Jul 1851 (fl 5), Spruce 1648 (B-n.v., BM-
23 Sep 1930 (fr), Mexia 5091 (F, G, GH, MO, NY, S, U, n.v., G-n.v., K-n.v., LE-n.v., NY-n.v., OXF-n.v., P-
SYSTEMATICTREATMENT
n.v., US-n.v.); ibid., Apr 1851 (fl d juv), Spruce
1433 = 'Nectandra(6)' (B-n.v., BM-n.v., E, G-n.v.,
GH-n.v., K-n.v., LE-n.v., M, NY-n.v., OXF-n.v.,
P-n.v.). Guyana. Without locality and date (fl Y),
Schomburgk784 (BM-n.v., B-n.v., F-n.v., G-n.v.,
K, MO, OXF-n.v., P, U, US, W-n.v.).
Ocotea simulans C. K. Allen, Mem. New York Bot.
Gard. 10(5): 99. 1964. Type. Venezuela.Amazonas:
Rio Casiquiare,just above Capihuara,100-130 m,
24 Jun 1959 (fl 6), Wurdack& Adderly43169A (ho-
lotype: NY; isotype: US).
Trees to 15 m or shrubs.Branchletsslender,mid-
way along flush 2-3 mm diam., distally weakly an-
gular,soon terete,densely pubescent,the surfacecon-
cealed by the indument cover to barely visible, the
hairs short,to 0.5 mm, straight,appressed,ascending,
or erect, silvery-grey to rust-brown;terminal buds
slender, 3 X 1 mm, densely pubescent, the hairs as
on branchlets. Leaves alternate,widely and evenly
spaced along currentflush; petioles slender, to 1 X
0.2 cm, semi-terete, the indument as on branchlets;
laminae chartaceous,plane, ovate, 8-17 X 2-4 cm,
the base obtuseto rounded,brieflydecurrent,the apex
acute, acuminatefor up to 2 cm, the marginsminutely
recurvedthroughout;uppersurfacedull greyishgreen
to dark olive-brown, waxy, the midrib sunken, sec-
ondary and higher-ordervenationraised, their prom-
inence decreasing with rank; lower surface densely
sericeous or tomentose, the hairs appressed,ascend-
ing, or erect, uniformly distributed,all vein orders
raised, their prominence decreasing with rank; sec-
ondaryveins 4-6 per side, + evenly spaced, slightly
more distantaroundmidlamina,ascendingat 50-60?
(more obtuse aroundmidlamina),arcuate,distalpairs
loop-connected; tertiaries roughly horizontal to
oblique to midrib, between secondaries straight to
forked. Staminateinflorescencesevenly spaced along
currentflush in the axils of foliage leaves, to 10 cm
long with 8 lateral branches,branchorders 3-4, the
highest order dichasial, lax, the flowers distant, the
axes sparsely rusty to grey-strigillose; bracts and
bracteoles caducous by anthesis, lanceolate, the in-
dument as on axes, denser; pedicels terete, to 3 mm
long, those supporting secondary flowers slightly
shorter.Flowers rotate, 5 mm diam., sparsely rusty
strigillose outside; receptacle patelliform, 0.5 X 1
mm, densely rusty tomentose inside. Tepals charta-
ceous, obovate,2 X 1 mm, spreading,the androecium
exserted at anthesis, the outer surface sparsely rusty
to grey-strigillose,the inner surfacesparselyrusty to
grey-strigillose near base, otherwise minutely papil-
lose throughout,the margins papillose. Stamens of
whorlsI and II stipitate,0.7 mm tall, the antherstrans-
versely oblong to obovate, 0.3 X 0.4 mm, glabrous,
75
the apex truncate,the connectiveslevel with or broad
above the 2 locelli, these suborbicular,the filaments
laminar, narrower than anthers, slender, glabrous;
whorl III stamens stipitate,ca. 1 mm tall, the anthers
oblong, 0.5 X 0.3 mm, bowed towards the outer
whorls, locelli 4, upper pair introrse-latrorse,lower
pair extrorse-latrorse,the filamentsnarrowerthanan-
thers, laminar,glabrous,the basal glands sessile, glo-
bose; whorl IV wanting;pistillode fusiform.Pistillate
inflorescencewith indumentand color as in staminate
plants,but shorterandwith fewer lateralbranches,the
flowers similar in size and shape; stamens sterile,
smaller; ovary glabrous;style slender, distinct from
ovary; stigma tri-lobed, 0.4 mm diam. Fruits borne
on claviform pedicels of up to 1 X 0.5 cm; cupules
scarcely broader,patelliform,to 0.6 cm diam., stri-
gose inside, the marginslobed, tepal bases persisting;
drupesellipsoid, to 1.2 X 0.6 cm.
Distribution (Fig. 24) and ecology. Small to
medium-sized trees or shrubs frequentin inundated
lowland forests (ca. 50-300 m) throughoutthe Am-
azon basin. Flowering and fruiting throughoutthe
year.
Representative specimens examined. COLOMBIA.
AMAZONAS:Puerto Narifio, Rio Loretoyacu, Trapecio
Amaz6nico, 100 m, Jun 1973 (fl d), Soejarto et al. 4198
(NY); Rio Yari,QuebradaEl Mochilero, 120-200 m, 21 Apr
1986 (fr), Galeano et al. 1024 (MO). META:PuertoL6pez,
240 m, 27 Aug 1944 (fl Y), Little & Little 8259 (COL).
VAUPES:Rio Vaupes,Mitd, 9 Jul 1976 (fl d), Zarucchiet
al. 1830 (COL, MO); 2 mi upriverfrom Mitui,1 Nov 1976
(fl 6), Davis 216 (COL, K). VICHADA:ParqueNacional
Natural"El Tuparro,"La Linea Roja, just S of Rio Tomo,
100 m, 12 Mar 1985 (fr), Zarucchiand Barbosa3679 (MO).
VENEZUELA.AMAZONAS: Atabapo, Rio Asisa, 100
m, Oct 1989 (fl Y), Delgado 868 (MO);Atures,PuertoAya-
cucho, 37 m, 23 Jul 1981 (fl 6), Castillo 1282 (MO, NY).
ApuRE:PedroCamejo,Lagunala Guacharaca,70 m, 24 Feb
1979 (fl d), Davidse & Gonzales 15701 (MO); ParqueNa-
cional "SantosLuzardo,"5 Apr (fr), G6mezet al. 649 (MO).
BOLiVAR: Cedefno,Rio Parguaza,30 m, 25-28 Jan 1989 (fr),
Cuello 637 (MO, NY); Rio Pargueni,1-10 km above mouth,
90 m, 10 Dec 1955 (fl 6, most diseased), Wurdack& Mon-
achino 39775 (NY); SierraIchdn,cercaniasdel Salto Maria
Espuma(Salto Ichdn)del Rio Ichdn,base de la sierraIchdn,
tributarydel Rio Paragua,500 m, 28 Dec 1961 (fr), Stey-
ermark90315 (NY).
ECUADOR. NAPO:Aguarico,ParqueNacionalYasuni,
200 m, 22 Sep 1988 (fl Y, fr juv), Ceron & Gallo 4898
(MO, NY); Rio LagartaCocha, 6 Aug 1992 (fl 6), Delprete
et al. 6141 (MO). SUCUMBiOS: Lago Agrio, ReservaFaun-
istica Cuyabeno,230 m, 4 Oct 1991 (fl d), Palacios et al.
8144 (MO).
PERU. LORETO: Loreto, Nauta, Reserva Nacional
Pacaya-Samiria,Rio Yanayacu,campamentoPalizada,90 m,
76
FIG. 24. Distributionof Endlicheriaanomala.
3 Nov 1992 (fl 6), Del Carpio & Ruiz 1620 (MO); Maynas,
Iquitos, 100 m, 3-11 Aug 1929 (fl c), Killip & Smith27192
(NY); Requena, Avispa Cocha, Rfo Tapiche, 180 m, 9 Jan
1984 (fl 6), Vdsquezet aL 4778 (MO). UCAYALI: Coronel
Portillo, Yarinacocha, Cafnode Alijandriam vecindad de
Lago Yarinacocha,210 m, 22 Nov 1997 (fl Y, fr), Graham
& Schunke365 (MO).
BRAZIL. ACRE:Cruzeiro do Sul, Rio Jurua,Igarape
Preto, 29 Oct 1966 (fl juv), Prance et aL 2972 (HBG, NY,
R). AMAZONAS: Barcelos, Rio Araca, 29 Jul 1985 (fl 6),
Cordeiro309 (INPA, MO); Humaita,Rio Madeira,lago do
Purusinho,15 May 1985 (fl 6), Hendersonetal. 458 (INPA,
K, MO, NY); Manaus, 28 Mar 1937 (fl Y, fr), Ducke 440
(K, MO, NY, R, U); Bom Jardim,near Flutuantedo INPA,
7 Apr 1971 (fl 6), Prance et al. 11737 (COL, INPA, MO,
NY, R, U); Rio Negro, Ilha da Costa Arirarra,28 June 1979
(fl 6), Poole 1717 (INPA,MO); Ilha das Onqas,24 Jun 1992
(fl 6), Mori & Gracie 22473 (INPA, MO, NY); Tefe, lago
de Tefe, 27 Feb 1972 (fl 6), Albuquerqueet al. 570 (INPA).
PARA:Oriximink, Rio Trombetas,9 Jul 1980 (fl J), Cid
Ferreira et al. 1388 (INPA, MO, NY); Rio Paru do Oeste,
Lago Ara,c, 8 Sep 1980 (fl 6), Cid Ferreira et al. 2321
(INPA, MO, NY). ROND6NIA:Rio Jaciparand,4 km above
FLORA NEOTROPICA
. | . . .|.f. ~~~~~~~~~~~
f~~~~~~~~~~~~~~~~.
T
.X
r|0iw
O
_^-S
', .. .. . .
7 4 - -
Jaciparana,28 Jun 1968 (fl d), Prance et al. 5295 (MO,
NY); Rio PacaasNovos, 8-25 km above mouth,6 Aug 1968
(fl d), Prance et al. 6834 (MO, NY).
BOLIVIA. BEM: Guayaramerin,Rio Mamor6, 15 Feb
1978 (fl 9, fr), Anderson 12052 (NY). LA PAZ:Prov. Itur-
ralde, Luisita, 180 m, 29 Feb 1984 (fl 9, fr), Beck & Haase
10129 (MO). PANDO:Manupiri,Rio Manupiri,278 m, Sep
1996 (fr), Paniagua & Foster 690 (MO).SANTA CRUZ:
Prov.Velasco, ReservaEcologfia El Refugio, 160 m, 24 Apr
1995 (fl d), GuillUn& Chore 3292 (MO); Campos de San
Ram6n, 8-10 Aug 1995 (fl d), Halloy et aL 4319 (MO).
Local names. Colombia: ko-ma-nee-nee-ko
(Makuna),chaviaco negro, pee-shee'. Venezuela:lau-
rel blanco, laurel blanco de orilla del rio, laurel de
revalta,laureldel rio, laurelfino, laurelrebalsero,lau-
rel oriyera, laurelito. Peru: canela, isma moena,
moena de bajo, muena blanca, sanango,yacu muena.
Brazil: louro canela, louro do igap6, louro flor, louro
tambaqui.
Endlicheriaanomala is the only species of the ge-
nus in which whorl HI stamens are consistently pro-
SYSTEMATICTREATMENT 77

vided with four-locellate anthers.The rotate flowers long with 14 lateralbranches,branchorders3-4, the
with an androeciumin which all stamenshave distinct highest order dichasial, lax, the flowers distant, the
filaments and whorl III stamens are closely grouped axes densely reddish to rusty tomentose; bracts and
with their anthersextrorselybowed, are also unmis- bracteoles caducous by anthesis, lanceolate, densely
takable.The claviformpedicels and shallow platelike rusty strigose; pedicels terete, to 2 mm long, those
cupules with persistenttepal bases resemblethose of supportingsecondary flowers slightly shorter.Flow-
E. lorastemonbut are much smallerin size. ers hypocrateriform,5 mm diam., sparselyrusty stri-
Most specimens, including the type materialand gose outside; receptacleinfundibuliform,1 X 1 mm,
the type of Ocotea simulans, bear an appressedin- densely grey-tomentose inside. Tepals chartaceous,
dument of silvery hairs, the denser forms of which narrowly elliptic or ligulate, 2 X 1 mm (the inner
are reminiscent of the Endlicheria sericea species whorl slightly broader),ascendingto spreadingat an-
group, but several specimens, including the type ma- thesis, the inner surface and margins sparsely rusty
terial of Goeppertiapolyantha, have a rusty tomen- papillose, otherwise glabrous. Stamens of whorls I
tose vestiture much like that typical of Endlicheria and II stipitate, 1 mm tall, the anthers transversely
paniculata. However,ascendinghairsof intermediate oblong, 0.3 X 0.4 mm (slightly narrowerin whorl II),
orientationalso appear(e.g., Cavalcante& Silva 1782 the apex truncate,the connectives broad above the 2
andMori & Gracie 22473), and flowersand fruitsare locelli, these suborbicular,introrse,the filamentslam-
uniformirrespectiveof indumentorientation. inar, narrowerthan anthers, sparsely whitish grey-
pilose, the hairs straight, erect; whorl III stamens
broadlystipitate, 1 mm tall, pressed together,the an-
36. Endlicheria williamsii 0. C. Schmidt, Repert. thers oblong, bowed towardsthe outerwhorls, locelli
Spec. Nov. Regni Veg. 31: 177. 1933. Type. Peru. 2, extrorse-latrorse,occasionally a second introrse-
Loreto: Maynas, Rio Nanay, Maquisapa, 5 Jul latrorsepair above, the filamentsalmost as broad as
1929 (fl d), Williams1193 (holotype:F; isotype: anthers,wider towardsbase, the indumentas in outer
NY). whorls, the basal glands sessile, globose; whorl IV
wanting; pistillode fusiform. Pistillate inflorescence
Trees to 20 m. Branchletsrelativelystout,midway
with indumentand color as in staminateplants, but
along flush 4-5 mm diam., distally weakly angular,
shorter and with fewer lateral branches,the flowers
soon terete,densely tomentose,the surfaceconcealed
slightly deeper; stamens sterile, smaller; ovary gla-
by the indumentcover, the hairs relatively short, to
brous, ovoid; style slender, distinct from ovary;
0.3 mm, straight to crisped, erect, rusty to reddish
stigma tri-lobed,0.4 mm diam. Fruitsbormeon short
brown;terminalbuds plump, 4 X 3 mm, densely pu-
claviform pedicels of up to 0.5 X 0.3 cm; cupules
bescent, the hairs as on branchlets,ascending.Leaves
hemispherical,to 1 X 1.5 cm, glabrousoutside,tawny
closely spiraled at tips of currentflush; petioles ro-
sericeous inside, the marginsentire;drupesovoid, to
bust, to 2.5 X 0.3 cm, terete, slightly flattenedabove,
2 x 1.2 cm.
the indumentas on branchlets;laminae chartaceous,
plane, broadly elliptic, 12-20 X 6-12 cm, the base Distribution (Fig. 25) and ecology. Medium-
obtuse, briefly decurrent,the apex obtuse, abruptly sized trees of seasonally flooded forest in western
acuminatefor up to 1.5 cm, the marginsminutelyre- Amazonian at ca. 100-200 m. Flowering specimens
curved throughout;upper surface greyish to olive- collected in June and July, fruits in January,March,
brown, minutely punctulate,the midrib and second- April, and September.
aries prominent, tertiary and higher-ordervenation
immersed; lower surface obscured by a short Representativespecimensexamined.COLOMBIA.
tomentose cover of 0.3 mm long creamish hairs in- AMAZONAS: Rio Yari,QuebradaEl Mochilero, 120-200 m,
terspersedwith straighterect reddishhairsof 0.7 mm, 21 Apr 1986 (fr), Galeano et al. 1065 (MO).
all vein orders raised, their prominence decreasing PERU. LORETO: Maynas, Rio Ampiyacu,between Esti-
r6n and TierraFirme, 24 Apr 1977 (fr), Plowmanet al. 7013
with rank; secondary veins 5-7 per side, ? evenly
(F, GH, MO, NY); Rio Nanay, Iquitos, Almendro, 17 Jun
spaced, slightly more distant around midlamina, as-
1976 (fl d), McDaniel & Rimachi 20744 (MO); Mishana,
cending at 50-60O (more acutely towards apex), the 120 m, 11 Jan 1976 (fr), Gentry et al. 15865 (F, G, MO,
lowermost pair sometimes asymmetric (one or both NY); QuebradaYarana,130 m, 10 Jul 1988 (fl d), Vdsquez
merging with margin at base), distal pairs loop- etal. 10939(MO);Timbuchi, Jun-Jul1929(fljuv),Williams
connected;tertiariesroughlyhorizontal,between sec- 1002 (F), (fl d), Williams1003 (F, G), (fl Y, fr), Williams
ondaries straightor forked. Staminateinflorescences 1004 (F, G); Rio Ampiaco, 24 Sep 1972 (fr juv), Croat
distally clustered in the axils of cataphylls,to 20 cm 20680 (HBG, MO, NY).
78 FLORA NEOTROPICA

..........~~~.t .

-7-

4~~~~~~~~~~~~~ $~~~~~~~~~~~~

FIG. 25. Distributionof Endlicheriawilliamsii E canescens, and E. xerampela.

Local names. Colombia:jigua. Peru:isma muena, Trees to 35 m. Branchletsslender, midway along

pampa muena. flush 2-3 mm diam., angular,densely tomentose, the
surface concealed by the indument cover, the hairs
Endlicheria williamsii is immediately recognized
relatively short, to 0.3 mm, straightto crisped, erect,
by the mixtureof shortcreamishtomentoseindument
that completely obscures the lower surfaceof the ter- reddish to rust brown; terminal buds plump, 3 X 2
minally clusteredleaves. Similar indumentcombines mm, densely rusty tomentose. Leaves alternate,
with widely spaced leaves in E. duotincta, and on widely and evenly spaced along currentflush;petioles
account of its phyllotaxy E. williamsii may be mis- slender,to 2.5 X 0.3 cm, semi-terete,the indumentas
taken for a member of the Ampelodaphne species on branchlets; laminae stiff chartaceous, plane,
group, but its flowers suggest affinity with E. anom- broadlyelliptic, 10-20 X 5-8 cm, the base acute, of-
ala. The receptacle is deeper in E. williamsii but the ten asymmetric, briefly decurrent, the apex acute,
androeciumis very similar, even occasionally show- acuminatefor up to 1.5 cm, the marginsminutely re-
ing the four-locellate whorl m antherstypical of E. curved throughout;upper surface reddish to olive-
anomala. (e.g., Vdsquezet al. 10939). brown, minutely punctulate,the midrib and second-
aries depressed,the higher-ordervenationimmersed;
lower surface densely tomentose, the hairs erect,
37. Endlicheria canescens Chanderbali,Novon 6: straight,tan to light brown, slightly denser on main
328. 1996. Type. Suriname.SaramaccaRiverToe- veins, all vein orders raised, their prominence de-
koemoetoe Creek, 5 Oct 1994 (fl di), Maguire creasing with rank;secondaryveins 4-7 per side, +
24898a (holotype: MO; isotypes: A, F, G, K, NY, evenly spaced, slightly more distant aroundmidlam-
U, US, W). ina, ascendingat 50-60o (more acutelytowardsapex),
SYSTEMATICTREATMENT
arcuate,the lowermostpair often asymmetric(one or
both merging with marginat base), distal pairs loop-
connected;tertiariesoblique to midrib,between sec-
ondaries once-forkedor straight.Staminateinflores-
cences evenly spaced along currentflush in the axils
of foliage leaves, to 15 cm long with 8 lateral
branches,branchorders2-3, the highest orderdicha-
sial or irregular,lax, the flowersdistant,the axes rusty
tomentose;bracts and bracteolescaducous by anthe-
sis, lanceolate, rusty tomentose; pedicels terete, to 3
mm long, those supportingsecondaryflowersslightly
shorter. Flowers broadly rotate, to 5 mm diam.,
sparsely rusty to grey-strigillose outside; receptacle
broadly infundibuliform,2 X 3 mm, slightly con-
strictedbelow tepals, densely grey-velutinousinside.
Tepals fleshy, obovate, 1.9 X 1.3 cm, spreading at
anthesis, the inner surface densely greyish white to-
mentose, the margins densely reddish brown papil-
lose. Stamensof whorls I and II broadlystipitate,rel-
atively short, 0.6 mm, the anthers depressed-ovate,
0.3 X 0.5 mm (slightly narrowerin whorl II), gla-
brous,the apex sharplyapiculate,the connectivespro-
longed between the 2 locelli, these obliquely hemi-
spherical, introrse-latrorse,the filaments clavate,
taperingtowardsbase, densely grey-tomentelloseout-
side; whorl III stamens sessile, 0.6 mm tall, the an-
thers depressed-oblong,0.3 X 0.6 mm, erect, locelli
2, extrorse-latrorse,the filamentsas broadas anthers,
columnar;densely grey-tomentelloseinside, the basal
glands sessile, globose, apiculate;whorl IV wanting;
pistillode filiform. Pistillate inflorescencewith indu-
ment and color as in staminateplants,but shorterand
with fewer lateral branches, the flowers slightly
deeper; stamens sterile, smaller; ovary glabrous, el-
lipsoid, 1 X 0.6 mm; style stout,weakly distinguished
from ovary; stigma minutelytri-lobed,0.3 mm diam.
Fruitsborne on claviformpedicels of up to 1.5 X 0.7
cm; cupules hemispherical,to 1 X 1.5 cm, glabrous
outside, rusty sericeous inside, the margins entire;
drupesellipsoid, to 1.3 X 0.9 cm.
Distribution (Fig. 25) and ecology. Largetreesof
terra firme forests from the Guianas, western Ama-
zonia, and easternAndean slopes at ca. 100-1700 m.
Flowers collected in May, June, September,October,
November, and January,and fruits in October,No-
vember, April, May, and June. If not a sampling ar-
tifact, it is conceivable the May-June flowering sea-
son results in October-November fruits, while the
September-Januaryflowering period provides fruits
from April to June.
Representativespecimensexamined.COLOMBIA. type anthers, of which E. ferruginosa has similar in-
dument,but in E. canescens whorl I and II anthersare
AMAZONAS: Leticia, Tarapaca,Parque Nacional Natural
79
Amacayacu, CabaniaPamate, Rfo Cotuhe, 100 m, 29 Jun
1991 (fl d), Rudas et al. 2642 (MO). PUTUMAYO: Mocoa,
San Antonio, Alto Campucaca,La Mariposa, 1350 m, 20
April-I May 1994 (fr), Ferndndezet al. 11101 (MO).
VENEZUELA. BOLiVAR: Between San Ignacio de Ya-
runi and San Francisco de Yaruni, 1200 m, 4 Jan 1975 (fl
Y), Steyermark111387 (NY).
GUYANA. ESSEQUIBO: Upper Mazaruni River Basin,
KamarangR., 24 Oct 1960 (fl d), Tillett& Tillett45789 (F,
K, NY, US).
SURINAME. Natuurpark Brownsberg, 16 Jun 1917 (fl
d ), SurinameForestBureau2936 (A, MO, U), 23 Jun 1925
(fl d), SurinameForestBureau6884 (MO, US).
ECUADOR. NAPO:Aguarico, Reserva Etnica Huaor-
ani, 247 m, 10-14 Nov 1993 (fl 6), Dik 767 (MO); Reserva
FaunisticaCuyabeno,Zancudo, 230 m, 5 Oct 1991 (fl Y),
Palacios et al. 8164 (MO); Orellana,ParqueNacional Ya-
sunf, 230 m, 6 Oct 1993 (fr), Dik 665 (QCNE-n.v., MO).
PASTAZA: Pastaza,Pozo petrolero"Danta2" de UNOCAL,
50 km SSE of Cararay,365 m, 1-20 Oct 1990 (fl 6), Es-
pinoza & Coba 378 (MO). ZAMORA-CHINCHIPE: Nangar-
itza, Pachicutza, Camino al Hito, Cordillera del C6ndor,
1200-1300 m, 20 Oct 1991 (fl Y), Palacios et al. 8399
(MO).
PERU. CAJAMARCA: San Ignacio,RicardoPalma, 1650
m, 20 May 1998 (fl 6), Campos & L6pez 4930 (MO); Ta-
baconas,La Bermeja,1600-1700 m, 19 Nov 1997 (fr), Cam-
pos & Cano 4695 (MO). LORETO: Alto Amazonas,Cerros
Campanquiz,Pongo de Manseriche,300-550 m, 19-21 Oct
1962 (fl 6), Wurdack2362 (F, GH, NY, UC, US).
BRAZIL. AMAZONAS: Benjamin Constant, s.d. (st),
Brees 40 (INPA, MO).
Local names. Colombia: amarillo. Brazil: louro
comum.
This species was first broughtto my attentionas
Endlicheria endlicheriopsis(Mez) Kosterm.,a com-
binationbased on Ocotea endlicheriopsisMez. How-
ever, Kostermanshad overlooked the four-locellate
anthersin the pistillatetype materialof the basionym,
and Endlicheriacanescens was described to accom-
modate the two-locellate plants formerly associated
with Ocotea endlicheriopsis(Chanderbali,1996).
From the several congeners with similar rusty to-
mentose branchletsEndlicheria canescens is easily
recognized by its rotate flowers where fleshy obovate
tepals have densely greyish white tomentose inner
surfaces.Immersedtertiariesabove and a tendencyto
have asymmetricleaf bases with a pairof secondaries
emerging from the junctureof leaf base and petiole
are also useful for identification.Flower shape and
subsessile whorl I and II stamens with rathersmall
locelli arereminiscentof species with Microlocellata-
80 FLORA NEOTROPICA

provided with sharply apiculate ratherthan truncate truncateto emarginate,the connectives level with or
apices. reducedbetween the 2 locelli, these suborbicular,in-
trorse, the filaments laminar,narrowerthan anthers,
sparselyrustypilose; whorl III stamenssessile, 0.6 m
tall, the anthers depressed-oblong,0.3 X 0.4 mm,
38. Endlicheria xerampela Chanderbali,sp. nov.
erect,locelli 2, extrorse-latrorse,the filamentsbroader
Type. Colombia. Choco: Halfway between Certe-
than anthers,ligulate, sparselyrustypilose nearbase,
gui and Las Animas, 100 m, 17 Aug 1976 (fl d),
the basal glands sessile, globose, apiculate;whorl IV
Gentry & Fallen 17803 (holotype:MO; isotypes:
wanting; pistillode minute, filiform. Pistillate inflo-
F, G, HBG, NY). Fig. 26
rescence with indument and color as in staminate
Endlicheriae canescentis aemulans, floribus infundibu- plants, but shorter and with fewer lateral branches.
laribuset antherissex externislateoblongisconnectivonec Mature pistillate flowers unknown. Fruits borue on
supraloculosproductodiffert. claviformpedicels of up to 7 X 4 mm; cupules hem-
ispherical,to 1 X 1.5 cm, glabrousoutside, rusty se-
Trees to 18 m. Branchletsslender,midway along riceous inside, the margins entire; drupes ovoid, to
flush 3-4 mm diam., angular,densely reddishbrown 1.5 X 1 cm.
tomentose, the surface concealed by the indument
cover, the hairs relatively short, to 0.3 mm, straight, Distribution (Fig. 25) and ecology. Medium-
erect; terminalbuds plump, 3 X 2 mm, densely pu- sized trees from the lowland (0-150 m) forests of the
bescent, the hairs as on branchlets,ascending.Leaves Pacific coast of Colombia.Flowering specimens col-
alternate, widely and evenly spaced along current lected in April, August, and October,fruitsin March
flush;petioles slender,to 1.5 X 0.15 cm, semi-terete, and April.
the indumentas on branchlets;laminae chartaceous,
Additional specimens examined. COLOMBIA.
plane, ovate, 8-15 X 3-7 cm, the base obtuse, the CAUCA:Bajo Calima, Concesi6n Pulapel/Buenaventura,
apex acute, acuminatefor up to 1 cm, the margins 100m, Oct 1987(fl 9, juv),Monsalve1975(MO);Lopez,
minutelyrecurvedthroughout;uppersurfacerusty to Rio Naya,"ElCarmen," 50-150 m, Apr1992(fr),Cogollo
yellowish green, sparsely rusty strigose, waxy, the et al. 5167 (MO).VALLE: Buenaventura, Concesi6ndePul-
midrib and secondaries immersed, the higher-order papel,0-50 m, Apr1992(st),Cogolloet al. 5190(MO),(fl
venation prominulous;lower surface sparsely rusty , juv),Cogolloet al. 5195(MO);Corregimento SanIsidro,
tomentose, the hairs as on branchlets, scattered on Quebrada Ordofiez,40 m, Mar1989 (fr),Devia & Prado
lamina, denser on main veins, all vein ordersraised, 2669 (MO);RioCajambre, "ElChorro," 0-50 m, Apr1992
(fr), Cogollo et al. 5182 (MO).
their prominence decreasing with rank; secondary
veins 4-6 per side, ? evenly spaced, slightly more Endlicheriaxerampela is most similar to E. ca-
distantaroundmidlamina,ascendingat 50-60o (more nescens. The two share a dense reddish tomentose
obtuse aroundmidlamina),arcuate,distal pairs loop- vestiture and deeply hemispherical cupules with a
connected;tertiariesroughlyhorizontal,betweensec- rusty sericeous indumentinside. However,the upper
ondaries straightto forked. Staminateinflorescences leaf surface of E. xerampela is smooth ratherthan
evenly spaced along currentflush in the axils of fo- minutely punctulate,the flowers are infundibuliform
liage leaves, to 7 cm long with 8 lateral branches, ratherthanrotate,and apices of whorl I and II anthers
branch orders 3-4, the highest order essentially di- are truncateratherthan apiculate.None of the other
chasial, but the internodes reduced and flowers ar- species of Endlicheriaknown from the Choco region
ranged in pseudo-umbellate fascicles, the axes are remotely similar.
sparsely rusty tomentose; bracts and bracteoles ca-
ducous by anthesis, ovate, the indumentas on axes;
pedicels terete, to 2 mm long, those supportingsec- 39. Endlicheria citriodora van derWerff,Ann. Mis-
ondary flowers slightly shorter.Flowers infundibuli- souri Bot. Gard. 78: 415. 1991. Type. Peru. Lor-
form, 2.5 mm diam., sparsely rusty strigose outside; eto: Requena, Sapuena,ArboretoJenaroHerrera,
receptacle broadly infundibuliform,0.5 X 1 mm, 140 m, 7 Aug 1988 (fl d), van der Werf et al.
densely rustypilose inside. Tepalschartaceous,ovate, 9991 (holotype:MO; isotypes: AMAZ-n.v., F, G,
I x 0.6 mm (the inner whorl slightly narrower),as- HBG-n.v., MO, US).
cending at anthesis, the inner surface sparsely grey-
tomentose,the marginsand tips papillose. Stamensof Trees to 25 m. Branchlets stout, midway along
whorls I and II stipitate, 0.4 mm tall, the anthers flush 4-6 mm diam., angular,striate, densely rusty
depressed-oblong,0.3 x 0.4 mm, glabrous,the apex tomentellose, the surface concealed by the indument
SYSTEMATICTREATMENT 81

FIG. 26. Endlicheria xeramipela(A-E, Gentry & Fallen 17803; F, Cogollo et aL 5167). A. Habit.B. Flower. C.
Flower l.s. D. Whorl III stamen seen from without. E. Whorl I stamen seen from within. F. Fruits.
82
cover, the hairs short, to 0.2 mm, straight,erect; ter-
minal buds plump, 3 X 1.5 mm, densely brown to-
mentellose. Leaves alternate, widely and evenly
spaced along currentflush;petioles striate,robust,to
2 X 0.6 cm, semi-terete, the indument as on bran-
chlets; laminaechartaceous,plane, elliptic-ovate,15-
30 X 7-14 cm, the base obtuse to truncate,briefly
decurrent,the apex acute, acuminatefor up to 1.5 cm,
the marginsminutelyrecurvedthroughout,often rev-
olute near leaf base; upper surface olive-brown,the
midrib and secondariesflat to immersed,the higher-
order veins prominulous;lower surface sparsely pu-
bescent, the hairs 0.2 mm long, curved, weakly as-
cending to appressed,uniformlydistributed,all vein
ordersraised, theirprominencedecreasingwith rank;
secondary veins 7-10 per side, ? evenly spaced,
slightly more distantaroundmidlamina,ascendingat
50-60O (more acutely towards apex), arching after
midcourse, distal pairs loop-connected; tertiaries
oblique to midrib, between secondariesonce-forked
to straight. Staminate inflorescences evenly spaced
along currentflush in the axils of foliage leaves, to
15 cm long with 14 lateralbranches,branchorders2-
3, the highest order essentially dichasial, but the in-
termodesreduced and flowers arrangedin pseudo-
umbellate fascicles, the axes densely brown
tomentellose; bracts and bracteoles caducous by
anthesis,lanceolate,the indumentas on axes; pedicels
terete, to 2 mm long, those supporting secondary
flowers slightly shorter.Flowers hypocrateriform,2
mm diam., densely rusty strigillose outside; recepta-
cle deeply infundibuliform,0.7 X 0.5 mm, slightly
constrictedbelow tepals, rusty tomentose inside. Te-
pals chartaceous,ovate, 1 X 0.6 mm, both whorls
equal, spreadingat anthesis,the tips incurved,the in-
ner surface densely rusty tomentellose. Stamens of
whorlsI andII stipitate,0.8 mm tall, the anthersovate,
0.5 X 0.3 mm, glabrous,the apex apiculate,the con-
nectives prolonged between the 2 locelli, these
obliquely hemispherical, introrse-latrorse,the fila-
ments clavate, narrowerthan anthers,densely grey-
tomentose; whorl III stamens broadly stipitate, 0.6
mm tall, the anthersovate, 0.3 X 0.3 mm, erect,locelli
2, extrorse-latrorse,the filamentsas broadas anthers,
wider towardsbase, ligulate, densely grey-tomentose
inside, the basal glandsabsent;whorlIV wanting;pis-
tillode wanting. Pistillate inflorescence with indu-
ment, color, and branchingas in staminateplants,the
flowers slightly deeper; stamens sterile, smaller;te-
pals erect; ovary glabrous,ovoid; style stout, weakly
distinguished from ovary; stigma discoid, 0.5 mm
diam. Fruitsborne on claviformpedicels of up to 1.3
x 0.3 cm; cupules hemispherical,to 1 x 2 cm, gla-
FLORANEOTROPICA
brous inside and outside, the margins entire; drupes
ellipsoid, to 3 X 2 cm.
Distribution (Fig. 27) and ecology. Medium-
sized trees from lowlandforests of westernAmazonia
at ca. 130-215 m. Flowering material known from
June,August, andSeptember,andfruitsfromJanuary.
Representativespecimensexamined.COLOMBIA.
VAUPES:Rio Paca, Wacaricuaraand vic., 215 m, 1-3 Jun
1953 (fl d), Schultes & Cabrera19540 (US).
PERU. LORETO: Maynas,Iquitos,Estaci6nExperimen-
tal IIAP,Allpahuayo,130m, 24 Aug 1988(fl d), vander
Werfet al. 10242(F,G, MO);RfoNanay,Mishana,140m,
10 Jan 1983 (fr), Gentry et al. 39301 (G, MO), Aug 1988
(fl Y), vander Werifet al. 10187(G, MO);Requena,Sa-
puena,Arboreto JenaroHerrera,130m, 18 Sep 1980(fl 6),
Castillo34 (F,MO),170m, 15 Sep 1987(fl 6), Vdsquez &
Jaramillo9593(G,MO,US).
Local names. Peru:anis moena, limon moena.
Endlicheriacitriodorahas ovate-apiculatewhorlI
and II anthers typical of the E. canescens species
group(species 37-45), butthe very shortreddishhairs
that cover its striatebranchletsand petioles are with-
out equal therein. Further,the truncateleaf base is
immediately diagnostic whenever evident. As noted
in the protologue(van der Werff, 1991), the tendency
for ultimatecymes to collapse into pseudo-umbelsis
unusual. Elsewhere in Endlicheria, pseudo-umbels
appearin relativelypauciflorousinflorescencesin E.
acuminataandE. xerampela,species furtherdiffering
from E. citriodora by truncateto emarginateanther
apices.
Collectors of this species consistently report a
strong lemonlike scent and glaucous lower leaf sur-
faces, but the odordisappearsin herbariumspecimens
andthe glaucouscast is seldomretained(e.g., Castillo
34).
40. Endlicheria rubra Chanderbali,sp. nov. Type.
Peru. San Martin:Naranjillo, along rd. between
Rioja and PedroRuiz, 950 m, 22 Mar 1998 (fl d),
van der Werifet al. 15436 (holotype: MO; isoty-
pes: F, G, GH, HBG, MO, NY, QRS, U, US).
Fig. 28
Ex affinitateEndlicheriae canescentis et specierumaf-
finium foliis obtusissimisdistinguenda.
Treesto 25 m. Branchletsrelativelystout,midway
along flush 3-5 mm diam., angular,densely tomen-
tose, the surface barely visible, reddish brown, the
hairs short,to 0.3 mm, straightto crisped, erect, dark
red; terminalbuds slender,2 x 0.6 mm, densely pu-
SYSTENMATIC
TREATMENT
~~~~~~~~~.
V
AR~~~~
4 - ----------
4, Irc
W,
T.~~~~~~~~~~~~~
lv
FI.7.Dsriuin.fEdl.e.a....oaE..r
bescent, the hairs as on branchlets,ascending.Leaves
alternate,evenly spaced along currentflush; petioles
slender,to 1 x 0.2 cm, semi-tereteto canaliculate,the
indumentas on branchlets;laminae stiff chartaceous,
plane, obovate, 8-15 X 2-5 cm, the base acute, the
apex bluntto rounded,the marginsminutelyrecurved
throughout;uppersurfaceolive to reddishbrown,the
primaryto fourth-orderveins raised,theirprominence
decreasingwith rank,but the midrib sunken towards
petiole; lower surface sparsely tomentose, the hairs
erect, straight to curved, dark red, denser on main
veins, all vein orders raised, their prominence de-
creasing with rank;secondary veins 5-8 per side, ?
evenly spaced, slightly more distant aroundmidlam-
ina, ascending at 50 60? (more obtuse aroundmid-
lamina), arcuate,distal pairs loop-connected;tertiar-
ies oblique to midrib,between secondariesstraightto
forked.Staminateinflorescencesevenly spaced along
currentflush in the axils of foliage leaves, to 12 cm
long with 10 lateralbranches,branchorders3-4, the
highest order dichasial, lax, the flowers distant, the
83
I~~~~~~~~~~~~~~~~~~~~.
7
~
Evnticaan recl
axes densely red tomentellose; bracts and bracteoles
caducous by anthesis, lanceolate, the indumentas on
axes; pedicels gradually increasing in diameter api-
cally, to 1.5 mm long, those supporting secondary
flowers slightly shorter.Flowers hypocrateriform,to
2 mm diam., densely rusty tomentellose outside; re-
ceptacle cyathiform, 1 X 1 mm, slightly constricted
below tepals, densely greyish tomentose inside. Te-
pals fleshy, broadly ovate, 0.8 X 0.6 mm (the inner
whorl slightly broader),spreading,the inner surface
densely greyish rusty tomentellose, the hairs erect,
crisped, the margins and apex inside rusty papillose.
Stamens of whorls I and II broadly stipitate, ca. 0.6
mm tall, the anthers ovate, 0.4 X 0.3 mm (slightly
narrowerin whorl II), glabrous, the apex apiculate,
the connectivesprolongedbetween the 2 locelli, these
obliquely hemispherical, introrse-latrorse,the fila-
ments fleshy, slightly narrowerthan anthers,densely
grey-tomentellose;whorl Ill stamens sessile, 0.6 mm
tall, the anthersnarrowlyovate to lanceolate, 0.3 X
0.15 mm, erect, locelli 2, narrow, slitlike, extrorse-
84 FLORA NEOTROPICA

FIG. 28. Endlicheria rubra (van der Weriffetal. 15436).A. Habit. B. Leaf base below. C. Flower. D. Flower I.s. E.
Whorl I stamen seen from within. F. Whorl III stamen seen from without.
SYSTEMATICTREATMENT
latrorse, the filaments apically as broad as anthers,
widening towards base, fleshy, densely grey-
tomentellose, the basal glands sessile, laminar,apic-
ulate; whorl IV wanting;pistillode wanting.Pistillate
inflorescencewith indumentand color as in staminate
plants,but shorterandwith fewer lateralbranches,the
flowers slightly deeper; stamens sterile, smaller;
ovary glabrous, ellipsoid; style stout, weakly distin-
guished from ovary, 0.2 mm long; stigma tri-lobed,
0.5 mm diam. Fruitsunknown.
Distribution (Fig. 27) and ecology. Medium-
sized treesknownfrompoorlydrainedforestsin west-
ern Amazonia and eastern Andean slopes at ca. 800
m. Flowering at least in March,May and July.
Additional specimens examined. BRAZIL. ACRE:
Cruzeirodo Sul, Rio Juruaand Rio Moa, vic. of Porangaba,
17 May 1971 (fl juv), Maas et al. P13058 (HBG, MG, NY,
U). AMAZONAS: Rfo Ituxi, vic. of B6ca do Curuquete,12
Jul 1971 (fl 2), Prance et al. 14150 (HBG, K, MO, NY,
US).
Among species with dense reddish vestitureEn-
dlicheria rubra is immediatelyrecognized by its ob-
ovate leaves with blunt apices and recurvedmargins.
The type materialis made even more conspicuousby
a glaucous cast on the leaves below. This featuredoes
not appearin the Brazilian specimens, possibly lost
through preservationin alcohol, but otherwise they
provide good vegetative matches and the narrowly
ovate anthersof the type also appearin the pistillate
flowers of Prance et al. 14150.
The unusual leaf morphology of Endlicheria
rubra,complete with glaucous cast below, appearsin
a fruiting collection from Amazonian Colombia,
Garcfa-Barriga13944 (NY, US), but four locelli ar-
rangedin a shallow arc in whorl I and II staminodes
assign this specimen to Rhodostemonodaphne.
Garcia-Barriga 13944, apparently representing an
undescribed species, is so similar to Endlicheria
rubra that my initial impressionwas that it provided
the fruitsof this new species. These areimmature,but
as they already show deeply hemisphericalcupules,
this suggests deep floralreceptaclesin pistillateflow-
ers, as found in E. rubra. Given these similarities,it
would be surprisingif the two were not sisterspecies.
41. Endlicheria vinotincta C. K. Allen, Mem. New
YorkBot. Gard. 10(5): 63. 1964. Type.Venezuela.
Amazonas: Rio Yatua, Cerro Neblina, Cafion
Grande, below Cumbre Camp, 1650 m, 26 Nov
1957 (fl ), Maguire et al. 42244 (holotype:NY).
85
Scandentshrubsto 3 m. Branchletsrelativelyslen-
der, midway along flush 2-4 mm diam., distally
weakly angular,soon terete, densely tomentose, the
surface concealed by the indument cover, the hairs
short, to 0.5 mm, crisped, erect, dark red; terminal
buds plump, 5 X 3 mm, reddishtomentose-pannose.
Leaves alternate,widely andevenly spacedalong cur-
rent flush; petioles robust, to 1.5 X 0.4 cm, semi-
terete, the indumentas on branchlets;laminaecoria-
ceous, plane to subbullate,ovate, 8-2 X 3-8 cm, the
base rounded,briefly decurrent,the apex acute, acu-
minate for up to 4.5 cm, the margins minutely re-
curvedthroughout;uppersurfaceolive-brown,waxy,
areolate, the primary to fourth-orderveins raised,
their prominencedecreasingwith rank;lower surface
sparselytomentose,the hairsas on branchletsbutpale
brown, denser on main veins, all vein ordersraised,
their prominence decreasing with rank; secondary
veins 3-4 per side, ? evenly spaced, slightly more
distantaroundmidlamina,patent,divergingat 70-85?
(more acutely towards apex), arcuate, distal pairs
loop-connected;tertiarieslaxly reticulatingbetween
secondaries. Staminateinflorescencesevenly spaced
along currentflush in the axils of foliage leaves, to
15 cm long with 9 lateralbranches,branchorders3-4,
the highest order dichasial, lax, the flowers distant,
the axes reddishtomentose;bractsand bracteolesca-
ducous by anthesis, lanceolate, the indument as on
axes; pedicels terete, to 3 mm long, those supporting
secondaryflowers slightly shorter.Flowers hypocra-
teriform, 5 mm diam., densely reddish tomentose
throughout;receptacleinfundibuliform,0.6 X 1 mm.
Tepalschartaceous,elliptic to obovate, 1.6 X 1.3 mm
(the inner whorl slightly narrower),spreadingto re-
curvedat anthesis.Stamensof whorlsI and II broadly
stipitate,0.8 mm tall, the anthersovate,0.3 X 0.6 mm,
densely reddishtomentoseoutside,the apex glabrous,
truncateto emarginate,the connectives level with or
slightly reducedbetweenthe 2 locelli, these obliquely
hemispherical,introrse, the filaments stout, clavate,
gradually tapering below anthers, rusty tomentose,
provided with a pair of minute sessile basal glands;
whorl III stamens sessile, 0.8 mm tall, the anthers
ovate, 0.3 X 0.6 mm, erect,locelli 2, extrorse-latrorse,
the filamentsas broadas anthers,ligulate,fleshy,rusty
tomentose, the basal glands minute sessile; whorl IV
wanting; pistillode wanting. Pistillate inflorescence
with indument,color, and branchingas in staminate
plants, the flowers similar in size and shape;stamens
sterile, smaller;ovary glabrous,ovoid; style stout, in-
distinct from ovary; stigma discoid, 0.5 mm diam.
Fruitsborne on claviform pedicels of 0.4 X 0.3 cm;
cupules infundibuliform,to 0.7 X 0.4 cm, glabrous
86 FLORA NEOTROPICA

inside and outside, the marginsstronglylobed, tepals 1997 (fl d), Campos & Nuniez4669 (holotype:
persisting;drupesellipsoid, to 1.3 X 0.8 cm. MO; isotypes: F, HBG, NY). Fig. 29

Distribution (Fig. 27) and ecology. Scandent Nova species Endlicheriae vinotinctae proxima, fo-
shrubs from sandstone formations of the Guiana liorumtexturae gris-
et colorisimilis,sedfloribusindumento
Highlands at ca. 350-1650 m. Flowering specimens eis differt.
have been collected in April, October,andNovember,
Treelets or shrubsto 8 m. Branchletsstout, mid-
and the only fruitingspecimen in April.
way along flush 4-6 mm diam., angular, densely
Additional specimens examined. COLOMBIA. greenish yellow tomentose, the surfaceconcealed by
GUAINIA: Maimachi, Serranfa delNaqu6n.CerroMinas,al- the indumentcover, the hairs relatively short, to 0.2
rededoresdel Helipuerto-15 y caminohastala cima del mm, straightto crooked,erect; terminalbuds plump,
Cerro,900 m, 7 Apr 1993 (fl 5), Madrinidn& Barbosa 936 5 X 3 mm, densely pubescent,the hairsas on branch-
(COL), (fr), Madrinidn& Barbosa 948 (MO), (fl 5), Mad- lets, ascending. Leaves alternate,widely and evenly
rinan & Barbosa 953 (MO). spaced along currentflush; petioles slender, to 2 X
VENEZUELA.AMAZONAS: Atabapo,CerroDuida, 0.2 cm, semi-terete,distally furrowed,the indument
cuestade areniscade grandafino al pie del CerroDuida, as on branchlets;laminae coriaceous, subbullateto
360 m, 10 Nov 1982(fl 5), Gudnchez2154 (MO);meseta
plane, ovate, 9-15 X 3-7 cm, the base acute, briefly
de areniscagrandeubicadaal Surdel Rio Matakuni,1050
m, 19 Nov 1991 (fl 5), Huber13261 (MO);Cerrode la decurrent,the apex acute, briefly acuminatefor up to
Neblina,slopeN of RioMawarinuma, above"Pto.Chimo," 0.5 cm, the margins minutely recurvedthroughout;
500 m, 25 Apr 1984 (fl 5), Stein & Gentry 1660 (F, MO, upper surface yellowish green to reddish brown,
US);RioNegro,CerroAracamuni, summit,1550m, 16Oct waxy, areolate, all vein orders prominulous;lower
1987(fl i), Liesner& Delascio22008(MO),1415m, 16- surfacesparselytomentose,the hairsas on branchlets,
18Oct 1987(fl d), Delascio & Liesner 13508 (MO,VEN), scatteredon lamina, denser on main veins, all vein
600 m, 21 Oct 1987 (fl 5), Liesner & Carnevali 22292 ordersraised, theirprominencedecreasingwith rank;
(MO), 1400 m, 25 Oct 1987 (fl 5), Liesner & Carnevali secondary veins 4-6 per side, ? evenly spaced,
22414 (MO).
slightly more distantaroundmidlamina,ascendingat
In Endlicheria, E. vinotincta alone inhabits the 50-60O (moreobtuse aroundmidlamina),arcuate,dis-
summitsand upperslopes of the sandstonemountains tal pairsloop-connected;tertiariesroughlyhorizontal,
(tepuis)of the Guianahighlands.In its scandenthabit, between secondaries once-forkedto straight.Stami-
the rich dark red vestiture that covers its branchlets nate inflorescencesevenly spaced along currentflush
and inflorescences, including inner floral surfaces, in the axils of foliage leaves, to 3 cm long with 3
and stamens all provided with basal glands, this is a lateralbranches,branchorders2-3, the highest order
very distinctive species of Endlicheria.Yet all these dichasial,the flowersdistant,the axes densely yellow-
charactersare matched by the sympatricRhodoste- ish green tomentose;bracts and bracteolescaducous
the indumentas on axes; pedicels
monodaphne celiana. Leaves of Endlicheria vino- by anthesis,ovate,
terete, to 1 mm long, those supporting secondary
tincta have prominenttertiariesabove andpale brown
flowers slightly shorter. Flowers infundibuliform,3
indumentwhile in Rhodostemonodaphneceliana the
mm diam., densely greyish green tomentoseoutside;
tertiariesareimmersedabove anda darkredindument
receptacle broadly infundibuliform, 0.6 X 1 mm,
covers the lower leaf surface, but flowers of the two
densely grey-tomentoseinside. Tepalsfleshy,ovate, 1
differ mainly in locelli number.Another sympatric
X 0.6 mm, ascending at anthesis, the inner surface
species of Rhodostemonodaphne,R. steyermarkiana,
sparsely grey-tomentose.Stamens of whorls I and II
is vegetatively similar, but appears to be an erect
broadly stipitate,0.4 mm tall, the anthersovate, 0.3
shrub (Madrinian,1996b), and, like R. celiana, has
X 0.4 mm, glabrous,the apex apiculate,the connec-
darkred indumenton the leaves below.
tives prolongedbetween the 2 locelli, these obliquely
The faint glaucous cast persisting on lower leaf
hemispherical, introrse-latrorse,the filaments stout,
surfacesin Gudnchez2154 and Huber 13261 hints at
clavate, slightly narrowerthan anthers,densely grey-
a useful field characterfor this species.
tomentose;whorl III stamensstout, sessile, 0.5 m tall,
the anthers oblong, 0.3 X 0.2 mm, erect, locelli 2,
extrorse-latrorse,the filamentsbroaderthan anthers,
42. Endlicheria oreocola Chanderbali, sp. nov. columnar,densely grey-tomentose,the basal glands
Type. Peru. Cajamarca:San Ignacio, Romerillo, sessile, globose; whorl IV wanting; pistillode want-
base of Cordillerade Romerillo, 1570 m, 15 Nov ing. Pistillateinflorescencewith indument,color, and
 FIG. 29. Endlicheriaoreocola (Campos & Nuntez4889). A. Habit. B. Leaf base below. C. Flower. D. Flower 1.s. E.
Whorl I stamen seen from within. F. Whorl III stamen seen from without.
88
branchingas in staminateplants, the flowers similar
in size and shape;stamenssterile, smaller;ovary gla-
brous, ovoid; style slender, distinct from ovary;
stigma discoid, 0.3 mm diam. Fruits borne on clavi-
form pedicels of up to 1 X 0.4 cm; cupules shallowly
infundibuliform,to 0.3 X 1 cm, glabrousinside and
outside, the margins entire; drupes obovoid, to 2 X
1.5 cm.
Distribution (Fig. 27) and ecology. Small to
medium-sized trees or shrubs of pre-montanecloud
forest vegetation from ca. 1500-2500 m on the east-
ern Andean slopes of Peru and Ecuador. Flowers
known from November and January,fruits from Jan-
uary and April.
Additional specimens examined. ECUADOR.
ZAMORA-CHINCHIPE:Zamora,ParqueNacionalPodocar-
pus, carreteraLoja-Zamora,Estaci6n Cientifica San Fran-
cisco, 2250 m, Jan 1995 (fl 6), Palacios & Tirado 13425
(QCNE, MO), (fl Y), Palacios & Tirado 13439 (QCNE,
MO), (fl Y, fr), Palacios & Tirado 13459 (QCNE, MO),
2100 m, 20 Apr 2000 (fr), Neill et al. 12616 (QCNE, MO).
The dense greenish yellow tomentose indument
found on branchlets of the type material of Endli-
cheria oreocola is unmatchedin the genus. A glau-
cous cast appearing on the underside of younger
leaves offers anotherstriking featurebut appearsto
be lost with age, and older leaves attach to reddish
brown tomentose branches.Older leaves of E. oreo-
cola resemble those of E. vinotincta in their coria-
ceous texture, ovate shape, and areolate upper sur-
faces. Yet, this is certainly not its closest relative.
Flowers of E. oreocola are greyish green ratherthan
reddish tomentose outside, and although the shape
and size of the stamens in the two species are much
alike, in E. oreocola only whorl III stamens are pro-
vided with basal glands. Still, as no other species of
Endlicheriais remotely similarit seems best to com-
pare E. oreocola with E. vinotincta, if only to em-
phasize its peculiarities.
43. Endlicheria szyszylowiczii Mez, Jahrb.Konigl.
Bot. Gart. Berlin. 5: 121. 1889. Type. Peru. Ca-
jamarca:Tambillo,without date (fl d), Jelski 165
(lectotype, designated by Kostermans,1937: B-
n.v.; isolectotypes:L, MO, S, US).
Trees to 15 m. Branchlets stout, midway along
flush 4-6 mm diam., distally weakly angular,soon
terete,densely rustyto reddishtomentose,the surface
concealed by the indumentcover, the hairs relatively
long, to 0.6, straightto crisped, erect to ascending;
terminalbuds plump, 2 x 2 mm, densely pubescent,
FLORANEOTROPICA
the hairs rusty to red, ascending. Leaves alternate,
widely andevenly spacedalong currentflush;petioles
slender,to 2.5 X 0.3 cm, semi-terete,the indumentas
on branchlets; laminae coriaceous to chartaceous,
plane to subbullate,ovate to elliptic, 9-25 X 4-10
cm, the base acute to rounded,briefly decurrent,the
apex acute to obtuse, acuminatefor up to 2.5 cm, the
marginsminutelyrecurvedthroughout;uppersurface
dull greyish green to olive-brown,minutely punctu-
late, the primary to fourth-orderveins raised, their
prominence decreasing with rank, or tertiary and
higher-orderveins immersed;lower surface densely
pilose, the hairs soft, erect, up to 1 mm long, rust
brown, slightly shorterand denser on main veins, all
vein ordersraised, their prominencedecreasingwith
rank;secondaryveins 4-6 per side, ? evenly spaced,
slightly more distantaroundmidlamina,ascendingat
50-60? (moreobtuse aroundmidlamina),arcuate,dis-
tal pairsloop-connected;tertiariesroughlyhorizontal,
between secondariesstraight,or forked.Staminatein-
florescencesevenly spaced along currentflush in the
axils of foliage leaves or cataphylls, to 25 cm long
with 14 lateralbranches,branchorders3-4, the high-
est order dichasial, lax, the flowers distant, the axes
densely rusty tomentose;bractsand bracteolescadu-
cous by anthesis, narrowovate to lanceolate, the in-
dument as on axes; pedicels terete, to 2 mm long,
those supportingsecondary flowers slightly shorter.
Flowers depressed-globoseto urceolate, 1.5 to 3 mm
diam., sparsely rusty to grey-strigoseoutside; recep-
tacle deeply cyathiform,1 X 2, densely rusty red ve-
lutinous inside. Tepals chartaceous,broadly ovate, 1
X 1.5 mm (the inner whorl slightly narrower),in-
curvedto erect at anthesis,the innersurfaceminutely
papillose. Stamensof whorlsI andII broadlystipitate,
0.6-1 mm tall, the anthersovate, 0.3-0.5 X 0.3-0.6
mm, glabrous, the apex apiculate, the connectives
prolongedbetween the 2 locelli, these obliquelyhem-
ispherical,introrse-latrorse,the filamentsligulate, al-
most as broad as anthers, densely grey-tomentose;
whorl III stamens sessile, 0.6-1 mm tall, the anthers
depressed-oblong,0.2-0.3 X 0.3-0.5 mm, erect, lo-
celli 2, extrorse-latrorse,the filamentsas broadas an-
thers, wider towards base, the indumentas in outer
whorls, the basal glands sessile, globose; whorl IV
wanting; pistillode filiform. Pistillate inflorescence
with indument,color, and branchingas in staminate
plants, the flowers similarin size and shape;stamens
sterile, smaller;ovary glabrous,ovoid; style slender,
distinctfrom ovary;stigmaminutelytri-lobed,ca. 0.2
mm diam., emerging beyond incurved tepals. Fruits
borneon shortclaviformpedicels of up to 5 X 5 mm;
cupules hemispherical,to 0.5 x 1 cm, glabrousinside
SYSTEMATICTREATMENT 89

and outside, the margins undulate, tepal bases per- cional Alexander von Humboldt, 300 m, 25 Nov 1977 (fl
sisting; drupesellipsoid to obovoid, to 1.5 X 1 cm. ), Froehner67 (HBG, MO). MADRE DE DIos: Tambopata,
Lago Sandoval, 13 km NE of PuertoMaldonado,200 m, 25
Distribution (Fig. 30) and ecology. Medium- Jul 1989 (fr), Nuiiez 11195 (HBG, MO); ParqueNacional
sized trees ranging throughoutthe Amazonian low- "Bahuaja-Sonere,"Ex Santuario Nacional Pampas del
lands and reaching the surroundinghighlands of the Heath, 10 Jul 1997 (fl S), Diaz & Pereira 9020 (MO).
Guianashield to the north,the Brazilianshield to the PAsco: Oxapampa, 1800 m, 4 Mar 1986 (fl d), van der
south, and lower montaneAndes to the west at 200- Werifet al. 8340 (HBG, MO, NY). SAN MARTiN: Rioja,
2200 m. Flowering materialcollected from February Pardo Miguel, Venceremos,CaserfoEl Afluente, Bosque de
through November, fruits in July, and September Protecci6nde Alto Mayo, 1440-1520 m, 14 Nov 1996 (fr),
Sdnchez & Dillon 8687 (MO).
throughNovember.
BRAZIL. AcRE: Brasileia, Reserva ExtrativistaChico
Representative specimens examined. COLOMBIA. Mendes, Seringal Porongaba, 15 Apr 1992 (fl d), Saraiva
ANTIoQJIA: San Antonio de Prado, 2200 m, Nov 1989 (fl & de Lima 1523 (MO); Cruzeiro do Sul, BR 364, km 42,
d ), de Escobar et al. 8851 (HUA). ramal 4 do Projeto Santa Luzia (INCRA), Sep 1985 (fr),
VENEZUELA. AMAZONAS: Rfo Negro, Canyon Rosas et al. 216 (MO, NY). AMAPA:Rio Jari, Sep 1968 (fl
Grandede La Neblina, 5 Feb 1984 (fld ), Funk6102 (MO). d6), Silva 968 (HBG, NY). AMAZONAS: Novo Aripuana,BR
GUYANA. EssEQuIBo: Wassarai Mtns., 14 km S of 230, Rod. Transamaz8nica400 km de Humaita, Projeto
KassikaituR., 1135 m, 10 Sep 1999 (fl Y), Clarke et al. INCRA-RfoJuma,vicinal dos Gadchos,4 May 1985 (fl 6),
8397 (MO). Cid Ferreira et aL 6033 (F, GH, INPA, K, MO, NY, US);
PERU. AMAzONAS: Mendoza, 1600 m, 16 Aug 1963 (fl Presidente Figueiredo, Rio Uatuma, UHE Balbina, 7 Jul
d ), Woytkowski8323 (GH, MO). CAJAMARCA: Tambillo, 1986 (fl Y), Thomaset aL 5370 (F, INPA, K, MO, NY, US).
s.d. (fl juv), Jelski 196 (L). LORETO:Maynas, Bosque Na- MATOGRosso: Sinop, 25 Sep 1985 (fr), Thomas et al.

It

A~~~~~~~~~~~~~~~~~~~~~~~~~

4,~~~~~~~~~~~~~~~~~

FIG~~~~~~~~~~~~~~~~~~~~~~
30DsrbtoFfEdihrasyzlwcuEdoicaadElrseo
90
4059 (HBG, INPA,MO, NY). PARA:Belem, 27 Jul 1944
(fl Y), Silva 309 (NY, UC, US); Rio Mapua, 18 Jul 1950 (fl
6), Black et al. 50-9809 (INPA,NY, US). ROND6NIA: Gua-
jara Mirim, 175 m, 9 Apr 1987 (fl Y), Nee 34692 (F, INPA,
GH, K, MO, NY, US).
BOLIVIA. LA PAZ: Prov. Franz Tamayo, 1600 m, 23
May 1992 (fl Y), Perry 999 (MO); Prov. Larecaja,Copa-
cabana, 10 km S of Mapiri,850-950 m, 8 Oct-15 Nov 1939
(fr), Krukoff11103 (A, F, G, K, MO, NY,S, U, US). SANTA
CRUZ: Prov. Caballero,ParqueNacional Ambor6, 31 Mar
1996 (fl 6), Jardimet al. 2576 (NY).
Local names. Peru: moena, roble fusi. Brazil:
louro de f61hapeluda, louro vermelho.
Endlicheria szyszylowiczii is readily distinguished
from other species with dense reddish tomentose
branchletsby its depressed-globoseto campanulate
flowers with tepals incurved to erect at anthesis. In
vegetatively similar species, ascending tepals may
yield infundibuliform flowers, but horizontally
spreading tepals predominate. At first glance the
higher-elevationmaterial of E. szyszylowiczii (e.g.,
van der Werif et al. 8340) may be mistaken for E.
duotinctabecause the pale lower leaf surfacecontrast-
ing with the erectreddishhairsproducean aspectsim-
ilar to that bestowed by the mixed indumentof the
latter. Despite a wide geographic and altitudinal
range, the only noteworthyinternalvariationappears
as slightly reducedfloralsize andpalerpubescencein
extra-Andeancollections.
44. Endlicheria duotincta Chanderbali, sp. nov.
Type. Peru. Cajamarca:San Ignacio, Huarango,
Nuevo Mundo, road between Caserio Gosen and
Nuevo Progress,1350-1400 m, 23 Jul 1997 (fl d),
Rodrigues & Reyes 1807 (holotype: MO; isotypes:
F, G, HBG, NY, US, U). Fig. 31
Foliis subtusindumentocum Endlicheriaewilliamsiiop-
time congruens,sed floribuscampaniformiset foliis alternis
differt.
Trees to 15 m. Branchlets stout, midway along
flush 4-6 mm diam., angular,densely rusty red to-
mentose, the surface concealed by the indument
cover, the hairs moderatelylong, to 0.5 mm, crooked
to crisped, erect; terminalbuds plump, 2 X 2 mm,
densely pubescent, the hairs as on branchlets, ap-
pressed. Leaves alternate,widely and evenly spaced
along currentflush; petioles robust,to 3.5 X 0.4 cm,
striate, the indument as on branchlets;laminae stiff
chartaceous,plane to subbullate,ovate to elliptic, 10- sized trees of pre-montaneforests from ca. 1300-
30 X 5-13 cm, the base acute to rounded,the apex 1900 m on the easternAndeanslopes of Ecuadorand
acute, acuminatefor up to 1.5 cm, the margins flat Peru. Flowering materialcollected in February,July,
FLORA NEOTROPICA
throughout;uppersurfacelight green to olive-brown,
minutely punctulate, the midrib prominulous, the
higher-ordervenation immersed; lower surface ob-
scured by a short creamish tomentose indument
interspersedwith straighterect reddish hairs of 0.7
mm, all vein ordersraised,theirprominencedecreas-
ing with rank;secondaryveins 6-8 per side, ? evenly
spaced, slightly more distant aroundmidlamina,as-
cending at 50-60o (more acutely towards apex), ar-
cuate, distal pairs loop-connected;tertiariesroughly
horizontal, between secondaries straight or forked.
Staminateinflorescencesevenly spacedalong leafless
flushes in the axils of cataphylls,to 20 cm long with
14 lateral branches, branch orders 2-3, the highest
order dichasial, lax, the flowers distant, the axes
densely rusty tomentose; bracts and bracteoles per-
sistent at anthesis, lanceolate, the hairs as on axes,
appressed;pedicels terete, to 2 mm long, those sup-
porting secondary flowers slightly shorter.Flowers
campanulate,to 2.5 mm diam., densely pubescent
outside, the hairs rusty red, appressedto ascending,
reducedto a basal triangularpatchin the innerwhorl
of tepals; receptacle cyathiform,2 X 1.5 mm, rusty
pilose inside. Tepals chartaceous,ovate, 1.3 X 0.6
mm (the inner whorl slightly narrower),ascendingat
anthesis,the inner surfacesparselypapillose towards
the marginsand apex, otherwiseglabrous.Stamensof
whorlsI and II broadlystipitate,1 mm tall, the anthers
ovate, 0.5 X 0.4 mm, glabrous,the apex apiculate,the
connectivesprolongedbetween the 2 locelli, these su-
borbicular,introrse,the filamentslaminar,hardlynar-
rowerthananthers,densely grey-tomentose;whorlIII
stamensbroadlystipitate,1 mm tall, the anthersovate,
0.5 X 0.4 mm, erect, locelli 2, extrorse-latrorse,the
filamentsalmost as broadas anthers,laminar,the in-
dument as in outer whorls, the basal glands sessile,
globose; whorl IV wanting;pistillode filiform.Pistil-
late inflorescence with indument and color as in
staminateplants, but shorter and with fewer lateral
branches,the flowers slightly deeper;stamenssterile,
smaller, sessile, the filaments as broad as anthers;
ovary glabrous,ellipsoid; style slender,distinct from
ovary, 1 mm long; stigma minutelytri-lobed,0.2 mm
diam. Fruits borne on claviform pedicels of up to 5
mm long; cupules hemispherical,to 1 X 1.5 cm, gla-
brous outside, densely rusty strigose inside, the mar-
gins sharplylobed, tepal bases persisting;drupesel-
lipsoid, to 2 x 1 cm.
Distribution (Fig. 30) and ecology. Medium-
 4~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~

ttI t~ ~ ~ ~ ~ ~ ~ ~~~V

FIG. 31. Endlicheriaduotincta(Rodr(gues& Reyes 1807). A. Habit. B. Leaf base below. C. Flower.D. Flower with
facing tepal turneddown to reveal androecium.E. Flower 1.s. F Whorl I stamen seen from within. G. Whorl HI stamen
seen from without.
92
August, and September,and fruitingmaterialin June
and August.
Additional specimens examined. ECUADOR.
ZAMORA-CHINCHIPE: QuebradaLas Pavas, 1800 m, 21
Aug 1975 (fl Y), Samaniego& Vivar90 (QAME,US); Que-
bradade Los Monos, cerca de Sabanilla,1600 m, 4 Sep 1975
(fl Y, fr juv), Little et al. 221 (COL, QAME, US).
PERU.SAN MARTiN:Rioja, Pedro Ruiz-Moyobamba
rd., km 390, Venceremos,1800-1900 m, 29-31 Jul 1993 (fl
d), Smith4491 (MO), 1790 m, 9-10 Aug 1983 (fr), Smith
& Vdsquez4775 (MO); Rioja-Pomacochasrd., below Ven-
ceremos, ca. 20 km NW of Rioja nearRestaurantEl Amigo,
1600 m, 8 Feb 1984 (fl ?), Gentry & Smith45169 (MO);
vic. of Puente Serranoyacu,1590-1840 m, Aug 1983 (fr),
Luna 112 (MO).
Local name. Ecuador:laurel blanco.
Endlicheriaduotinctais the second species in the
genus in which tomentose indument conceals the
lower leaf surface. This remarkablevestitureis also
found in E. williamsii, but may have originated in-
dependentlythere since its flowers are comparableto
those of E. anomala, while the campanulateflowers
of E. duotincta closely resemble those of E. rufora-
mula of the E. sericea species group.Although a po-
sition in the latteris suggested on floral grounds,the
rusty tomentose branchletsof E. duotinctawould be
anomalousthere, and a place among those with sim-
ilar indument,only less dense on leaves below-i.e.,
the E. canescens species group-is preferreduntil its
affinitiesbecome more clear.
45. Endlicheria lorastemon Chanderbali,sp. nov.
Type. Ecuador. Zamora-Chinchipe:Nangaritza,
Rio Nangaritza,Pachicutza,1000-1200 m, 6 Dec
1990 (fl J), Palacios & Neill 6577 (holotype:MO;
isotypes: F, GH, HBG, NY, QCNE, U, US).
Fig. 32
Species ramulis sub-sericeis foliis subtus pilosis a con-
generis diversa.
Trees, 5-40 m. Branchletsslender,midway along
flush 2-3 mm diam., distally weakly angular,soon
terete, densely pubescent, the surface barely visible
to concealed by the indument cover, the hairs rela-
tively short,to 0.3 mm, straight,appressedto ascend-
ing, yellowish to rustbrown;terminalbuds slender,3
X 1 mm, densely rusty sericeous. Leaves altemate,
widely andevenly spacedalong currentflush;petioles
slender,to 2.5 X 0.15 cm, semi-terete,the indument
as on branchlets;laminae chartaceous,plane, ovate,
10-25 X 4-9 cm, the base acute, briefly decurrent,
the apex acute, acuminatefor up to 4 cm, the margins
FLORA NEOTROPICA
flat throughout;upper surface olive-brown,minutely
punctulate,the primary to fourth-orderveins raised,
theirprominencedecreasingwith rank;lower surface
sparsely pubescent, the hairs as on branchlets,but
erect, denser on main veins, all vein orders raised,
their prominence decreasing with rank; secondary
veins 3-5 per side, + evenly spaced, slightly more
distantaroundmidlamina,ascendingat 50-60? (more
obtuse towards apex), arcuate, distal pairs loop-
connected;tertiariesroughlyhorizontal,between sec-
ondaries forked to straight.Staminateinflorescences
evenly spaced along currentflush in the axils of fo-
liage leaves, to 10 cm long with 8 lateralbranches,
branchorders2-3, the highest orderdichasial,lax, the
flowers distant,the axes densely pubescent,the hairs
reddishto light brown,appressedto ascending;bracts
andbracteolescaducousby anthesis,lanceolate,rusty
sericeous; pedicels terete, to 2 mm long, those sup-
porting secondary flowers slightly shorter.Flowers
hypocrateriform,3 mm diam., densely tawnyto rusty
pubescentoutside, the hairs ascending;receptaclein-
fundibuliform,0.6 X 1 mm, silvery velutinousinside.
Tepals chartaceous,ligulate, 1 X 0.5 mm (the inner
whorl slightly narrower),ascending to spreadingat
anthesis,the inner surfacesparselysilvery tomentose
near base, otherwise glabrous,the marginsminutely
papillose. Stamensof whorlsI and II broadlystipitate,
0.5 mm tall, the anthersdepressed-ovate,0.2 X 0.3
mm, glabrous,the apex broadlyapiculateabove the 2
locelli, these suborbicular,introrse,the filamentslig-
ulate, equaling or slightly narrower than anthers,
densely silvery-greytomentellose;whorl III stamens
columnar,0.6 mm tall, the anthersoblong, 0.3 X 0.2
mm, erect, locelli 2, extrorse-latrorse,the filaments
slightly broader than anthers, wider towards base,
densely silvery-grey tomentellose, the basal glands
stipitate, minute, globose; whorl IV wanting; pistil-
lode filiform. Pistillate inflorescencewith indument,
color, and branchingas in staminateplants, the flow-
ers similarin size and shape;stamenssterile, smaller;
ovary glabrous, ovoid; style stout, weakly distin-
guished from ovary; stigma minutely tri-lobed, 0.3
mm diam. Fruits borue on stout, claviform pedicels
of up to 1 X 0.5 cm; cupules patelliform,to 0.3 X 1
cm, glabrousinside and outside, the marginssharply
lobed, tepal bases persisting;drupes spheroidto ob-
ovoid, to 1 X 1 cm.
Distribution (Fig. 30) and ecology. Small to large
trees from western Amazonia and adjacent lower
montaneeastern Andean slopes, at ca. 120-1200 m.
Flowering specimens have been collected continu-
ously from August throughJanuaryand fruits from
Januaryto June.
SYSTEMATICTREATMENT 93

3 cm ~~~~~~~~~~

FIG. 32. Endlicheria lorastemon (Palacios & Neill 6577). A. Habit. B. Leaf base below. C. Flower. D. Flower 1.s.
E. Whorl I stamen seen from within. F. Whorl mIstamen seen from without.
94
Additional specimens examined. COLOMBIA. AMA-
ZONAS: Rio Apaporis, Soratama,near mouth of Rio Kan-
anari,250 m, 14 Dec 1951 (fl 9), Schultes& Cabrera14903
(COL,U); Rio Caqueta,Sep 1989 (fljuv), Urregoet al. 1012
(MO). CAQUETA:Araracuara,22 Dec 1993 (fl 3), Vester&
Roman865 (COL, MO).
ECUADOR. NAPO:Aguarico, Reserva Etnica Huaor-
ani, carreteray oleoducto de Maxus, km 108, 235 m, 18 Jan
1995 (fr), Aulestia & Omehuat3236 (MO), km 72, 270 m,
23-31 Jan 1994 (fr), Dik & Andi 1001 (MO, QCNE-n.v.),
km 75-76, entre el Rio Tivacunoy Rio Yasuni, 17-20 Feb
1994 (fr), Aulestia & Gonti 1755 (MO); Orellana,Parque
Nactional Yasuni, 230 m, 9-13 Jan 1987 (fl d), Ceron &
Coello 3257 (MO); Tena, Estaci6n Biol6gica JatunSacha,
450 m, 17 Aug 1993 (fl d), Palacios & Tipaz11074 (MO).
PASTAZA: Rio Chullana,ca. 15 km N of Puerto Sarayacu,
16 Oct 1974 (fl 9), Lugo 4188 (GB); Rio Curiacu,ca. 8 km
W of PuertoSarayacu,19 Oct 1974 (fl 9), Lugo 4241 (GB);
Rio Bobonaza, Caimito, between Puerto Sarayacuand Pa-
cayacu, 25 Oct 1974 (fl 9), Lugo 4302 (GB), ca. 4 km E of
Pacayacu, 31 Oct 1974 (fl 9), Lugo 4394 (GB), between
Pacayacuand Canelos, 3 Nov 1974 (fl 9), Lugo 4450 (GB);
Via Auca, 115 km al S de Coca, cerca del Rio Tigiiino, 320
m, 29 Apr 1989 (fr), Rubio 8 (MO). SANTIAGO-ZAMORA:
Cordillera Cutucd, 1600 m, 17 Nov-5 Dec 1944 (fl d),
CampE-1165 (NY). ZAMORA-CHINCHIPE:Nangaritza,Rio
Nangaritza,Miazi, 1000 m, 9 Dec 1990 (fl 3), Neill & Pa-
lacios 9618 (MO), (fl d), Palacios & Neill 6669 (MO), (fl
d), Palacios & Neill 6688 (MO), 20 Oct 1991 (fl 3), Pa-
lacios et al. 8492 (COL, MO), Pachicutza,6 Dec 1990 (fl
3), Palacios & Neill 6572 (MO); Zamora,ParqueNacional
Podocarpus,GuarderiaRio Bombuscaro,sendero al Mira-
dor, 1100 m, Jan 1995 (fl d), Palacios & Tirado 13310
(MO).
PERU. LORETO: Maynas, Iquitos, Puerto Almendras,
UNAP, 122 m, 17 Jan 1993 (fl 3), Grdndez et al. 5512
(MO).
BRAZIL. ACRE:Cruzeirodo Sul, sub-base do Projeto
RADAM/BRASIL, 8 Mar 1976 (fr), Ramos & Mota 350
(INPA);Mancio Lima, 12 Oct 1989 (fl 9), Cid Ferreiraet
al. 10031 (INPA,NY); Serrodo Moa, Local Central,30 Sep
1984 (fl 3), Cid Ferreiraet al. 5094 (F, INPA, K, MO, NY,
US); Rio Moa, 8 km above CachoeiraGrande,27 Apr 1971
(fr), Prance et al. 12563 (HBG, MO, NY). AMAZONAS:Rio
Solimoes, Tefe, entradado Lago Tefe, Santa Missoes, 16
Oct 1982 (fl d), Amaral et al. 109 (INPA, MO); Parandde
Tefe, 16 Oct 1982 (fl d), Cid Ferreira& Lima 3270 (INPA,
K, MO, R); Fonte Boa, sub-base do Projeto RADAMI
BRASIL, CartaSA-l9-XD-PT? 06, 4 Jun 1976 (fr), Ramos
450 (INPA).
Local names. Ecuador: ocatoe, yahuemomo (both
are Huaorani names).
Endlicheria lorastemon typically has straplike
whorl I and II stamens where filaments equal anthers
in width, but occasional stipitate stamens (e.g.,
Amaral et al. 109 and Cid Ferreira & Lima 3270)
reduce the diagnostic value of an otherwise unique
FLORA NEOTROPICA
feature. Still, no other species of Endlicheria com-
bines erect hairs on the leaves below with a dense
subsericeouscover of appressedto ascendinghairson
the branchlets.
As for Endlicheriaduotincta,affinitiesareunclear
since the subsericeous branchlets of E. lorastemon
recall the E. sericea species groupwhile the claviform
pedicels below shallow platelike cupules resemble
those of E. anomala.
46. Endlicheria sericea Nees, Linnaea 8: 38. 1833.
Goeppertiasericea (Nees) Nees, Syst. laur. 369.
1836. Type. Trinidad.Without locality and date
(fl), Sieber 175 (holotype:B-n.v.; isotypes: BM-
n.v., E, G, GH, L, LZ-n.v., MO, NY-n.v., P, W-
n.v.).
Goeppertiasericea (Nees) Meisn. var.opaca Meisn. DC.
Prodr.15 (1): 174. 1864.Type.Dominica.Without
localityanddate(fl),Imray457 (holotype:K-n.v.).
Endlicheria guadaloupensis Mez, Jahrb.Konigl. Bot.
Gart.Berlin5: 124. 1889.Type.Guadeloupe.
With-
out locality and date (fr), Duchassaing s.n. (holo-
type:B-n.v.isotype:W-n.v.).
Treesto 20 m. Branchletsrelativelystout,midway
along flush 3-5 mm diam., distally weakly angular,
soon terete, silvery to golden sericeous, the surface
concealed by the indumentcover, the hairs short, to
0.2 mm, straight,appressed;terminalbuds plump, 3
X 2.5 mm, sericeous. Leaves alternate,widely and
evenly spaced along currentflush;petioles slenderto
robust, to 2 X 0.3 cm, semi-terete,the indumentas
on branchlets; laminae coriaceous to chartaceous,
plane, ovate, 10-20 X 2-10 cm, the base obtuse,
briefly decurrent,the apex acute, acuminatefor up to
2 cm, the marginsminutelyrecurvedthroughout;up-
per surface light olive-green, waxy, the midrib flat,
sunken towards petiole, drying dark, conspicuous
against the lamina, the secondaries immersed;terti-
aries prominulous;lower surface densely sericeous,
the hairs as on branchlets,uniformlydistributed,all
vein ordersraised, their prominencedecreasingwith
rank;secondaryveins 4-5 per side, ? evenly spaced,
slightly more distantaroundmidlamina,or basalpairs
closer, subopposite, all ascending at 50-60? (more
obtuse aroundmidlamina),arcuate,distal pairs loop-
connected;tertiariesroughlyhorizontal,between sec-
ondaries once-forked to straight. Staminate inflor-
escences evenly spaced along current flush in the
axils of foliage leaves or cataphylls, to 20 cm long
with 15 lateralbranches,branchorders3-4, the high-
est order dichasial, lax, the flowers distant, the axes
silvery to golden sericeous;bractsand bracteolesca-
SYSTEMATICTREATMENT 95

ducous by anthesis, lanceolate, the indument as on 690 m, 21, 27, & 28 Mar 1978 (fr), Steyermark& Davidse
axes; pedicels terete, to 2 mm long, those supporting 116957 (G, MO); Los Guayabitos(arribade Baruta), 1350
secondary flowers slightly shorter.Flowers rotate,to m, Mar 1958 (fr), Aristeguieta3012 (MO, NY). TACHIRA:
4.5 mm diam., densely silvery sericeous outside; re- Cerro Las Minas, 17 km SE of Santa Ana, 1150-1250 m,
Nov 1979 (fr), Steyermarket al. 119895 (G, MO, VEN).
ceptacle shallowly cyathiform,2 X 4 mm, silvery pi-
lose inside. Tepals fleshy, ovate, 1.6 X 1 mm, spread- Local names. Dominica:bois marble,laurierbord
ing, the androeciumexserted at anthesis, the inner de mer, laurier p6te. St. Lucia: laurier gris, laurier
surfacedensely grey-tomentose,the marginsandapex gwa gwen, laurier marbre,sweetwood. St. Vincent:
inside rusty papillose. Stamens of whorls I and II sweetwood.
broadly stipitate,0.6 mm tall, the anthersovate, 0.4
X 0.3 mm, glabrous,the apex apiculateor truncateto Endlicheriasericea is distinguishedfrom all other
emarginate, the connectives prolonged between, species where silvery to golden sericeous indument
obscures the lower leaf surface by its rotate flowers
broad above, or level with the 2 locelli, these subor-
with fleshy spreadingtepals thatbear a dense greyish
biculate, introrse-latrorse, the filaments laminar,
tomentose cover on the inner surfaces. Stout clavi-
slightly narrowerthan anthers, basally grey-pilose;
form pedicels below hemispherical cupules and a
whorl III stamens sessile, 0.7 mm tall, the anthers
sunkenmidribin the lower laminaarealso diagnostic.
oblong, 0.4 X 0.3 mm, erect, locelli 2, extrorse-
This species has been collected most frequentlyin
latrorse,the filamentsas broad as anthers,columnar,
the Lesser Antilles and the type is from Trinidad,but
grey-pilose, the basal glands sessile, globose; whorl
presence on the South American mainland is sug-
IV wanting; pistillode wanting. Pistillate inflores-
gested by three collections from the VenezuelanAn-
cence with indumentand color as in staminateplants,
des (viz., Steyermark& Davidse 116957, Steyermark
but shorterand with fewer lateralbranches,the flow-
et al. 119895, andAristeguieta3012). All arefruiting
ers slightly deeper; stamens sterile, smaller; ovary
specimens that are vegetatively indistinguishable
glabrous; style slender, distinct from ovary; stigma
from insular material and, moreover,show truncate
tri-lobed, 0.3 mm diam. Fruits borne on stout
antherapices in whorl I and II staminodes.Exceptfor
claviformpedicels of up to 2 X 0.5 cm; cupuleshem-
E. griseo-sericea with elliptic to obovate leaves and
ispherical,to 1 X 1.3 cm, glabrousoutside, sericeous
much larger cupules, and E. bracteolata with tripli-
inside, the margins entire to undulate;drupes ellip-
nervedleaves, all otherspecies with densely sericeous
soid, to 2 x 1.3 cm.
lower leaf surfaceshave definitely apiculateanthers.
Distribution (Fig. 33) and ecology. Medium- Endlicheria guadaloupensis is based on fruiting
sized trees from lower montaneforests in the Lesser material from Guadeloupe (Duchassaing s.n.) that
Antilles, Trinidad,and northernAndes in Venezuela could not be located.FromMez's (1889) descriptions
at ca. 300-1350 m. Flowering throughoutthe year, and key to species it is apparentthathe distinguished
fruits collected from FebruarythroughApril and in this species from E. sericea by ellipsoid ratherthan
October. ovoid fruits and nonpersistenceof tepals on cupule
margins.Without floral correlatesthe specific value
Representativespecimens examined.DOMINICA. of these differences is questionable,and Kostermans
Laudat, Jul1881(fl 6), Eggers403 (G,GH,HBG,P);Castle (1937), who examinedthe type of E.
guadaloupensis,
BruceTrail,400 m, 25 Feb 1946(fl ?, frjuv),Beard628
consideredit conspecific with that of E. sericea. His
(A, MO, NY, U).
GUADELOUPE. Basse-TerrenearDuclos, PetitBourg, synonymy
is maintainednot only because the differ-
200 m, 30 Mar 1956 (fr), Smith 10356 (K, NY); Pidgeon, ences in fruit shape are ratherslight, but also because
21 Aug 1973 (fl ?), Sastre & Sastre 2046 (NY, P). tepal persistenceappearsto be temporaryin E. seri-
MARTINIQUE. Trace des Jesuites NW of Fond St.- cea.
Denis, entranceE of Morne Rouge-Fond St. Denis rd., 450 The type material of E. sericea appearsto be a
m, 22 Jul 1987 (fl Y), Daly 5293 (MO, NY); MamneRose, mixed collection since the E, GH, and P duplicates
600 m, 28 Sep 1984 (fl 2), Rollet 1672 (A). bear staminateinflorescences,the MO duplicatepis-
ST. LUCIA. Savanne Edmund district, SE of Piton tillate inflorescences,andthe G duplicatediseasedin-
Troumass6e,1800-2000 ft, 20 Nov 1960 (fi 2, fr), Procter florescences.
21585 (A); Zeno Top Soufriere, 3 Oct 1986 (fr), Pierre &
Slane 357 (A).
ST.VINCENT.SilverSpoon,aboveThreeRivers,1-7
Apr 1960 (fl 2), Howard11145 (GH, NY); Mt. St. Andrews 47. Endlicheria bracteolata (Meisn.) C. K. Allen,
summit, Feb 1972 (fr) Howard & Howard 18005 (A, NY). Mem. New York Bot. Gard. 10(5): 64. 1964.
VENEZUELA. MIRANDA: Cerros del Bachiller, 200- Goeppertia sericea (Nees) Nees var. bracteolata
96
- e-- . . . . .. .--
FIG.
FIG 33
Meisn. DC. Prodr. 15(1): 174. Type. Venezuela.
Amazonas: San Carlos, Rio Negro, 1853-4 (fl c,
juv), Spruce 3092 (syntypes: B-n.v., BM-n.v., E,
G, GH, L, NY, OXF-n.v., P-n.v., U, W).
Trees to 15 m. Branchletsslender,midway along
flush 3-4 mm diam., distally weakly angular, soon
terete, silvery to golden sericeous, the surface con-
cealed by the indumentcover, the hairs rathershort,
to 0.15 mm, straight,closely appressedto the surface;
terminalbuds plump, 3 X 3 mm, sericeous. Leaves
alternate, widely and evenly spaced along current
flush; petioles slender, to 1.5 X 0.3 cm, semi-terete,
the indument as on branchlets;laminae chartaceous
to coriaceous, plane, ovate, 10-20 X 5-8 cm, the base
obtuse to rounded,briefly decurrent,the apex acute,
acuminatefor up to 2 cm, the margins minutely re-
curvedthroughout;uppersurfacelight green to olive-
brown, the midrib immersed, secondary and tertiary
veins prominulous;lower surface densely silvery to
golden sericeous, the hairs uniformly distributed,all
vein orders raised, their prominencedecreasing with
FLORA NEOTROPICA
* * - -- -.-
.,,~ ~ ~ ~ ~~.
33. Distributionof Endlicheria sericea arld E. bracteolata.~ .7'
Distribution of Endlicheria Iserice
bracteota. and E
rank; secondary veins 2-4 per side, the lowermost
pair(s) subopposite shortly above the leaf base, as-
cending at 5060O, arcuate, distal pairs loop-
connected;tertiariesroughlyhorizontal,between sec-
ondaries straightor forked. Staminateinflorescences
all evenly spaced along leafless flushes in the axils of
cataphylls, or the distal ones in the axes of foliage
leaves, to 10 cm long with 10 lateralbranches,branch
orders 3-4, the highest orderdichasial, lax, the flow-
ers distant,the axes golden sericeous;bractsandbrac-
teoles caducous by anthesis, lanceolate, sericeous;
pedicels terete, to 1 mm long, those supportingsec-
ondary flowers slightly shorter.Flowers infundibuli-
form, 2 mm diam., golden sericeous outside; recep-
tacle infundibuliform, 1 X 1 mm, densely silvery
pilose inside. Tepals chartaceous,ovate to ligulate, 1
x 0.5 mm (the inner whorl slightly broader),ascend-
ing, surroundingandroecium at anthesis, the inner
surface moderately silvery tomentose, the margins
minutely papillose. Stamens of whorls I and II stipi-
tate, 0.8 mm tall, the anthers depressed-elliptic to
SYSTEMATICTREATMENT 97

ovate, 0.5 X 0.5 mm, glabrous,the apex roundedto ori and Crique Sapokay,26 Jul 1968 (fl Y), OldemanT-35
truncate,the connectives broad above the 2 locelli, (MO, P), entre le Saut et la crique Couata, 31 Jul 1968 (fl
these suborbiculate, introrse-latrorse,the filaments Y), OldemanT-55 (NY, P, U).
laminar,narrowerthananthers,densely silvery pilose; PERU. LORETO:Maynas, Rfo Nanay, Puerto Almen-
dras, 122 m, 4 Jun 1986 (fl d), Vdsquez& Jaramillo 7644
whorl III stamens sessile, 0.8 mm tall, the anthers
(MO); Upper Rio Mazan, near Base Araguana,8 Jul 1976
oblong, 0.4 x 0.3 mm, erect, locelli 2, extrorse-
(fl d), Gentry & Revilla 16539 (F, HBG, MO).
latrorse,the filamentsbroaderthan anthers,laminar, BRAZIL. AMAPA: Colonia do Torrao,28 Aug 1962 (fl
densely silvery pilose, the basal glands sessile, glo- d ), Pires & Cavalcante52649 (NY, US); Rio Oiapoque,Rio
bose, apiculate;whorl IV staminodial;pistillode fili- Ingarari,15 Sep 1960 (fl Y), Irwin et al. 48290 (F, NY, U).
form. Pistillateinflorescencewith indumentandcolor AMAZONAS: Jauarete,Vaupes, Rio Negro, 22 Oct 1945 (fl
as in staminateplants, but shorterand with fewer lat- d), Froes 21243 (F, MO, NY); Manaus,Cachoeirabaixa do
eral branches,the flowers campanulate;stamensster- Taruma, 12 Sep 1966 (fl Y), Prance et al. 2269 (F, GH,
ile, smaller; ovary glabrous; style slender, distinct HBG, INPA, K, MG, NY, R). PARA:Porto Trombetas,12
from ovary; stigma tri-lobed, 0.3 mm diam. Fruits Aug 1986 (fl d), Soares 189 (INPA).
borne on narrowlyclaviformpedicels of up to 0.5 X Local names. Colombia:jigua. Venezuela:dimu-
0.2 cm; cupules hemispherical,to 0.5 X 0.7 cm, gla- kuima(Ye'kwana),laurelblanco, laurelplateado,lau-
brous outside, sericeous inside, the margins entire; rel rebalsero.FrenchGuiana:cedre. Brazil:louroced-
drupesellipsoid, to 1 X 0.5 cm. inha.
Distribution (Fig. 33) and ecology. Medium- Endlicheria bracteolata is easily distinguished
sized trees from seasonally inundatedlowland and from otherspecies with sericeous indumentby its tri-
lower montaneforests (50-1000 m) throughoutnorth- plinerved leaves. Occasional productionof a second
ern South America. Flowers and fruits availableyear pair of basal secondaries shortly above the leaf base
round. results in quintuplinervationsimilar to that frequent
in E. sericea. Indeed,Meissner(1864) consideredthis
Representative specimens examined. COLOMBIA.
species a mere varietyof E. sericea, butrotateflowers
AMAZONAS: Rio Apaporis, Jirijirimo,250 m, 25-26 Nov
in the lattercontrastinginfundibuliformflowers with
1951 (fl d), Garcfa-Barriga13703 (COL, NY, US), 27 Nov
1951 (fl d), Schultes & Cabrera 14606 (GH, U). ANTIO- only ascending tepals in E. bracteolata supportsAl-
QUiA: Urrao, Parque Nacional Natural "Las Orquideas," len's (1964) elevation to specific status.The inflores-
1300-1500 m, 3 Apr 1992 (fl 6), Cdrdenas& Alvarez3267 cence bracteolesto which the specific epithetalludes
(MO), Sector Calles, margenderechadel Rfo Calles y de la are presenton the type materialand other specimens
quebrada"El Guaguo,"12 Feb 1989 (fl Y, fr juv), Cogollo with immatureinflorescences,but fall by the onset of
et al. 3928 (MO), camino de Venados arribahacia Calles, anthesis. Specimens from Valle and Antioquiain Co-
30 Jul 1988 (fl ?), Cogollo et at. 3638 (MO). VALLE: Barco, lombiahave slightly darkerindumentandtendto have
Rio Cajambre,Apr 1944 (fr), Cuatrecasas 17216 (F, US); larger leaves but otherwise cannot be distinguished
Buenaventura,Caserfo"SanIsidro,"CONIF,100-150 m, 23
from materialeast of the Andes.
Mar 1992 (fr), Cogollo et al. 5127 (INPA, MO), 25 Mar
1992 (fl juv), Cogollo et al. 5129 (MO).
VENEZUELA. AMAZONAS: Atabapo, Rio Asisa, 100
m, Oct 1989 (fl d), Delgado 835 (MO);Atures,PuertoAya- 48. Endlicheria tschudyana (Lasser)Kosterm.,Bol.
cucho, Rio Cataniapo, 100-110 m, 11 Nov 1980 (fl d), Tecn. Inst. Agron. N. No. 28, Addendapost p. 75.
Gua'nchez388 (HBG, MO, VEN). BOLiVAR: Cedeno,20 km 1953. Aniba tschudyanaLasser, Bol. Soc. Venez.
E of Turiba,6-11 Dec 1970 (fl d), Marcano-Berti2581 (G, Ci. Nat. 11(72): 181. 1948. Type. Venezuela.Ar-
MO); Rio Las Ahallas, 14 Mar 1985 (fr), Liesner 18665 agua:ParqueNacional de RanchoGrande,14 Feb
(MO, NY). 1946 (fl Y), Lasser 2036 (holotype:VEN; isoty-
GUYANA. ESSEQUIBO: Cuyuni-MazaruniRegion, Pa-
pes: MO, VEN).
ruima Falls to ParuimaMission, Kamarang-Wenamu Trail,
600 m, 8 Aug 1951 (fl Y), Maguire& Fanshawe32463 (GH, Trees to 12 m. Branchlets stout, midway along
MO, NY, U, US); KamarangR., Utschi River mouth, 500 flush 5-7 mm diam., angular,silvery to pale yellow
m, 20 Oct 1960 (fl Y), Tillett & Tillett 45705 (F, K, NY,
sericeous, the surface concealed by the indument
US); KurupungR., MakrebaFalls, 24 Feb 1939 (fr), Pinkus
263 (F, G, GH, MO, NY, U, US). cover, the hairs short,to 0.1 mm, straight,appressed;
SURINAME. Tafelberg,610 m, 3 Sep 1944 (fl 6), Ma- terminalbuds plump, 4 X 3 mm, sericeous. Leaves
guire 24712 (A, K, MO, NY, U, US); MarowijneR., Wane alternate, widely and evenly spaced along current
Ck., Feb 1918 (fr), Gonggrijp3692 (U). flush; petioles robust,to 4 X 0.4 cm, semi-terete,the
FRENCH GUIANA. Fleuve Approuague,GrandCan- indument as on branchlets;laminae coriaceous, or
98
chartaceous,plane, ovate to elliptic, 13-35 X 8-15
cm, the base obtuse to subcordate,briefly decurrent,
the apex acute, acuminatefor up to 1.5 cm, the mar-
gins minutely recurved throughout; upper surface
greyish green to olive-brown,the primaryto fourth-
order veins raised, their prominencedecreasingwith
rank;lower surfacedensely sericeous, the hairs as on
branchlets, uniformly distributed, all vein orders
raised, their prominence decreasing with rank; sec-
ondaryveins 5-8 per side, + evenly spaced, slightly
more distantaroundmidlamina,ascendingat 50-60'
(more acutely towardsapex), arcuate,the lowermost
pairs sometimes patent, diverging at 70-85?, and
abruptlyascendingaftermidcourse,distal pairsloop-
connected;tertiariesroughlyhorizontal,between sec-
ondaries once-forkedto straight.Staminateinflores-
cences evenly spaced along currentflush in the axils
of foliage leaves or cataphylls,to 12 cm long with 8
lateralbranches,branchorders3-4, the highest order
dichasial, lax, the flowers distant, the axes rusty to
grey-sericeous;bractsandbracteolescaducousby an-
thesis, ovate, the indumentas on axes; pedicels terete,
to 3 mm long, those supportingsecondary flowers
slightly shorter.Flowers cyathiform to infundibuli-
form, to 3 mm diam., silvery-grey to rusty sericeous
outside;receptaclecyathiformor infundibuliform,1.5
X 2.5 mm, grey-piloseinside. Tepalschartaceous,el-
liptic, 0.8 X 0.5, erect to ascending, surroundingan-
droeciumat anthesis,the outersurfacedensely silvery
or rusty sericeous, the indumentreduced to a basal
triangularpatch in the inner whorl, the inner surface
glabrous or sparsely grey-strigillose. Stamens of
whorls I and II narrowly stipitate, 0.5 mm tall, the
anthersovate, 0.3 X 0.3 mm, glabrous,the apex apic-
ulate, the connectivesprolongedbetweenthe 2 locelli,
these suborbicular, introrse-latrorse,the filaments
laminar,narrowerthananthers,grey-piloseor tomen-
tose; whorl III stamens columnar,0.6 mm tall, the
anthers depressed-oblong,0.2 X 0.3 mm, erect, lo-
celli 2, extrorse-latrorse,the filamentsas broadas an-
thers, columnar,the indumentas in outer whorls, the
basal glands sessile, globose; whorl IV wanting;pis-
tillode fusiform. Pistillate inflorescence with indu-
ment and color as in staminateplants,but shorterand
with fewer lateral branches, the flowers slightly
deeper;stamenssterile, smaller;ovary glabrous;style
stout, weakly distinguishedfrom ovary; stigma dis-
coid, 0.3 mm diam. Fruitsborue on stout terete ped-
icels of up to 0.8 X 0.3 cm; cupules shallowly hem-
ispherical, to 0.8 X 1.5 cm, glabrous outside and
inside, the marginsundulateto lobed, tepal bases per-
sisting; drupesellipsoid, to 1.7 X 1.2 cm.
FLORA NEOTROPICA
Distribution (Fig. 34) and ecology. Small trees
of lower montane and cloud forests from NW South
America to Panama at ca. 300-1600 m. Flowering
and fruitingspecimens collected throughoutthe year.
Additional specimens examined. PANAMA. BOCAS
DEL TORO:Along rd. to ChiriqufGrande,10 road-mifrom
continental divide, 300 m, 9 Feb 1987 (fr), McPherson
10444 (MO). COCLE:Near sawmill 16.7 km N of turnoff
to Coclesito from Llano Grande,700 ft, 7 Mar 1978 (fl 5),
Hammel 1858 (MO). DARIEN: Alturasde Nique, 850-1100
m, 24 Aug 1987 (fl 5), McPherson 11582 (HBG, MO);
Trochade RanchoFrio, 900 m, 23 Sep 1989 (fl Y), Aranda
et al. 978 (MO). PANAMA: CerroJefe, 650 m, 27 Aug 1986
(fl Y), McPherson10000 (MO), 750-850 m, 19 Jul 1987 (fl
Y), McPherson11310 (MO), 800 m, 13 Jan 1988 (fl 9 juv,
fr), McPherson 11936 (MO), 900 m, 27 Dec 1986 (fl), Val-
despino et al. 278 (MO).
COLOMBIA.ANTiOQuIA: San Francisco, carreteraa
Aquitania,500-900 m, 8 Apr 1990 (fl 5, juv), Ca'rdenaset
al. 2645 (MO); San Luis, carreterahacia Aquitania,a 12 km
de la Autopista Medellin-Bogota, 850 m, 24 Nov 1988 (fl
5), Cogollo et al. 3735 (MO).
VENEZUELA. ARAGUA:ParqueNacional Henri Pit-
tier,4 Aug 1991 (fl 5, juv), Cardozoet al. 1822 (MO), Fila
de Paraiso,above Portachuelo,1500 m, 5 Jul 1963 (fr), Stey-
ermark 91524 (K, NY), Periquito Trail, 18 Jun 1962 (fr),
Allen 20 (NY), Mar 1959 (fr), Aristeguieta3845 (NY), Pico
Periquito, 12 Dec 1979 (fr), Manara s.n. VEN 172708
(VEN), 1250-1600 m, 4 Sep 1960 (fl 5, juv), Steyermark
& Agostini 7 (F, NY, US), Rancho Grande,Aug 1963 (fr),
Aristeguieta5107 (VEN), 15 Jun 1962 (fr), Lasser & Allen
13 (NY), 29 May 1975 (fr), Ferrari& Benitezde Rojas 1456
(F). CARABOBO:Aut6nomo Mora, cuenca hidrograficadel
Rio Moran,700-1100 m, 3-5 May 1991 (fr), Diaz & Ninio
289 (MO); Rio San Gian, arribade La Toma, 750-850 m,
30 Mar 1966 (fl 5, juv), Steyermark& Steyermark95347
(G, NY, VEN). YARACUY: CerroLa Chapa, 1200-1400 m,
9-10 Nov 1967 (fl 9), Steyermarket al. 100298 (HBG,NY),
21 Oct 1982 (fl 5, juv), Davidse et al. 20840 (MO); San
Felipe, SerraniaSanta Maria-CerroLa Chapa, 1150-1250
m, 12 Aug 1992 (fr), Meier & Walter-Weisbeck 2586 (MO).
ECUADOR. NAPO: Estaci6n Biol6gica Jatun Sacha,
Rio Napo, 8 km al E de Mishualli,450 m, 4 Sep 1987 (fr),
Cer6n et al. 2090 (MO), 3-4 Jun 1988 (fr), Neill et al. 8475
(MO); ParqueNacionalYasuni,Rio Tiputini,200-300 m, 5
Oct 1996 (fl 5), Romolerouxet al. 2567 (MO). SUCUMBiOS:
Lago Agrio, Reserva Faunistica Cuyabeno, Tarapoa-
Tipishca, cruce del Rio Cuyabeno,230 m, 14 Nov 1991 (fl
5), Palacios et al. 8938 (MO, NY, US), 8940 (MO).
PERU. HUANUCO: Pucallpa, Sira Mtns., 800 m, 9 Feb
1988 (fl 5), Morawetz& Wallnofer13-9288 (MO).
Local names. Colombia: laurel bobo, laurel
bongo.
The Venezuelan type material of Endlicheria
tschudyanawas takenfrom a pistillateplantwith pat-
SYSTEMATIC TREATMENT
..
W'
'At~
'2
-J ~ ~ ~ ~
FIG 3 Disributon o Endlchera tscudyaa
ent, almost perpendicular,secondary veins in the
lower lamina of cordiform leaves. Such leaves are
found again in Aristeguieta3845, but all other spec-
imens from the type locality have ovate to elliptic
leaves with more ascending secondaries. Since flow-
ers from the type locality are either pistillate or im-
mature,the concept of E. tschudyanaadoptedhere is
largelybased on plantsfrom elsewhere.Althoughthis
recourse increases its geographicalrange considera-
bly, it is not withoutjustification.In the type material,
the floral receptacle is infundibuliformand incurved
tepals close aroundthe stigma. These flowers are un-
usual but are matched by pistillate plants from Pan-
ama (e.g., McPherson 10000). It thereforebecomes
plausible that vegetatively indistinguishable stami-
nate plants from Panama, Hammel 1858 and Mc-
Pherson 11582, provide maturestaminateflowers of
E. tschudyana.However,the two offer slightly differ-
ent manifestationsof this, with Hammel 1858 show-
ing cyathiform, and McPherson 11582 infundibuli-
99
"V. '~~~~~~
.. ~ - -
~
~.
andE grseo srice
form, flowers. As the most advanced stage of floral
developmentfound in the type locality, Steyermark&
Agostini 7 shows an infundibuliformreceptacle, it is
likely that McPherson 11582 provides the more typ-
ical representation.Furthermore,with infundibuli-
form staminate flowers also in Cogollo et al. 3735
from Colombia, the disjunction between Venezuela
and Panamais considerablyreduced.
Yet, it is possible that flower shape is unstable in
E. tschudyana.An apparentcorrelationbetween cy-
athiform flowers and robust branchlets with coria-
ceous leaves can be disregardedsince three collec-
tions from Ecuador,Palacios et al. 8938 and 8940,
and Romoleroux et al. 2567, combine cyathiform
flowers with slender branchlets and relatively thin
leaves. Furthermore,branchletsand leaves also vary
in the type locality. With both floral shapes accom-
modatedin E. tschudyana,a staminateplant identical
to Hammel 1858 in both vegetative and floral terns,
Morawetz& Wallnofer13-9288, extends the rangeof
100 FLORA NEOTROPICA

the species to Peru. With these specimens all in- grey-sericeous inside. Tepals chartaceous,narrowly
cluded,E. tschudyanaaccommodatesall of the E. ser- ovate to triangular,1.5 X 0.5 mm (the inner whorl
icea species group material with silvery sericeous slightly narrower),erect to spreadingat anthesis,the
branchlets,pinnate venation, campanulateor infun- outer surface densely pubescent, the hairs as on re-
dibuliformflowers,with erect to ascending,internally ceptacle, reduced to a basal triangularpatch in the
glabrous tepals, ovate antherswith apiculate apices, inner whorl, the inner surface grey-tomentose, the
and shallowly hemisphericalcupules. marginsand apex inside minutelypapillose. Stamens
of whorls I and II stipitate,0.6 mm tall, the anthers
obovate, 0.3 X 0.3 mm, glabrous,the apex truncate,
the connectives broad above the 2 locelli, these su-
49. Endlicheria griseo-sericea Chanderbali,sp. nov. borbicular,introrse,the filaments laminar,narrower
Type. Ecuador.Napo: Archidona,VolcainSumaco, than anthers,glabrous;whorl III stamens sessile, 0.6
km 50 along road between Hollin and Loreto, mm tall, the anthersoblong to obovate,0.4 X 0.3 mm,
GuaguaSumaco Community,1300 m, 5 Feb 1992 erect, locelli 2, extrorse-latrorse,the filaments as
(fl 6), Rubio & Alvarado 2395 (holotype: MO; broadas anthers,columnar,glabrous,the basal glands
isotypes: F, G, GH, HBG, K, MO, NY, P, QCNE, sessile, globose; whorl IV wanting;pistillode want-
U, US). Fig. 35 ing. Pistillate inflorescencewith indumentand color
as in staminateplants, but shorterand with fewer lat-
Ex affinitateEndlicheriaesericeae et specierumaffinium eral branches,the flowers similar in size and shape;
floribusurceolatiset antherisobovatis distinguenda. stamens sterile, smaller, glabrous or the filaments
grey-tomentose;ovary glabrous; style slender, dis-
Trees to 30 m. Branchlets stout, midway along tinct from ovary; stigma minutely tri-lobed, 0.3 mm
flush 5-7 mm diam., angular,densely grey to yellow- diam. Fruitsbome on stotit claviformpedicels of up
ish sericeous, the surfaceconcealed by the indument to 1.5 X 0.4 mm; cupuleshemispherical,to 1.5 X 2.5
cover, the hairs short, to 0.2 mm, straight,appressed cm, glabrous outside, sericeous inside, the margins
to ascending; terminalbuds plump, 0.4 X 0.4 mm, entire;drupesellipsoid, to 4.5 X 2.5 cm.
densely sericeous. Leaves alternate, widely and
evenly spaced along currentflush; petioles robust,to Distribution (Fig. 34) and ecology. Medium-
4 X 0.3 cm, semi-terete, the indument as on bran- sized to large trees of lower montaneAndeanforests
chlets; laminae chartaceous, or membranaceous, at ca. 800-1500 m. Flowering specimenscollected in
plane, elliptic to obovate, 15-30 X 5-13 cm, the base August, November, and February, fruits in April,
obtuse, or acute, briefly decurrent,the apex acute, or May, June, October,November,and December.
obtuse, acuminatefor up to 2 cm, the marginsmin-
utely recurvedthroughout;upper surface darkolive- Additional specimens examined. ECUADOR. Mo-
brown,the primaryto fourth-orderveins raised, their RONA-SANTIAGO: Cordillerade Cutucu,Mendez-Morona
Rd., 800 m, 4 Feb 1989 (fl Y), van der Werif& Palacios
prominence decreasing with rank; lower surface
10385 (MO). NAPO:Archidona, faldas al sur del Volcan
densely grey to yellowish sericeous, the hairs as on
Sumaco, carreteraHollin-Loreto, km 45, 1100 m, 10 Nov
branchlets, sparser on main veins, all vein orders 1989 (fl Y), Palacios 4747 (G, K, MO, NY), km 50, Gua-
raised, their prominence decreasing with rank; sec- gua Sumaco, 1000 m, 29 Apr-2 May 1989 (fr), Cer6n &
ondaryveins 7-9 per side, ? evenly spaced, slightly Hurtado 6642 (MO), 1100 m, I May 1993 (fr), Neill et al.
more distantaroundmidlamina,ascending at 50-60? 8953 (G, MO, NY), km 31, Comuna ChalluaYacu, 1200
(moreobtusetowardsapex), arcuate,distalpairsloop- m, 4 Oct 1989 (fr), Palacios & Iguago 4541 (MO); carre-
connected;tertiariesroughlyhorizontal,between sec- teraHollin-Loreto-Coca, km 40, 1200 m, 11 Dec 1987 (fr),
ondaries straightor forked. Staminateinflorescences Neill et al. 8094 (HBG, K, MO, NY, US), km 25, 1230 m,
evenly spaced along leafless flushes in the axils of 10-19 Nov 1988 (fr), Hurtado & Alvarado 963 (MO); Or-
cataphylls, to 18 cm long with 12 lateral branches, ellana, Parque Nacional Yasunf, carreteray Oleoducto de
Maxus en construcci6n,km 53-54, 250 m(?), 13-16 Sep
branch orders 3-4, the highest order dichasial, the
1993 (fr), Dik 405 (MO). PASTAZA: Pastaza, via Puyo-
flowers loosely crowded,the axes densely pubescent,
Macas, Pitirishca, 800 m, Aug 1993 (fl 6, juv), Palacios
the indumentas on branchlets;bracts and bracteoles 11001 (F, MO).
persistentat anthesis,ovate, sericeous;pedicels terete, PERU. SANMARTiN: Rioja, km 399 of carreteraMar-
to 3 mm long, those supporting secondary flowers ginal, trail to QuebradaVenceremosand Rio Serranoyacu,
slightly shorter.Flowers urceolate,.to2.5 mm diam., 67 km E of Pomacochas, 8 km W of bridge over Rio Ser-
densely yellowish tomentellose; receptacle cyathi- ranoyacu,1400-1500 m, 13 Jun 1985 (fr), Knapp& Alcorn
form, 2 x 3 mm, constricted below tepals, densely 7778 (MO).
 I0~~~~~~~~~

FIG. 35. Endlicheria gniseo-sericea (Rubio & Alvarado 2395). A. Habit. B. Flower with facing tepal turneddown
to show androecium.C. Flower I.s. D. Whorl I stamen seen from within. E. Whorl HI stamen seen from without.
102
Local name. Ecuador:sacha palta (Quichua).
Endlicheria griseo-sericea is a remarkablenew
species in the E. sericea species group,differingfrom
all other members by its urceolate flowers with the
receptacle strongly constricted below spreading te-
pals, and obovate antherswith broadlytruncateto re-
tuse apices in whorl I and II stamens.The character-
istic darkolive-browncolor assumedby the upperleaf
surfacein the dry stateis a useful diagnosticcharacter,
and in plants with a shiny greyish sericeous vestiture
the contrastis especially striking.
Co-occurring with Endlicheria griseo-sericea in
Ecuadorbut reaching furthernorth to Colombia is a
vegetativelysimilarentity with flowersof similarsize
and shape, but in which whorls I and II anthersare
three-locellateandwhorlIIIanthersarefour-locellate.
I have refrainedfrom describingthis material,prefer-
ring instead to annotate it as cf. E. griseo-sericea,
because in the only staminatecollections, Acevedoet
al. 1370 (F, MO, US), Henao 9 (COL), and van der
Wertf& Palacios 9246 (HBG, MO, NY), the anthers
are devoid of pollen and the unusualandroeciummay
be a consequenceof disease or hybridization.Several
pistillateplantsare known-Boom & Beardsley8449
(MO, NY), Campos & Campos 3092 (MO) Cogollo
et al. 2531, 6556, 7061, & 7062 (MO), Henao 21 &
281 (COL), Jorge & Torres1270 (COL), Neill et al.
12614 & 12615 (MO), Pipoly et al. 17315 & 17453
(MO) Vdsquez& Rojas 21920 (MO)-and whether
in flower or fruit, locelli numberis as in staminate
plants,but I cannotrule out the possibility thatpollen
donors were two-locellate.
50. Endlicheria ruforamula Chanderbali,sp. nov.
Type.Peru.San Martin:Rioja, along roadbetween
Rioja and Pedro Ruiz, 850-1050 m, 26 Mar 1998
(fl d), van der Werifet al. 15736 (holotype:MO;
isotypes: AMAZ, F, G, GH, HBG, K, MO, NY, P,
QCNE, QRS, U, US). Fig. 36
Ex affinitate sericeaeet specierum
Endlicheriae affinium
et ramulistomentellisdistinguenda.
floribuscampanulatis
Trees to 40 m. Branchlets stout, midway along
flush 5-6 mm diam., angular,densely tomentellose,
the surfaceconcealedby the indumentcover,the hairs
short,to 0.15 mm, straight,ascending,reddishbrown;
terminalbuds plump, 5 X 4 mm, densely pubescent,
the hairs as on branchlets.Leaves alternate,widely
andevenly spacedalong currentflush;petioles robust,
to 5 X 0.4 cm, semi-terete, striate, the indumentas
on branchlets;laminaecoriaceous,plane, ovate to ob-
FLORA NEOTROPICA
ovate, 15-30 X 5-15 cm, the base obtuse to acute,
briefly decurrent,the apex acute to obtuse, acuminate
for up to 2 cm, the margins minutely recurved
throughout;upper surface greyish green to olive-
brown,waxy, the primaryto fourth-orderveins raised,
their prominencedecreasingwith rank;lower surface
densely pubescent, the hairs appressed, concealing
surface,rusty red to golden yellow, reducedon main
veins, all vein orders raised, their prominence de-
creasing with rank;secondaryveins 5-7 per side, ?
evenly spaced, slightly more distantaroundmidlam-
ina, ascending at 50-60o (more obtuse aroundmid-
lamina),arcuate,the lowermostpairsometimesasym-
metric (one or both merging with margin at base),
distal pairs loop-connected; tertiaries roughly hori-
zontal, between secondariesforked. Staminateinflo-
rescences distally clusteredin the axils of cataphylls,
or evenly spaced along currentflush in the axils of
foliage leaves or cataphylls, to 25 cm long with 30
lateralbranches,branchorders3-4, the highest order
dichasial, lax, the flowers distant, the axes rusty to-
mentellose; bracts and bracteoles persistentor cadu-
cous by anthesis,triangular,the indumentas on axes;
pedicels terete, to 4 mm long, those supporting
secondary flowers slightly shorter.Flowers campan-
ulate, to 3 mm diam., rusty to greyish tomentellose
outside; receptacle cyathiform,2 X 2 mm, rusty pi-
lose inside. Tepals chartaceous,ovate, 1 X 0.7 mm
(the innerwhorl slightly narrower),erectto ascending
at anthesis, the outer surface rusty to greyish
tomentellose-sericeous,this reducedto a basal trian-
gular patch in the inner whorl, the inner surfacegla-
brous. Stamensof whorls I and II stipitate,1 mm tall,
the anthersovate, 0.5 X 0.5 mm, glabrous,the apex
broadly apiculate,the connectives bluntly prolonged
between the 2 locelli, these suborbiculate,introrse-
latrorse,the filamentslaminar,narrowerthananthers,
glabrousor basally grey-tomentose;whorlIIIstamens
sessile, ca. 1 mm tall, the anthersdepressed-oblong,
0.3 X 0.5 mm, erect, locelli 2, extrorse-latrorse,the
filaments as broad as anthers, ligulate, glabrous or
sparsely grey-tomentosenear base, the basal glands
sessile, globose; whorl IV wanting; pistillode well-
developedor wanting.Pistillateinflorescencewith in-
dument and color as in staminateplants, but shorter
and with fewer lateral branches,the flowers slightly
deeper; stamens sterile, smaller or equaling those of
staminateflowers;ovary glabrous;style slender,dis-
tinct from ovary; stigma weakly tri-lobed, 0.2 mm
diam. Fruitsborue on stout terete pedicels of up to 1
x 0.4 cm; cupules hemispherical,to 1.5 x 2.5 cm,
glabrousoutside, rusty sericeous inside, the margins
entire;drupesellipsoid, to 4.5 X 2.5 cm.
 IL~~I

vC ( -IFT Gh
FIG. 36. Endlicheria ruforamula(A-G, van der Werffet al 15736; H, Vdsquezet al 11954). A. Habit.B. Leaf base
below. C. Flower. D. Flower with facing tepal turneddown to reveal androecium.E. Flower l.s. F. Whorl I stamen seen
from within. G. Whorl III stamen seen from without. H. Fruits.
104 FLORA NEOTROPICA

Distribution (Fig. 37) and ecology. Medium- & Aulestia 975 (MO, NY); Quininde,Fundaci6nParaisode
sized to large trees of flooded and nonflooded Ama- Papagayos, Centro de Rescate de Aves y Mamiferos, 6-8
zonian lowlands, easternlower montaneslopes of the Jul 1996 (fr), Clark et al. 2784 (MO). NAPo: Aguarico,
Andes, and Pacific coast of Colombia and Ecuadorat Reserva Etnica Huaorani,carreteray oleoducto de Maxus,
km 92-96, 250 m, 20 Mar 1994 (fl 9, fr juv), Aulestia &
ca. 100-1200 m. Flowers andfruitsavailablethrough-
Gonti 1986 (MO); Estaci6n Experimental INIAP-Napo,
out the year. Payamino, Reserva Floristica "El Chuncho,"250 m, 5 Apr
Additional specimens examined. COLOMBIA. AMA- 1986 (fr), Jaramillo 8372 (MO), 250 m, 10-11 Sep 1986
ZONAS:Leticia, Tarapacd,Parque Nacional Natural Ama- (fr), Palacios & Neill 1289 (HBG, MO, NY); Estaci6nCien-
cayacu, Rio Cotuhe, 100 m, 19 Jun 1991 (fl d), Rudas et tfficaYasuni, 200-300 m, 15 Oct 1998 (fl 9), Romoleroux
al. 2116 (MO), CabafiaLorena (Inderena)y CainoLorena, et al. 3502 (MO); La Joyade los Sachas, Pompeya,carretera
100 m, 21 Jun 1991 (fl d), Rudas et aL 2262 (NPA, MO), de Maxus, km 1-5, 220 m, 23-29 Nov 1992 (fr), Grijalva
100 m, 21 Jun 1991 (fl d), Rudas et al. 2288 (MO), Cabania et al. 243 (MO); Aiiangu, ParqueNacionalYasunf,260-350
Pamat6;CanioPamate, 100 m, 27 Jun 1991 (fl d), Rudas et m, Apr 1986 (fr), SEF 8740 (NY); Tena,Estaci6nBiol6gica
al. 2588 (MO). CHOC6:EntreCurundey San Jose de Pal- JatunSacha, 450 m, 8 Nov 1987 (fl 6), Cero$n2600 (HBG,
mar, 500 m, 30 Aug 1976 (fl 6), Foreroet al. 2379 (COL, K, MO, NY), 450 m, 17-24 Feb 1988 (fr), Cer6n 3704
MO). VALLE: Cordillera Occidental, Rio Anchicaya, 230- (MO), 400 m, 27 Nov 1992 (fr), Zuleta 42 & 50 (MO); Via
260 m, 13 Oct 1943 (fr), Cuatrecasas 15292 (F, U, US); Payamino-Loreto,250 m, 13 Sep 1986 (fl d, juv), Zaruma
Buenaventura,Caserfo "San Isidro," Reserva Forestal de 695 (MO). PASTAZA: Pastaza, 300 m, 21-28 Feb 1990 (fl
CONIF, 30-100 m, 23 Mar 1992 (fr), Cogollo et al. 5128 cI), Zak & Espinoza 4835 (MO, NY), Curaray,290 m, 13-
(GH, MO). 30 Nov 1990 (fr), Espinoza & Coba 588 (MO); Arajuno,
ECUADOR. ESMERALDAS:San Lorenzo, Parroquia Ri- 700 m, 3-14 Sep 1998 (fl c3), Freire & Santi 3294 (MO).
caurte,CentroPambilar,500 m, 21 Jan 1993 (fl d), Aulestia PERU. HuANuco: Pachitea,Dantas, trochasde estudio

4~~~~~

Z;
-----------3-DstiutonofE-l-hri rfoam l
SYSTEMATICTREATMENT
UNA La Molina, km 42, 22 Jan 1987 (fr), Diaz & Baldeon
2304 (MO, NY). JUNiN: Satipo, Gran Pajonal, E of Che-
quitavo on trial to Kotampaz, 1200 m, Apr 1984 (fl Y),
Smith6764 (MO). LORETO: Alto Amazonas,Rio Huallaga,
Shucushuyacu,250 m, 13 Sep 1981 (fr), Vdsquez& Jar-
amillo 2441 (F, MO, NY); Maynas, CaserfoGamitana,Re-
serva del Rfo Mazan, 116 m, 21 Jun 1990 (fr), Grdndezet
al. 1611 (MO), (fl ?), Grdndezet al. 1612 (MO); Iquitos,
Estaci6n ExperimentalIIAP, Allpahuayo, 130 m, 24 Aug
1988 (fl 6, juv), van der Weriffetal. 10237 (MO); carretera
Iquitos-Nauta,km 44, 150 m, 5 Apr 1989 (fr), Vdsquezet
al. 11954 (MO); Punchana,Rfo Mom6n, 2 Jul 1998 (fl 6),
Rimachi12301 (MO); Rfo Tigre-Rio Corrientes,Cocha Be-
lem, 116 m, 20 May 1987 (fl 6), Grandez& Chiquispama
982 (MO); SargentoLores, ConstanciaNorte, 116 m, 17 Apr
1997 (fr), Vdsquezet al. 23419 (MO). MADRE DE DIos:
Tambopata,Cusco Amaz6nico, 200 m, 8 Oct 1991 (fr), Ti-
mana & Jaramillo2476 (MO);Zona Reservede Tambopata,
280 m, 13 Aug 1990 (fr), Reynel & Meneses 5089 (MO), 14
Aug 1990 (fl 6), Reynel & Meneses 5125 (MO), Hermosa
Chica, Nativa de Infierno,260 m, 19 Dec 1990 (fl 6, juv),
Pesha 68 (K, MO).
BRAZIL. ACRE:Mancio Lima, ParqueNacional Serra
do Divisor, Rio Azul, 11 May 1996 (fl 6), Daly et al. 9024
(MO), (fr), Daly et al. 9025 (MO); Manoel Urbano, Rio
Purus, margemesquerda,seringal Nova Olinda, Colocaqao
Nova Olinda, 24 Nov 1966 (fr), Silveira et al. 1565 (MO);
Marechal Taumaturgo,Rio Jurua,Reserva Extrativistade
Alto Jurua,200 m, 31 Mar 1993 (fr), Daly 7656 (MO);Peix-
oto, 13 Aug 1989 (fr), Santos et al. 39 (INPA);Rio Macau-
han (tributaryof Rio Yacu), 4 Aug 1933 (fr), Krukoff5279
(F, G, MO, NY, U, US); Sena Madureira,Rio Macaua,Fa-
zenda Sao Jose, 30 Mar 1994 (fr), Daly et al. 8104 (MO),
SeringalCapital,7 Oct 1978 (fr), Lima & Sousa 230 (INPA).
AMAZONAS: Coari, Rio Urucu, 7 Dec 1993 (fr), Aguiar et
al. RUC-122 (INPA);ibid., Rio Urucu-Petrobras,SantoAn-
tonio, Porto Helio, 2 May 1988 (fl 6), Matos et al. 207
(INPA);Humaita,estradaHumaita-Ldbrea,km 80, Igarape
do Riozinho, 4 Jun 1982 (fl 6), Filho 82-2N & 82-3N
(HBG), near Tres Casas, 14 Sep-il Oct 1934 (fr), Krukoff
6116 & 6406 (A, F, G, K, MO, NY, U, US); Itapiranga,Rio
Pitinga, 24 Aug 1979 (fr), Cid Ferreiraet al. 690 (F, HBG,
INPA, K, MO, NY, US); Manaus, Cachoeirinha, 1929 (fl
6 ), Ducke 2480 (U); Novo Japura,entreTamandaree Man-
guari, Rio Japura,12 Nov 1982 (fr), Cid Ferreira & Lima
3624 (F, MO, NY, US); PresidenteFigueiredo,Rio Uatuma,
UHE Balbina, 23 Apr 1987 (fr), Bilby et al. 230 (MO); Rio
Cunhua,Deni IndianVillage, 29 Nov 1971 (fr), Prance et
al. 16527 (HBG, INPA, NY, US); Rio Ituxi, 8 Jul 1971 (fl
Y, fr juv), Prance et al. 14013 (F, HBG, INPA, NY, US);
Sao Paulode Oliven,a, 13 May 1945 (fl Y, frjuv), de Lemos
Froes 20899 (NY, US). PARA: Rio Brancode 6bidos, Barro
Vermelho,27 Dec 1913 (fr), Ducke s.n. M.G. 15252 (NY);
Rio Tapaj6s,31 May 1923 (fl 6), Ducke s.n. R 17542 (U);
Rio Itapacaru,4 Apr 1924 (fl Y, fr), Kulhmanns.n. R 18360
(U). RONDONIA: Ji-Parana,Gleba G, km 3, 29 Mar 1983 (fl
6), Silva 6059 (MO, NY), km 2, 29 Mar 1983 (fl 6), Filho
83-30 (HBG), 31 Mar 1983 (fr), Filho 83-49 (HBG), km 3,
105
Mar 1983 (fr), Silva 6078 (MG); PortoVelho, 18 Jun 1986
(fr), Cid Ferreira 7485 (K, MO, NY).
BOLIVIA. PANDO:Madre de Dios, 165 m, 21 May
1991 (fl Y, fr), Killeen 3880 (MO); Manuripi,35 km al N
de Puerto America, 200 m, 19 May 1994 (fr), Jardim 764
(MO); Triunfo, 54 km SW Cobija, 250 m, 3 Aug 1988 (fl
Y), Penningtonet al. 80 (F, MO).
Local names. Ecuador:ocatue (Huaorani),hon-
catohue (Huaorani), urcuayua (Quichua), plata gi-
gante. Peru: inchaquito, moena hoja ancha. Brazil:
louro branco, louro de folha cinzenta, louro pichuri,
louro seda.
Endlicheriaruforamulais distinguishedfrom oth-
ers of the E. sericea species groupby its campanulate
flowers and rusty tomentellose branchletsand inflo-
rescences. Further,its large hemispherical cupules
with entire margins dry a characteristicbrick-red
color that is unmistakableamong the darkbrown to
blackish cupules of similar size and shape produced
by E. griseo-sericea, E. sericea, and E. tschudyana.
Still, the species circumscribedby these threepe-
culiarities is a variable one. In flowers of the type
material and other staminateplants from lower An-
dean montane regions (e.g., Cer6n 2600 and Forero
et al. 2379), inflorescencebracts and bracteolesper-
sist at anthesis.Furthermore,the pistillode is remark-
ably well-developed and providedwith a clearly dif-
ferentiated, albeit small, stigma. However, in most
staminate plants from lowland Amazonia, inflores-
cence bractsandbracteolesarelost at anthesisandthe
pistillode is absent from flowers. The latter are also
slightly smaller and have greyish ratherthan reddish
irndument.Yet, a narrower species concept than
adopted here would provide two entities of which
only staminateplants may be distinguished.A third
group of specimens (e.g., Vdsquez et al. 11954),
mostly in fruit, have asymmetric basal secondaries
thatoriginatefrom thejunctionof the midriband leaf
base in a manner reminiscent of E. metallica and
members of Rhodostemonodaphnein the R. grandis
species group (sensu Madrifinan, 1996b). This vena-
tion is very distinctive and may be sufficient to cir-
cumscribe an easily recognizable species. However,
in the only staminateplant with this venation(Freire
& Santi3294) inflorescencebractsandbracteolesper-
sist at anthesis,and flowers, complete with pistillode,
are indistinguishablefrom typical material.
Representativesof Endlicheria ruforamulawere
previouslyassignedto E. lhotzkyi,a species thatmain-
tains an indument of much longer hairs, has rotate
flowers, and has smallerfruits in shallow patelliform
cupules with lobed margins.
106 FLORANEOTROPICA

51. Endlicheria aurea Chanderbali,sp. nov. Type. tomentose;whorl III stamenscolumnar,0.5 mm tall,
Bolivia. La Paz: Prov. Nor Yungas, 13.7 km NW the anthers ovate, erect, dimensions as in outer
of San Pedro, 1500 m, 15-16 Jan 1983 (fl d), whorls, locelli 2, subapical,the filamentsas broadas
Solomon 9255 (holotype: MO; isotypes: K, NY, anthers,columnar,densely grey-tomentose,the basal
U, US). Fig. 38 glands sessile, globose; whorl IV wanting;pistillode
wanting. Pistillate inflorescence with indument and
Exaffinitate
Endlicheriaesericeaeet specierum
affinium
color as in staminate plants, but shorter and with
locellisantherarum sub-apicalibus
distinguenda.
fewer lateralbranches,the flowers similarin size and
Trees, 4-25 m. Branchletsslender,midway along shape; stamens sterile, smaller;ovary glabrous;style
flush 3-4 mm diam., angular,densely golden to red- slender,distinctfrom ovary;stigmatri-lobed,0.3 mm
dish sericeous, the surfaceconcealedby the indument diam. Fruitsborne on slenderteretepedicels of up to
cover, the hairs short,to 0.2 mm, straight,appressed; 0.7 X 0.2 cm; cupules shallowly hemispherical,0.3
terminalbuds plump, 3 X 3 mm, the indumentas on x 1 cm; glabrousoutside and inside, the marginsun-
branchlets. Leaves alternate, widely and evenly dulate;drupesellipsoid, 1.5 X 1 cm.
spaced along currentflush;petioles slender,to 1.5 X
0.2 cm, semi-terete,striateabove, the indumentas on Distribution (Fig. 39) and ecology. Small to
branchlets;laminaechartaceousto coriaceous,plane, medium-sized trees of eastern Andean forests from
X
ovate to lanceolate,9-28 3-10 cm, the base obtuse Colombia to Bolivia at ca. 850-1530 m. Flowering
to acute, briefly decurrent,the apex acute, acuminate from OctoberthroughJanuary,fruitscollected in Jan-
for up to 2 cm, the margins minutely recurved uary and late April to early May.
throughout;uppersurfacedull greyish green to olive-
Additional specimens examined. COLOMBIA. Pu-
brown, waxy, shining, the primary to fourth-order TUMAYO: Mocoa, San Antonio, Alto Campucaca,La Mar-
veins raised, their prominencedecreasingwith rank; iposa, 1350 m, 20 Apr-1 May 1994 (fr), Pilar Franco et al.
lower surfacedensely sericeous, the hairs as on bran- 5499 (MO).
chlets, uniformly distributed,all vein orders raised, ECUADOR.ZAMORA-CHINCHIPE: Nangaritza,
Miazi,
their prominence decreasing with rank; secondary Rio Nangaritza,1200 m, 10 Dec 1990 (fl d), Palacios 6732
veins 4-5 per side, ? evenly spaced, slightly more (MO, NY); Zamora,ParqueNacional Podocarpus,1400 m,
distantaroundmidlamina,ascendingat 50-60? (more Jan 1995 (fl Y, fr juv), Palacios & Tirado13342 (MO).
obtuse aroundmidlamina),arcuate,distal pairs loop- PERU. PUNO:Moro, Jan 1866 (fl d), Pearce s.n. (K 52,
connected;tertiariesroughlyhorizontal,betweensec- MO 1611827).
BOLIVIA. LA PAZ: Prov.Larecaja,Cocacabana(about
ondaries straightto once-forked. Staminateinflores-
10 km south of Mapiri),850-950 m, 8 Oct-15 Nov 1939 (fl
cences evenly spaced along currentflush in the axils
5, juv), Krukoff11262 (A, F, G, K, MO, NY, U, US); Prov.
of foliage leaves, to 10 cm long with 10 lateral Nor Yungas,4 km NE above Incahuara,13.5 km above San
branches,branchorders3-4, the highest orderdicha- Pedro, 1500-1530 m, 23 Jan 1984 (fl d), Gentry& Solomon
sial, lax, the flowers distant, the axes densely rusty 44524 (MO); Mapiri, San Carlos, 850 m, 11 Dec 1926 (fl
sericeous;bractsandbracteolescaducousby anthesis, d), Buchtien 745 (HBG, NY, US), 9 Jan 1927 (fl d), Buch-
lanceolate, rusty sericeous; pedicels terete, to 1 mm tien 746 (GH), 31 Jan 1927 (fl 5), Buchtien 748 (F, NY,
long, those supporting secondary flowers slightly US).
shorter. Flowers infundibuliform,to 3 mm diam.,
rusty sericeous outside; receptacle shallowly infun- Endlicheria aurea has a most unusual androecium
dibuliform,0.5 X 1 mm, densely rusty pilose inside. where, in stamens of all whorls, depressed-ovatean-
Tepals chartaceous, ovate, 1 X 0.6 mm (the inner thers with almost apically positioned locelli sit atop
whorl slightly narrower),erect to ascendingat anthe- densely pubescentfilaments.Whorl I and II anthers,
sis, the tips slightly incurvedover stamens,the outer though never whorl III anthers,may be ovate in sim-
surface densely rusty sericeous (distal margins and ilarly sericeous species, but locelli are nevertheless
apex of whorl II glabrous),the inner surfacedensely more introrsethan apical. The position of the locelli
grey-tomentose(distal margins and apex of whorl I and the densely pubescent filamentsare reminiscent
glabrous). Stamens of whorls I and II stipitate, 0.5 of stamens in Aniba, and as is typical of that genus,
mm tall, the anthersovate, 0.3 X 0.4 mm (whorl II in E. aurea the tepals surroundthe stamensat anthe-
slightly smaller), glabrous, the apex apiculate, the sis. Advantagesof having locelli placed closer to the
connectives slightly prolongedbetween the 2 locelli, tops of stamensin such flowers seem obvious, but of
these obliquely hemispherical, subapical, the fila- the several species of Endlicheria with erect tepals
ments laminar, narrower than anthers, grey- only E. aurea has such anthers.
 I X
fA, H~~~~~~~~~~~~~~~3c
000
~~o'

0.5mm0

FIG. 38. Endlicheria aurea (Solomon 9255). A. Habit. B. Single leaf showing venation. C. Leaf base below. D.
Close up of lower leaf suface. E. Flower from above. F. Flower with facing tepal turned down to reveal androecium.G.
Flower l.s. H. Whorl III stamen seen from without. I. Whorl I stamen seen from within.
108 FLORA NEOTROPICA

*1~~~~~~~~~~~~~A

FIG 39 Distributionof Endlicheriaaurea E lhorzkyi and E klugu

........ ... \ ---- 4--- - --- ---

r~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~M

FIG. 39. Distribution of Endlicheria aurea, E. Ihotzkyi, and E. klugEi.~~~~~~~~~~~~~..

.. .. . . . .

Kostermans(1937) noted denser indumentinside straight to crooked, weakly appressedto ascending,

flowers of Buchtien's collections but assigned them yellowish to rusty red; terminal buds plump, 4 X 5
(and Pearce's) to Endlicheria lhotzkyi, a species of mm, the indumentas on branchlets.Leaves alternate,
Brazilian cerradowith rotate flowers, a conventional widely andevenly spaced along currentflush;petioles
androecium, and an indumentum of much longer, slender, to 2 X 0.3 cm, semi-terete, the indumentas
more ascending hairs. on branchlets;laminaecoriaceous, plane, ovate, 9-25
X 4-10 cm, the base obtuse to acute, briefly decur-
rent, the apex acute, acuminate for up to 1 cm, the
52. Endlicheria Ihotzkyi (Nees) Mez, Jahrb.K6nigl. marginsminutely recurvedthroughout;uppersurface
Bot. Gart. Berlin 5: 122. 1889. Ocotea Ihotzkyi dull greyish green to olive-brown,waxy, the primary
Nees, Syst. laur.475. 1836. Strychnodaphnelhotz- to fourth-orderveins raised, their prominence de-
kyi (Nees) Meisn., D.C. Prodr. 15(l): 143. 1864. creasing with rank;lower surface densely golden se-
Type. Brazil. Mato Grosso: Cujaba, Nov (fl 9), riceous, the hairs as on branchlets,uniformlydistrib-
Manso & Lhotzky84 (lectotype, designated by uted, all vein orders raised, their prominence
Kostermans,1937: B-n.v.; isolectotypes: G-n.v., decreasing with rank; secondary veins 5-6 per side,
GZU-n.v.; fragments,F, NY; photos [neg. 3813 ex ? evenly spaced, slightly more distant aroundmid-
B], F, GH, NY). lamina, ascending at 50 60? (more obtuse around
Trees to 15 m. Branchletsslender, midway along midlamina),arcuate,distal pairs loop-connected;ter-
flush 3-5 mm diam., distally weakly angular, soon tiaries roughly horizontal, between secondaries
terete, pubescent, the surface concealed by the indu- forked. Staminateinflorescencesevenly spaced along
ment cover, the hairs relatively long, to 0.5 mm, current flush in the axils of foliage leaves or cata-
SYSTEMATICTREATMENT
phylls, to 15 cm long with 10 lateralbranches,branch
orders3-4, the highest orderdichasial, lax, the flow-
ers distant,the axes densely pubescent,the indument
as on branchlets;bracts and bracteoles caducous by
anthesis,lanceolate, sericeous;pedicels graduallyin-
creasing in diameterapically, to 1.3 mm long, those
supportingsecondary flowers slightly shorter.Flow-
ers rotate,rusty sericeous outside, the indumentthin-
ning distally; receptacle shallowly infundibuliform,
0.5 x 1 mm, densely grey-pilose inside. Tepalschar-
taceous, ovate, 1.3 x 0.8 mm, spreadingat anthesis,
the androeciumexserted, the outer surface sparsely
rusty sericeous, the innersurfacesparselygrey-pilose
near base, the margins and apex minutely papillose,
otherwise, glabrous. Stamens of whorls I and II
broadly stipitate, 1 mm tall, the anthersovate, 0.5 x
0.5 mm, glabrous,the apex roundedto truncate,the
connectives hardly prolonged between the 2 locelli,
these suborbicular, introrse-latrorse,the filaments
laminar,slightly narrowerthananthers,densely grey-
pilose; whorl III stamensbroadlystipitate, 1 mm tall,
the anthers oblong 0.5 x 0.4 mm, erect, locelli 2,
extrorse-latrorse,the filamentsslightly narrowerthan
anthers,laminar,densely grey-pilose,the basal glands
sessile, globose; whorl IV wanting; pistillode want-
ing. Pistillateinflorescencewith indument,color, and
branchingas in staminateplants, the flowers similar
in size and shape;stamens sterile, smaller;ovary gla-
brous; style stout, indistinct from ovary; stigma tri-
lobed, 0.3 mm diam. Fruitsbome on shortclaviform
pedicels of up to 5 x 3 mm; cupules shallowly in-
fundibuliform,0.3 X 1 cm, glabrousinside and out-
side, the marginssharplylobed, tepalbases persisting;
drupesellipsoid, to 1.5 X 1 cm.
Distribution (Fig. 39) and ecology. Small trees
of gallery forests apparentlyrestrictedto the cerrado
vegetation found in the states of Mato Grosso and
Goia's,Brazil, at ca. 400-500 m. Flowers collected in
May, June, and November,fruits from May to Octo-
ber.
Representative specimens examined. BRAZIL.
GoLas: Araguaina,500 ft, 11Aug 1963 (fr), Maguireet al.
56091 (HBG, MO, NY). MATO GROSSO:Barrado Gargas-
400 m, 15Jun
Rd.,77 kmfromBarrado Garqas,
Xavantina
1966 (fr), Hunt & Ramos 6004 (NY); ca. 270 km N of Xav-
antina, 4 May 1968 (fl d), Ratter et al. 1278 (E, K, MO,
NY, U); close to Xavantina-Sao Felix Rd., 6 Apr 1968 (fl
d), Ratteret al. 840 (E, K, NY);Corregodo Surucucu,10
Oct 1986 (fr), Sidney & Onishi 1308 (NY); Rio Araguaia,
Xavantina,400 m, 8 Jun 1966 (fr), Irwin et al. 16777 (HBG,
MG,MO,NY).
Local names. Brazil: louro amarello, louro dor-
ado, louro dourado,louro roxo.
109
Unlike most members of the Endlicheriasericea
species group,E. Ihotzkyimaintainsa vestitureof as-
cending rather than closely appressed hairs. These
hairs provide a shaggy aspect unmistakableamong
the smoothly sericeous branchletstypical of vegeta-
tively similar species. Hairs are also ascending in E.
ruforamulabut aremorereddishin color andless than
half the length found in E. lhotzkyi.Flowers of E.
lhotzkyiresemblethose of AntilleanE. sericea in be-
ing rotate with horizontallyspreadingtepals, but the
tepals are chartaceousratherthan fleshy with inner
surfaceindumentrestrictedto the base andconsisting
of straightratherthancrookedor crinkledhairs.Fruits
of E. lhotzkyiare conspicuousfor the shallowcupules
with strongly lobed margins. Of vegetatively similar
species, these cupules are matched only by the An-
dean E. aurea and densely sericeous forms of E. an-
omala, both with very different flowers. Goeppertia
chrysophylla Meisn., cited in synonymy by Mez
(1889) and Kostermans (1937), is an illegitimate
name based on a duplicateof the type materialof E.
lhotzkyidepositedin DeCandolle'sherbarium,now in
Geneva (G).
53. Endlicheria klugii 0. C. Schmidt,Repert.Spec.
Nov. Regni Veg. 31: 173. 1933. Type. Colombia.
Putumayo:Umbria,325 m, Dec 1930 (fl d), Klug
1904 (lectotype,designatedby Kostermans,1937:
NY; isolectotypes:A, B-n.v., F-n.v., GH, K-n.v.,
MO, S, US).
Treesto 15 m. Branchletsrelativelystout,midway
along flush 4-5 mm diam., angular,silvery to tawny
sericeous, the surface concealed by the indument
cover, the hairs short,to 0.15 mm, straight,appressed;
terminal buds relatively plump, 1 X 0.6 mm, rusty
sericeous. Leaves alternate,widely andevenly spaced
along currentflush; petioles long and slender,to 7 X
0.4 cm, semi-terete, the indument as on branchlets;
laminae chartaceous,plane, broadly ovate to elliptic,
12-35 X 6-18 cm, the base obtuse to rounded,rarely
acute, briefly decurrent,the apex broadlyacute, acu-
minatefor up to 6 cm, the marginsminutelyrecurved
throughout;upper surface dull slate grey to olive-
brown, minutely punctulate,the midrib and second-
aries prominent,the higher-ordervenation prominu-
lous; lower surfacesparselyor densely pubescent,the
hairs silvery to rustbrown, appressed,uniformlydis-
tributed,the epidermisclearly visible to obscured,all
vein ordersraised, their prominencedecreasingwith
rank;secondaryveins 4-6 per side, ? evenly spaced,
slightly more distantaroundmidlamina,ascendingat
50-60o (more obtuse aroundmidlamina),arcuate,the
110
lowermostpair occasionally asymmetric(one or both
merging with margin at base), distal pairs loop-
connected;tertiariesoblique to midrib, between sec-
ondaries straightto once-forked. Staminateinflores-
cences evenly spaced along currentflush in the axils
of foliage leaves, to 25 cm long with 9 lateral
branches,branchorders4-5, the highest orderdicha-
sial, lax, the flowers distant, the axes rusty to tawny
sericeous;bractsandbracteolescaducousby anthesis,
lanceolate, the indumentas on axes; pedicels terete,
to 2 mm long, those supportingsecondary flowers
slightly shorter. Flowers hypocrateriform,2.5 mm
diam., densely pubescentoutside,the hairsappressed,
reddish to brown; receptacle narrowly infundibuli-
form, 2 X 1 mm, grey-velutinousinside. Tepalschar-
taceous,ligulate, 1 X 0.6 mm (the innerwhorlslightly
narrower), spreading at anthesis, the inner surface
grey-tomentosenear base, the margins and apex in-
side sparsely papillose, otherwise glabrous.Stamens
of whorls I and II stipitate,0.6 mm tall, the anthers
narrowlyovate, 0.3 X 0.25 mm, glabrous,the apex
apiculate, the connectives prolonged between the 2
locelli, these suborbiculate,introrse-latrorse,the fila-
ments S-shaped,narrowerthananthers,densely grey-
tomentose;whorl III stamensbroadlystipitate,1 mm
tall, the anthers depressed-obovate,0.3 X 0.3 mm,
erect, locelli 2, extrorse-latrorse,the filamentsslightly
narrower than anthers, ligulate, densely grey-
tomentose, the basal glands sessile, minute, globose;
whorl IV wanting;pistillode fusiform.Pistillateinflo-
rescence with indument and color as in staminate
plants,but shorterandwith fewer lateralbranches,the
flowers slightly deeper; stamens sterile, smaller;
ovary glabrous,ellipsoid; style slender,distinctfrom
ovary, 0.6 mm long; stigma broadlytri-lobed to dis-
coid, 0.5 mm diam. Fruitsborne on stout, claviform
pedicels of up to 2 X 0.8 cm; cupules shallowly in-
fundibuliformto patelliform, to 0.4 X 1.3 cm, gla-
brous outside, sparselyrusty strigose inside, the mar-
gins undulate;drupesellipsoid to obovoid, to 3 X 1.4
cm.
Distribution (Fig. 39) and ecology. Small trees
from the nonfloodedforests of westernAmazoniaand
adjacenteasternAndeanfoothills at ca. 150-1300 m.
Flowering September to following March, fruits
availableyear round.
Representative specimens examined. COLOMBIA.
PUTUMAYO: Umbrfa,325 m, Dec 1930 (fl d), Klug 1884
(G, GH, K, MO, NY, S).
ECUADOR. NAPO: Estaci6n Experimental INIAP-
Napo, Payamino,Reserva Florfstica"El Chuncho,"250 m,
16-26 Feb 1986 (fl Y), Neill 7152 (MO, NY, US); Tena,
FLORA NEOTROPICA
Estaci6n Biol6gica JatunSacha, 400 m, 2 Jan 1998 (fl d),
Neill 11028 (MO).
PERU. AMAZONAS: Bagua, Imaza, Comunidadde Ya-
mayakat,320 m, 14 Feb 1996 (fl d), Jaramillo et al. 1159
(MO); Rfo Santiago, Caterpiza,180 m, Sep 1979 (fr), Hu-
ashikat 436 (HBG). HUANUCO:Pachitea, Pucallpa, Sira
Mtns., 680 m, 8 Apr 1988 (fr), Wallnofer111-8488 (MO).
LORETO: Alto Amazonas, Rio Huallaga, Shucushuyacu,
250 m, 12 Sep 1981 (fl d), Vdsquez& Jaramillo 2425 (F,
MO, NY); Maynas, Iquitos, Estaci6n ExperimentalIIAP,
Allpahuayo, 150 m, 24 Mar 1992 (fl d), Vdsquez et al.
18095 (MO). MADRE DE Dios: ManU,Atalaya,500-700 m,
8 Dec 1983 (fl Y, fr), Foster & Wachter7286 (G, MO).
PAsco: Oxapampa,Palcazu, 380 m, 24 Jan 1984 (fl Y),
Foster 9483 (G, MO, NY), 300-600 m, 13 Nov 1985 (fr),
Hartshornet al. 2824 (MO).
BRAZIL. AcRE: Bujari,FlorestaEstadualdo Antimari,
10 Mar 1997 (fr), Daly et al. 9425 (MO). AMAZONAS:Rio
Ia, 24 Feb 1977 (fl d), Mori et al. 9097 (F, HBG, INPA,
MO, NY, U, US); Rio Javari,behindPalemeirasArmy Post,
31 Jul 1973 (fr), Lleras et al. P16960 (HBG, K, MO, NY,
R, US).
Local name. Peru:yuwich (Huambisa).
The S-curved whorl I and II filaments of Endli-
cheria klugii are unique in the genus. These curious
filamentscombined with the ovate anthersconfer on
stamens an aspect that resembles a cobra raised in
strike position (Fig. IF). The flowers are themselves
remarkablefor their slender receptacles below hori-
zontally spreadingtepals.
The long petioles supportingbroadlyovate leaves
with arcuatesecondaries are sometimes matchedby
E. ruforamula,from which E. klugii can be distin-
guished, even without flowers, by its closely ap-
pressed indumenton branchletsand shallow cupules
with clavate pedicels.
54. Endlicheria argentea Chanderbali, sp. nov.
Type. Peru. Loreto: Requena, Sapuena,Arboreto
JenaroHerrera,130 m, 14 Jul 1989 (fl 6), Vdsquez
& Jong 12397 (holotype: MO; isotypes: AMAZ,
HBG, MO). Fig. 40
InterEndlicheriaesericeae et specierumaffiniumflori-
bus infundibularibusad E. klugii accedens sed foliis supra
costa impressiset staminibusrectis differt.
Trees to 5 m. Branchlets slender, midway along
flush 2 mm diam., distally weakly angular,soon te-
rete, sericeous, the surfaceconcealedby the indument
cover, the hairs short,to 0.3 mm, straight,appressed,
silvery to straw-colored,shining;terminalbuds slen-
der, 2 X 0.6 mm, sericeous. Leaves altemate,widely
and evenly spaced along currentflush; petioles slen-
 -5

1mm~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
FIG. 40. Endlicheria argentea (Vdsquez & Jong 12397). A. Habit. B. Single leaf showing venation. C. Leaf base
below. D. Rlower.E. Rlower with tepals removed. F. Whorl I stamen seen from within. G. Whorl III stamen seen from
without.
112 FLORA NEOTROPICA

der, to 1.5 X 0.2 cm, terete, the indument as on silvery sericeous ratherthan reddish or brown and
branchlets; laminae chartaceous, plane, narrowly filamentsof whorl I and II stamensare straightrather
ovate to lanceolate, 14-17 X 3-5 cm, the base acute, than curved.Endlicheriaargentea also appearsto be
briefly decurrent,the apex acute, acuminatefor up to a less robustspecies with slenderbranchletsand pet-
4 cm, the marginsflatthroughout;uppersurfaceolive- ioles less thanhalf the dimensionsfound in E. klugii.
brown, minutely punctulate, the midrib impressed, Urrego et al. 1218 (MO), from Rio Caquetaiin
the higher-orderveins prominulous;lower surfacese- Amazonian Colombia, is an exact vegetative match
riceous, the hairs silvery, uniformlydistributed,con- and likely belongs here but its sterile condition pre-
cealing the epidermis, all vein orders raised, their vents definite identification.
prominencedecreasingwith rank;secondaryveins 3-
4 per side, closer near base and apex, widely spaced
aroundmidlamina,ascendingat 50-60? (moreacutely 55. Endlicheria robusta (A. C. Sm.) Kosterm.,Re-
towards apex), arcuate, distal pairs loop-connected; cueil Trav.Bot. Neerl. 34: 556. Cryptocaryaro-
tertiaries roughly horizontal, between secondaries busta A. C. Sm., Bull. TorreyBot. Club 58: 97.
once-forked to straight. Staminate inflorescences 1931. Type. Peru. Junin: San Nicholas, Pichis
evenly spaced along currentflush in the axils of fo- Trail, 1100 m, 4-5 Jul 1929 (frjuv), Killip & Smith
liage leaves, to 10 cm long with 8 lateral branches, 26077 (holotype:NY; isotypes: F, US).
branchorders2-3, the highest orderdichasial,lax, the
flowers distant,the axes silvery sericeous;bractsand Trees to 20 m. Branchletsrelativelyslender,mid-
bracteolescaducousby anthesis,narrowlylanceolate, way along flush 2-3 mm diam., weakly angular,
sericeous; pedicels terete, to 2 mm long, those sup- densely strigillose, the surface barely visible, dark
porting secondary flowers slightly shorter.Flowers brown, the hairs very short, to 0.1 mm, appressed,
tubiform,to 5 mm diam., sericeous outside; recepta- tawny or light brown;terminalbuds slender,2 X 0.2
cle infundibuliforn,2 X 1.5 mm, densely silvery pi- mm, reddish sericeous. Leaves alternate,widely and
lose inside, tepals chartaceous,ovate, 1 X 0.6 mm evenly spaced along currentflush;petioles slender,to
(the inner whorl slightly narrower),ascending at an- 2 X 0.2 cm, semi-tereteto broadly canaliculate,the
thesis, surrounding androecium, the inner surface indument as on branchlets; laminae chartaceous,
densely pubescent,the hairs as in receptacle,ascend- plane, ovate to elliptic, 8-20 X 3-10 cm, the base
ing to erect, the margins and apex inside papillose. acute,briefly decurrent,the apex acute, acuminatefor
Stamens of whorls I and II stipitate, 1 mm tall, the up to 3 cm, the margins flat or weakly recurvedin
anthersbroadly ovate, 0.3 X 0.5 mm, glabrous,the lower half; upper surface olive-brownto light green,
apex apiculate, the connectives prolonged between minutelypunctulate,the primaryto fourth-orderveins
the 2 locelli, these obliquely hemispherical,introrse,raised, their prominencedecreasingwith rank;lower
the filamentslaminar,narrowerthan anthers,densely surface sericeous to sparsely strigillose, the hairs
tawny to light brown,dull to lustrous,uniformlydis-
silvery pilose, the hairs as in receptacle;whorlIII sta-
mens sessile, 1 tall, the anthers depressed-obovate, tributedon lamina, sparseron midrib and secondar-
0.3 X 0.3 mm, erect, locelli 2, extrorse-latrorse,the ies, all vein ordersraised,theirprominencedecreasing
filaments as broad as anthers,laminar,the indument with rank, the midrib often darkerand crooked after
midcourse;secondary veins 5-7 per side, + evenly
as in outerwhorls, the basal glands sessile, ovate, the
apex apiculate;whorl IV wanting;pistillode filiform. spaced, slightly more distant aroundmidlamina,as-
Pistillate plants unknown. cending at 50-60O (more obtuse towards apex), ar-
cuate, distal pairs loop-connected;tertiariesoblique
Distribution (Fig. 41) and ecology. Known only to midrib, between secondaries straight or forked.
from the type (but see discussion), a small plant of Staminateinflorescencesevenly spaced along current
seasonally inundatedlowland (ca. 130 m) forest in flush in the axils of foliage leaves, to 10 cm long with
Peru, found flowering in July. 8 lateralbranches,branchorders2-3, the highest or-
der dichasial, lax, the flowers distant,the axes grey-
Endlicheria argentea differs from similarly seri- strigillose, distalmost branches and pedicels more
ceous species by its tubiformflowerswith tall slender densely so; bracts and bracteolespersistentat anthe-
stamens,but the impressedmidrib above is also dis- sis, lanceolate, silvery sericeous; pedicels terete,to 3
tinctive. Perhapsits closest relationshiplies with E. mm long, those supportingsecondaryflowersslightly
klugii, with which it sharesnarrowlyinfundibuliform shorter.Flowers obconical, or urceolate,2 mm diam.,
receptacles and depressed-obovatewhorl III anthers. sparsely grey-strigillose outside; receptacle deeply
However, the flowers of E. argentea are externally cyathiform,the base abruptlytruncate,1.3 x 2 mm,
SYSTEMATICTREATMENT
L /~~~~~~~~~~~~~~~~~~~~~~~
.Api ' ' '';
F Db o c
FIG. 41. Distnbution of Endllicheriaargentea and E. robusta.
densely grey-pilose inside. Tepals chartaceous,
broadly ovate, 0.6 x 1 mm (the inner whorl slightly
narrower),inflexed to erect at anthesis, the androe-
cium included, the inner surface sparsely grey-
strigillose, the marginsand apex inside sparselypap-
illose. Stamens of whorls I and II stipitate, 0.6 mm
tall, broadly anthersovate, 0.5 X 0.5 mm, glabrous,
the apex sharply apiculate, the connectives narrowly
prolongedbetween the 2 locelli, these suborbiculate,
introrse-latrorse,the filaments laminar, slightly nar-
rowerthananthers,sparselygrey-tomentellose;whorl
III stamens sessile, 0.7 mm tall, the anthersnarrowly
oblong, 0.4 X 0.2 mm, erect, locelli 2, extrorse-
latrorse, the filaments broader than anthers, the in-
dument as in outer whorls, the basal glands sessile,
globose; whorl IV wanting; pistillode fusiform.
Pistillate inflorescencewith indumentand color as in
staminateplants, but shorter and with fewer lateral
branches,the flowers similar in size and shape; sta-
mens sterile, smaller; ovary glabrous, ovoid; style
slender, distinct from ovary; stigma minutely tri-
113
r a r
lobed, 0.5 mm diam. Fruits borne on claviform ped-
icels of up to 1 X 0.3 cm; cupules shallowly hemi-
spherical,to 0.5 X 1 cm, glabrousinside and outside,
the marginsentire;drupesovoid, to 1 X 0.8 cm.
Distribution (Fig. 41) and ecology. Small to
medium-sized trees from western Amazonia and ad-
jacent lower montane slopes of the eastern Andes at
ca. 100-1800 m. Flowering specimens collected from
MarchthroughJuly and from NovemberthroughDe-
cember, fruits from April throughJuly and from Oc-
tober throughJanuary.If not a sampling artifact,col-
lections of this species suggest two discrete
reproductivecycles per year, with an early flowering
period from Marchto July providingfruits from Oc-
tober to January,and a late flowering period in No-
vember and December responsiblefor fruit availabil-
ity from April to July.
Representative specimens examined. ECUADOR.
NAPO:Huaorani,Rio Shiripuno,Quehueiri-ono,300 m, 14
May 1995 (fl 6), Miller et al. 638 (MO). PASTAZA:Auca,
114
Rio Tiguino, 115 km S de Coca, 320 m, 29 Apr 1989 (fl d),
Rubio 12 (MO); Lorocachi, 200 m, 25 May 1980 (fl d),
Brandbyge & Asanza 30885 (AAU). SUCUMBiOS: Lago
Agrio, Reserva Faunistica Cuyabeno, 265 m, 11 Mar-13
May 1990 (fl d3), Balslev et al. 97016 (MO). ZAMORA-
CHINCHIPE: Nangaritza,faldas de las Cordilleradel C6n-
dor, 1600-1800 m, 5 Dec 1990 (fl d), Palacios & Neill 6522
& 6531 (MO).
PERU. AMAZONAS: Condorcanqui,Cenepa, Comuni-
dad de Tutino, 500 m, 18 Jul 1997 (fl Y), Vdsquezet al.
24341 (MO), 24 Jul 1997 (fr), Rojas et al. 172 (MO). JUNiN:
San Nicolis, Pichis Trail, 1100 m, 4-5 Jul 1929 (fr), Killip
& Smith25975 (NY). LORETO:Maynas, Iquitos, carretera
Iquitos-Nauta,km 44, 150 m, 14 Dec 1988 (fr), Vasquez&
Jaramillo 11442 (GH, HBG, MO); Rio Napo, San Pedro,
Isla de Mangua, below caserfo JuanchoPlaya, 6 Apr 1979
(fl d), Rimachi4397 (MO). PASCO:PEPP ForestReserve,
PuertoMairo,300-600 m, 5 Jun 1986 (fl J, juv), Hartshorn
al.
et 2971 (MO); SantiagoSoto, Buenos Aires, 300-600 m,
18 May 1986 (fl 6, juv), Hartshorn& Quijano2940 (MO).
BRAZIL. AcRE: Cruzeiro do Sul, Rio Jurua & Rio
Moa, Serrade Moa village, 22 Apr 1971 (fr), Prance et al.
12233 (HBG, MG, MO, NY, U). AMAZONAS: Labrea,sub-
base do Projeto RADAM/BRAZIL,SG-20-YA, Ponto 02,
20 Jun 1976 (fl 6), Mota s.n. INPA 60364 (INPA); Vila
Bittencourt,Rio Japura,Serrinha,16 Nov 1982 (fr), Amaral
et al. 488 (INPA, MG, MO).
Local names. Peru:moenablanca,moenilla,tikis,
tindu, tunchi, tinchi.
Endlicheria robusta is alone among species with
dense sericeous indumentin having obconical or ur-
ceolate flowers with incurvedor erect tepals thatpro-
vide only a narrowterminalpore at anthesis. In the
E. browniana species group, E. formosa and E. par-
adoxa have similar flowers well as as appressed in-
dument; but although E. robusta can be subsericeous,
its leaves are never obovate as in E. formosa, and the
tertiariesnever reticulateas in E. paradoxa. Similar
flowerscombine with rustytomentosevegetativeves-
titurein E. szyszylowiczii.
A few specimens from Jenaro Herrera are placed
here with hesitation. In Spichiger & Loizeau 4161 and
4108 (A) and Valcarcel & Chota 6/619 (MO), sta-
minate flowers are more densely pubescent than ever
encounteredin E. robusta,and the fruitingspecimen,
Encarnaci6n 26150 (G, MO, US), shows rather small
shallow cupules with spheroiddrupes.Such fruitsare
also found in Gentry et al. 37246 (F, G, MO) from
Junfn, Peru, and Berg & Steward P19913 (HBG, K,
MO, NY, US) from Mato Grosso, Brazil, extending
considerably the geographic range of this variant.
Similar cupules and densely pubescent flowers are
found in E. tschudyanabut the shape of the flowers
and sharplyapiculatewhorl I and II antherspoint to
E. robusta.
FLORA NEOTROPICA
56. Endlicheria paniculata (Spreng.) J. F. Macbr.,
Publ. Field Mus. Nat. Hist., Bot. Ser. 13 (2, 3):
850. 1938. Citrosmapaniculata Spreng.Syst. Veg.
2: 545. 1825. Type. Brazil. Withoutlocality, Sel-
low s.n. (holotype: B [fide Kostermans, 1937]-
n.v.).
Cryptocaryahirsuta Schott, Syst. Veg. 4(2): 405. 1827.
Endlicheria hirsuta (Schott) Nees, Linnaea 8: 38.
1833. Goeppertiahirsuta (Schott) Nees, Syst. laur.
366. 1836. Type. Brazil. Rio de Janeiro:San Cris-
tovao, Pohl 5611 (lectotype, designated by Koster-
mans, 1937: W-n.v.; isolectotype:U).
Nectandralucida Nees, Linnaea8: 47. 1833. Type. Bra-
zil. Without locality, Sellow s.n. (holotype: B [fide
Kostermans,1937]-n.v.; fragment,GZU [fide Roh-
wer, 1993a]-n.v.).
Goeppertia longifolia Nees, Syst. laur. 368. 1836. En-
dlicheria longifolia (Nees) Mez, Jahrb.Konigl. Bot.
Gart.Berlin 5: 119. 1889. Type. Peru.Cuchero,Nov
1821 (fl 5), Poeppig 1520 (syntypes: B-n.v., BM-
n.v., G-n.v., GOET-n.v., KIEL-n.v., LE-n.v., LZ-
n.v., NY-n.v., OXF-n.v., P-n.v., W-n.v.; fragment,
U).
Goeppertiapanicularis Nees, Syst. laur.368. 1836. En-
dlicheria panicularis (Nees) Mez, Jahrb. Konigl.
Bot. Gart.Berlin 5: 128. 1889. Type. Brazil. Rio de
Janeiro:Withoutlocality,Oct-Dec (fl 5), Lhotzky43
(holotype:B [fide Kostermans,1937]-n.v.).
GoeppertiacantagallanaMeisn., DC. Prodr.15(1): 173.
1864. Type.Brazil. Minas Gerais:CantaGallo, 1859
(fl 5), Peckholt115 (holotype:M-n.v.; isotype: U).
Goeppertiahirsuta (Schott) Nees, var.coriacea Meisn.,
DC. Prodr.15(1): 172. 1864. Syntypes.Brazil.Minas
Gerais, Sellow s.n. (G-n.v., M-n.v.); Weddells.n.
(G-n.v., M-n.v.); Claussen 454 (G-n.v., M-n.v.);
Aug 1840 (fl juv), Claussen 2454 [cited as 1454 by
Meissner] (G-n.v., K, M-n.v.).
Goeppertiahirsuta (Schott) Nees, var.hirsutiorMeisn.,
DC. Prodr. 15(1): 172. 1864. Endlicheriapoeppigii
Kosterm.,Recueil Trav.Bot. Neerl. 34: 555. 1937.
Type. Peru. Loreto:Maynas,Yurimaguas,Jun 1831
(fl 5), Poeppig2298 (holotype:W-n.v.; isotypes:B-
n.v., G-n.v., LE, LZ-n.v., OXF-n.v., U).
Goeppertiahirsuta (Schott) Nees, var. latifolia Meisn.,
DC. Prodr.15(1): 172. 1864. Type. Brazil. Near Vi-
t6ria, withoutdate (fl 5), Sellow 433 (holotype:B-
n.v.; isotype: K).
Endlicheriahirsuta(Schott)Nees, var.glabrataGlaziou,
Bull. Soc. Bot. France 12 (3): 590. 1912. Syntypes.
Brazil.Rio de Janeiro:TijucaandCorcovado,19 Oct
1868 (fl 5), Glaziou 3092 (B-n.v., K-n.v., P); road
to Macaco, nearVista Chineza, 26 Dec 1886 (fl 5),
Glaziou 16315 (B-n.v., K-n.v., P).
Endlicheriahirsuta (Schott) Nees, var. robustaGlaziou,
Bull. Soc. Bot. France 12(3): 590. 1912. Syntypes.
Brazil. Minas Gerais: Biribiry, 28 Mar 1892 (fr),
Glaziou 19795 (B-n.v., K-n.v., P)
SYSTEMATICTREATMENT 115

Endlicheria boliviensis Kosterm., Recueil Trav. Bot.

grey-pilose;whorl III stamensbroadlystipitate,1 mm
Neerl. 34: 553. 1937. Type. Bolivia. Santa Cruz:tall, the anthersoblong, 0.5 X 0.4 mm, erect, locelli
Buena Vista, 500 m, 14 Mar 1925 (fl d), Steinbach
2, extrorse-latrorse,the filamentsequal to or slightly
6985 (holotype: B-n.v.; isotypes: GH, K, MO, NY,narrowerthan anthers, ligulate, the indument as in
S, U).
outer whorls, the basal glands sessile, globose; whorl
Endlicheriaracemosa Lasser,Bol. Tecn. Minist. Agric.
3: 8. 1942. Type.Venezuela.Carabobo:Hospitalnear
IV wanting; pistillode fusiform. Pistillate inflores-
San Joaquin, 1000 m, 15 Aug 1918 (fl d), Pittiercence with indumentand color as in staminateplants,
8017 (holotype:VEN-n.v.; isotypes: GH, MO). but shorterand with fewer lateralbranches,the flow-
ers similarin size and shape;stamenssterile, smaller;
Trees to 10 m (rarely20 m). Branchletsslenderto ovary glabrous, ovoid; style stout, indistinct from
stout, midway along flush 3-6 mm diam., angular, ovary; stigma broadlytri-lobed,0.6 mm diam. Fruits
densely pubescent,the surfacebarely visible to con- borne on claviform pedicels of up to 2 X 0.5 cm;
cealed by the indument cover, the hairs relatively cupules hemispherical,to 1 X 1.5 cm, glabrousout-
short to long, (0.3-)0.6(-1.5) mm, straight,erect to side, strigose inside, the marginsentire,or tepalbases
appressed,rusty red to grey; terminalbuds plump, 3 persisting;drupesovoid, to 2.5 x 1.2 cm.
X 2 mm, densely pubescent,the hairs as on branch-
lets, ascending. Leaves alternate,widely and evenly Distribution (Fig. 42) and ecology. Small to
spaced along currentflush; petioles slender, to 2 X medium-sized trees distributed from the Atlantic
0.3 cm, terete,the indumentas on branchlets;laminae coastal forests of SE Brazil, throughthe lower slopes
chartaceousto membranaceous,plane, ovate to obo- of the Andes in tropicalSouth America to Panamain
vate, 7-30 X 2-12 cm, the base acute to obtuse, the CentralAmerica.Occurringat 50-1000 m throughout
apex obtuse to acute, acuminatefor up to 1.5 cm, the its geographicrangeand reachingover 2000 m in the
marginsminutelyrecurvedthroughout;uppersurface Andes. Flowers and fruits available throughoutthe
reddishto olive-brown,waxy, the midriband second- year.
aries sunken,the higher-ordervenation raised or im-
mersed;lower surfacedensely pubescent,the hairsas Representative specimens examined. PANAMA.
PANAMA: El Llano-CartiRd., 8-11 km from Inter-
on branchlets, uniformly distributed or denser on
AmericanHwy., 300-400 m, 13 Aug 1975 (fl d), Mori 7708
main veins, all vein orders raised, their prominence
(MO);Serranfa de Maje,5-6 hrswalkfromChoc6Village,
decreasing with rank;secondary veins 4-6 per side, 650-800m, 31 Mar1982(fr), Knappet al. 4496 (MO).SAN
? evenly spaced, slightly more distant aroundmid- BLAS: El Llano-Carti Rd.,Pacificside,350m, 13Feb1983
lamina, ascending at 50-60O (more obtuse around (fr), Hamilton & Stockwell2905 (MO); Pemasky,carretera
midlamina),arcuate,distal pairs loop-connected;ter- Nusagandi-Cartf, 5 Jun1994(fl 9, juv), Galdameset al.
tiaries roughly horizontal, between secondaries 1362 (MO).
straight to forked. Staminate inflorescences evenly COLOMBIA. CUNDINAMARCA:Laguna dePedroPalo,
spaced along current flush in the axils of foliage 4 kmfromrd.Bogotato La Mesa,2070 m, Oct 1990(fr),
leaves, to 20 cm long with 14 lateralbranches,branch Wijninga580 (U). HUILA: RioNegro,30 kmESEof Baraya,
orders3-4, the highest orderdichasial, lax, the flow- ca. 2300 m, Oct 1944(fl 9), Little8878 (A), 30 Oct 1944
(fl 9), Little 8886 (A). MAGDALENA: SierraNevadade
ers distant,the axes densely pubescent,the indument
SantaMarta,1700-1900m, 5 Oct1972(fr),Kirkbride 2387
as on branchlets;bracts and bracteoles caducous by (US). META: La Macarena, ParqueNacionalNaturalTini-
anthesis,rarelypersisting,ovateto lanceolate,densely guas,s.d. (fr),Polanco222 (MO).
pubescent, the hairs as on axes, ascending;pedicels VENEZUELA. LARA: DistritoMoran,Quebrada Los
terete, to 4 mm long, those supporting secondary Cedros, 1500 m, 8 Jul 1974 (fl d), Steyermark& Espinoza
flowers slightly shorter. Flowers rotate, 2-4 mm 110275(MO,NY, VEN);DistritoYaritagua, nearborder
diam., sparsely to densely silvery or rusty strigose withDistritoUrachiche of EstadoYaracuy, 1450m, 28 Mar
outside; receptacle infundibuliform,0.5 X 1 mm, 1975(fl 6), Steyermarket al. 111746 (F,G,NY).TACHIRA:
densely rusty or grey-pilose inside. Tepals charta- MataMula,N of Deliciason rd.to Bramon,1750m, 26 Jul
& Liesner 118687 (MO). YARA-
ceous to membranaceous,ovate to ligulate, 1.5 X 0.7 1979 (fl d), Steyermark
cuy: Cocorote, carreteraCocorote-Fila Las Cumaraguas,
mm, spreadingat anthesis, the inner surfacedensely
1500-1600 m, 13 May 1994 (fr), Benftez de Rojas et al.
silvery grey-strigose,the marginsminutelypapillose. 5096 (MO); Nirgua, al NW de Montalban,entre torres de
Stamens of whorls I and II stipitate, 1 mm tall, the relevo (TV), y Cumbrede Capotillo, 1350-1500 m, 19 Mar
anthersovate, 0.6 x 0.4 mm, glabrous, the connec- 1999 (fr), Meier & Kunert4632 (MO).
tives broadabove, reducedbetween, or homlike over, ECUADOR. NAPO: Estaci6n Experimental INIAP-
the 2 locelli, these suborbicular,introrse-latrorse,the Payamino,250 m, 18-26 Feb 1986 (fr), Palacios et al. 1030
filaments laminar, narrower than anthers, sparsely (MO);RioWai-si-aya, 300 m, 28 Aug 1981(fl d), Brand-
116 FLORA NEOTROPICA

|._-................ .....; -- -- -- - - ------- ---

1........ .- .------ M.,4 , -,P{

I f-ioB''!?t-i 4;257.

...................... --- ---e- --:-- --- i-

* f

FIG.
42. Distributionof Endlicheria paniculata.

byge et al. 36206 (AAU). SucuMBios: San Pablo de Kan- cerca a Rio Chino, 1400-1600 m, Jun 1976 (fr), Schunke
tesiya, Rfo Aguarico, 250 m, 23 May 1992 (fl d), Pfrommer 9242 (MO); Bosque Nacional Alexandervon Humboldt;km
6 (MO). 86 Pucallpa-Tingo Maria Rd., 270 m, Jan 1978 (fl d),
PERU. AMAZONAS: Above mouth of QuebradaCikan- Froehner 171 (HBG, MO).
inci, N of Rio Cenepa, 30 Dec 1972 (fl S), Berlin 753 (F, BRAZIL. BAHIA:Rod. PortoSeguro-Eunapolis,km 31,
MO); Vista Alegre, puente sobre el Rio Salas, 1525 m, 30 26 Nov 1970 (fl 6), Filho & Emmerich2934 (R). DisTiuTo
Jun 1998 (fl d), Sdnchez et al. 9575 (MO). HuANUco: FEDERAL: Brasflia, bacia do Rio SaioBartolomeu, 24 Jan
Pachitea,Honoria, Bosque Nacional de Iparia,300-400 m, 1980 (fl 6), Heringer et al. 3187 (MO); C6frego Quilombo,
30 Jan 1967 (fl &), Schunke 1587 (GH, INPA, K); Puerto 18 Sep 1980 (fl 6), Heringer et al. 5513 (MO). EsPiRfRo
Inca, Llullapichis,270 m, 1 Dec 1989 (fl juv), Kroll Saldania SANTO: Concei,caodo Castelo, Rod. BR-262, Dec 1984 (fl
795 (G, MO). LORETO:Alto Amazonas, above Pongo de dc),Hatschbach& Silva 48651 (MO); SantaTeresa,Reserva
Manseriche, Rio Santiago, 200 m, 23 Dec 1931 (fl &), Biologica de Nova Lombardia,700-750 m, 4 Feb 1985 (fl
Mexia 6329 (F, GH, K, MO, NY); Yurimaguas,140 m, 7 Jan d), Peixoto et aL 3466 (MO). GoIAs: Serrado Caiap6, ca.
1933 (fl J), Klug 2833 (GH, K, MO). MADRE DE Dios: 12 km S of Caiap6nia,840 m, 2 May 1973 (fl d), Anderson
Mand, ParqueNacional Mand, Rfo Manu, 350 m, 10 Sep 9604 (NY); Campinacu,400 m, 10 Oct 1991 (fr), Cavalcanti
1986 (fr), Foster 11339 (INPA, NY); Tambopata,Comuni- et al. 947 (MO). MATOGRosso: Caba.do Rio Taquaraussu
dad Nativa de Infierno,260 m, 17 Jul 1989 (fr), Alexiades (Chapada),Mar 1911 (fl Y), Hoehne 3517 (R); SantaAnna
& Diaz 849 (MO). PASCO:Oxapampa,Chantabamba,1890 da Chapada, Aug 1902 (fr), Robert 4856 (K). MATO
m, 28 May 1982 (fl f), Smith et al. 1748 (MO); Palmazu, GRosso Do SuL: Ponta Pora, Ponto Alto, 12 Feb 1983 (fl
2200 m, 2 Oct 1984 (fl 9), Smith et al 8670 (MO). SAN c), Hatschbach 46151 (MO); Rod. BR-163, 1 km L de
MARTiN: Mariscal Caceres, Tocache Nuevo, Quebradade Mundo Novo, 7 Nov 1993 (fl d), Hatschbach et al. 58562
Canuto, 250 m, 2 Jan 1979 (fl 9), Schunke 10657 (MO). MINASGERAIS: Serrodo Espinhaqo,1100 m, 12 Feb
(MO).UCAYALI: CoronelPortillo,PadreAbad, La Divisoria 1969 (fl Y), Irwin et al. 23151 (MO); Rio Acima, Feb 1994
SYSTEMATICTREATMENT
(fl 6), Costa s.n. BHCB26334 (MO). PARANA: CerroAzul,
Caeceirado Ribeiraodo Tigre, 14 Apr 1987 (fl 6), Hatsch-
bach et al. 51226 (MO); Palotina,Perola Independente,13
Mar 1985 (fl Y), Hatschbach& Silva 48978 (MO). Rio DE
JANEIRO:Angra dos Reis, ReservaBiologica Est. da Praia
do Sul, 2 Mar 1985 (fl 6), Araujo & de Oliveira601 (MO);
SantaMariaMadalena,PedraDubois, 900-1195 m, 22 Feb
1983 (fl 9), Plowman & de Lima 12895 (K, MO). Rio
GRANDEDO SUL: Itacolumi, prope Gravataf,11 Jan 1950
(fl 6), Rambo45269 (K, MO); PortoAlegre, Morroda Pol-
icia (Caseata),9 Dec 1901 (fl 6), Malme 755 (R). SANTA
CATARINA: Ararangua,Passo do Sertao, 10 m, Feb 1952 (fl
6), Reitz 4423 (MO); Horto Florestal I.N.P., Ihirama,350
m, 2 Nov 1953 (fr), Reitz & Klein 1140 (MO). SAOPAULO:
ParaguacuPaulista, Fazenda Sao Jose, 425-450 m, 8 Feb
1865 (fl 9), Eiten et al. 5899 (K, MO); Morro das Pedras,
Iguape, Dec 1922 (fl 6), Brade 8195 (R).
BOLIVIA. BENI:Prov.Ballivian,Serraniadel Pil6n La-
jas, 850-900 m, 19-26 Feb 1990 (fl 9), Smithet al. 14052
(MO), 16 Feb 1992 (fl 9), Killeen & Smith3634 (MO). LA
PAZ: Prov. Itturalde,Buena-VistaTacana, 17 Feb 1997 (fl
6), Serato 305 (MO); Prov. Murillo, Zongo valley, Cahua
hydroelectricplant, 1200-1400 m, 23 Dec 1984 (fl 9), Sol-
omon 12948 (MO). PANDO:Puerto Nuevo, Rio Ortan,Feb
1924 (fr), Meyer 146 (MO). SANTACRUZ:Prov. Ichilo,
Buena Vista, 300 m, 22 Mar 1995 (fl 9), Abbott & Isaacs
16484 (MO);Prov.Sara,Rio Surutd,400 m, 2 Oct 1925 (fr),
Steinbach 7264 (GH, K, MO, U).
PARAGUAY.ALTOPARANA:Centro Forestalalto Pa-
rand, 12 km 0 de Puerto PresidenteStrossner,1 Feb 1984
(fl 6), Little 40105 (MO); Reserva Biol6gica Itabo, 9 Oct
1990 (fr), Schinini & Marmori 27009 (MO). AMAMBAY:
Parque Nacional Cerro Cord, 300 m, 20 Feb 1982 (fl 6),
Solomon et al. 7140 (MO); Torin, Aug 1985 (fr), Hatsch-
bach & Cervi 48903 (MO). CAAGUAZU: S.loc., 12 Nov
1874 (fr), Balansa 2026 (K); s.loc., 5 Feb 1905 (fl 6), Hass-
ler 8935 (K). CAAZAPA: Linea del Bosque Parquehacia Ao
Ita y Ao Jakuy,20 Jul 1986 (fr), Molas 771 (MO); Tavai,
Prop. Trosiuk, 17 Mar 1989 (fl 6, juv), Soria 3419 (MO).
CANENDIYUJ: MbaracayuNaturalReserve, 13 Jan 1998 (fl
9), Zardini & Guerrero47737 (MO); Lagunita, Sendero
Arroyo Moroti, 13 Mar 1997 (fl 9), Jimenez et al. 1809
(MO). SANPEDRO:Colonia 8 de Diciembre, 12 km al SE
de Chore, 30 Sep 1987 (fr), Zardini & Ben(tez3182 (MO);
Colonia Naciente, 8 km al SE de Chore, 2 Oct 1987 (fr),
Zardini& Benftez3429 (MO).
Local names. Ecuador: shiringochy (Siona). Peru:
moena rosada, moenilla, roble, sacha muena, tinchi.
Brazil: canela amarella, canela frade, canela garuva,
canela paluda. Bolivia: laurel. Paraguay: laurel agua-
cate, laurel moroti.
As circumscribed here, Endlicheria paniculata ac-
commodates considerable variation in leaf size and
shape, but more disturbing to any sense of intraspe-
cific uniformity are the different manifestations of
vestiture. The type material and most collections of
117
this species arefrom the Atlanticcoastalforestsof SE
Brazil where one usually finds rusty tomentose in-
dumentcovering the lower surfacesof narrowlyovate
leaves. However, this indument can appear with
broadly ovate (e.g., Araujo 6776), or even obovate
leaves (e.g., Heringeret al. 3187). Similarlyirrespec-
tive of leaf size and shape, a subsericeousvestitureof
pale appressedhairs, e.g., Sellow 433 (type of E. hir-
suta var. latifolia) and Glaziou 3092 (type of E. hir-
suta var.glabrata), or a hirsutecover of longer,more
stiffly erect, dark red hairs (e.g., Rambo 45269 and
Brade 8195), are both found in SE Brazil.
These vegetative instabilities are repeated
throughoutthe range of E. paniculata. The typical
form, with rusty tomentose ovate leaves, is wide-
spread,ranging from SE Brazil throughlower mon-
tane Andean slopes from Bolivia (e.g., Solomon
12948) to Venezuela (e.g., Steyermark& Espinoza
110275). The othervariantsarealso widespread.Sub-
sericeous indument reappearswith ovate leaves in
NW South America, including the type of E. race-
mosa, andrangesinto Panama,while in westernAma-
zonia it combines with obovate leaves in the type of
E. longifolia. Likewise, hirsuteindumentreappearsin
western Amazonia with either ovate (e.g., Schunke
1617) or obovate leaves (e.g., Poeppig 2298, the type
of E. poeppigii). However,as in SE Brazil,tomentose
forms that are otherwiseindistinguishablefrom these
subsericeousandhirsuteforms arefoundin sympatry.
Flowers also vary in E. paniculata, but only slightly
and not in correlationwith indument.All are rotate
with horizontally spreadingtepals, stipitate whorl I
andII stamens,andbroadlystipitatewhorlIIIstamens
with relatively large globose glands, but the anther
apices in whorl I and II may be truncateor emargi-
nate. The latterconditionmay combine with hornlike
lobes persisting above each locule, as was indicated
as diagnostic for E. boliviensis (Kostermans,1937),
and appearsthroughoutthe rangeof E. paniculata. In
Mexia 6329, staminateflowersareprovidedwith four
whorlsof stamens(i.e., whorlIV membersarefertile).
This remarkablesituationappearsto be an abnormal-
ity never repeated in the species, nor elsewhere in
Endlicheria.
Despite the internalvariation,E. paniculatais eas-
ily recognizedby its pinnatevenationsince otherspe-
cies with rotate flowers and stipitate stamens with
truncateantherapices, E. acuminataand E. gracilis,
have triplinervedleaves.
57. Endlicheria gracilis Kosterm., Recueil Trav.
Bot. NMerl.34: 528. 1937. Type. Colombia. San-
118 FLORA NEOTROPICA

tander:Vicinity of BarrancaBermeja,Magdalena mm tall, the anthersdepressed-oblong,0.3 X 0.5 mm,

Valley, between Sogamoso and Colorado Rivers, erect, locelli 2, extrorse-latrorse,the filamentsslightly
100-500 m, 14 Nov 1934 (fl Y), Haught 1414 narrower than anthers, clavate, densely grey-
(holotype: U; isotypes: A, G). strigillose inside, the basal glands sessile, globose,
Endlicheriagrisea Kosterm.,Reinwardtia6: 285. 1962.
relatively large, filling the space between filamentsof
Type. Guyana.Essequibo:Wabuwak,KanukuMtns., the inner and outer whorls; whorl IV wanting;pistil-
ca. 650 m, Oct 1948 (fl Y), Wilson-Browne454 (ho- lode wanting. Pistillate inflorescencewith indument,
lotype: K; isotype: NY). color, and branchingas in staminateplants, the flow-
ers similarin size and shape;stamenssterile?,dimen-
Trees, or shrubs,to 7 m. Branchletsslender,mid-
sions as in staminateflowers;ovary glabrous,ovoid;
way along flush 2-3 mm diam., distally weakly an-
style stout, weakly distinguishedfrom ovary; stigma
gular,soon terete,rustytomentose,the surfacebarely
tri-lobed,0.3 mm diam. Fruitsborne on slenderclav-
visible or exposed, the hairsshort,to 0.3 mm, straight
ifonn pedicels of up to 1.5 X 0.4 cm; cupules shal-
to crooked,erect to ascending;terminalbuds slender,
lowly hemisphericalto patelliform,to 7 mm diam.,
1 X 0.6 mm, densely pubescent, the hairs as on
glabrous,the marginsundulate;drupesovoid, to 2 X
branchlets, ascending. Leaves alternate,widely and 1.3 cm.
evenly spaced along currentflush;petioles slender,to
1.3 x 0.2 cm, terete, the indumentas on branchlets; Distribution (Fig. 43) and ecology. Small trees
laminaechartaceous,plane to subbullate,narrowlyto or shrubsfrom westernAmazoniaand adjacentlower
broadly ovate, 5-10 X 1.5-5 cm, the base obtuse to montane slopes from Peru to Venezuela and east to
acute, briefly decurrent,the apex acute, acuminatefor the Guianas and NE Brazil at ca. 100-1100 m. Ap-
up to 1.5 cm, the marginsminutelyrecurvedthrough- parentlyabsentin the Amazon basin althoughknown
out; uppersurfacedull grey, minutelypunctulate,the to occur in both flooded and well-drainedsoils. Most
midribandsecondariesprominulous,the higher-order flowering specimens collected from September
venationimmersed;lower surfacesparselypubescent, through November, but also in January and May.
the hairs erect to ascending, slightly denser on main Fruitsavailableyear round.
veins, all vein orders raised, their prominence de-
creasing with rank; secondary veins 3-(4) per side, Representative specimens examined. COLOMBIA.
AMAZONAS: Leticia, ParqueNacional NaturalAmacayacu,
the lowermostpair suboppositeshortlyabove the leaf
120 m, 30 Mar 1992 (st), Rudas et al. 4057 (MO). ANTiO-
base, ascending at 50-60?, arcuate,upperpairs more QUIA:San Luis, Vereda Berlin, 400-530
m, 26 Oct 1989
obtuse, emerging around or beyond midlamina, (fr), Cogollo & Ram(rez4361 (MO).
abruptlyascending after midcourse,loop-connected; VENEZUELA. AMAZONAS: Atabapo, camino entre
tertiaries roughly horizontal, between secondaries Culebray la falda del extremo norte del Cerro Duida, 220
straight or forked. Staminate inflorescences evenly m, 9 Feb 1982 (fr), Steyermarket al. 126285 (HBG, MO,
spaced along current flush in the axils of foliage NY); CulebraSavanna,150 m, 14 Nov 1950 (fl Y), Maguire
leaves, to 3 cm long with 4 lateralbranches,branch et al. 29436 (F, G, GH, MO, VEN). BOLiVAR: Quebrada0-
orders2-3, the highest orderdichasial,lax, the flow- paru-ma, 1065-1220 m, 20-21 Nov 1944 (fl i, fr), Stey-
ers distant, the axes sparsely pubescent, the hairs as ermark60428 (F); Selva de Oparuma,Kavanay6n,30 May
1946 (fl 6), Lasser 1881 (F, US).
on branchlets;bractsand bracteolescaducousby an-
GUYANA. ESSEQUIBO:Rupununi,KuyuwiniLanding,
thesis, lanceolate,densely rustypannose;pedicels te- 250-350 m, 25 Oct 1992 (fl Y), Jansen-Jacobset al. 3085
rete, to 2 mm long, those supportingsecondaryflow- (B, MO, NY, U); Potaro-Siparuni,IwokramaRainforestRe-
ers slightly shorter. Flowers rotate, 3 mm diam., serve, 8-16 Jul 1997 (st), Chanderbaliet al. 250 (MO).
sparselypubescentoutside, the hairsappressedto as- SURINAME. Lucie R., small granitic islands in river
cending, silvery grey; receptaclebroadly infundibu- nearconfluenceof Oost R., 225 m, 12 Sep 1963 (fl Y), Irwin
liform,0.3 X 1.3 mm, densely rustytomentoseinside. et al. 55631 (F, NY, S, U, US).
Tepals chartaceous,ovate, 0.6 X 0.5 mm, spreading ECUADOR. NAPO: Tena,Estaci6nBiol6gica JatunSa-
to recurvedat anthesis,the inner surfacesparselysil- cha, 400 m, 3-6 Sep 1989 (fr), Palacios 4386 (G, MO, NY).
very grey-strigose. Stamens of whorls I and II stipi- PASTAZA: UNOCAL petroleumexplorationwell site "Ma-
zaramu,"390 m, Apr 1990 (fr), Beck et al. 1054 (MO). Suc.
tate, 0.5 mm tall, the antherstransverselyoblong, 0.3
UMBiOS: Gonzalo Pizarro,30 km NE of Lago Agrio, 1050
x 0.5 mm, glabrous,the apex truncateto emarginate, m, 23 Mar 1990
(fr), Cer6n et al. 9226 (MO); Lago Agrio,
the connectives level with or reduced between the 2 Reserva FaunisticaCuyabeno,230 m, 13 Nov 1991 (fl Y),
locelli, these suborbicular,introrse,the filamentslam- Palacios et al. 8840 (HBG, MO, NY). ZAMORA-
inar, much narrower than anthers, sparsely grey- CHINCHIPE:Nangaritze,Miazi, 900-1000 m, 21 Oct 1991
strigillose; whorl III stamens broadly stipitate, 0.5 (fl ?, fr), Palacios et al. 8585 (MO, NY); Zamora,Parque
SYSTEMATICTREATMENT
CmV. ...-.w,s.+W
.,,.
FIG. 43. Distributionof Endlicheniagracilis anldE. acuminata.
Nacional Podocarpus,GuardernaRio Bombuscaro,sendero
al Mirador, 1100 m, Jan 1995 (fl 9), Palacios & lirado
13298 (MO).
PERU. AMAZONAS: Condorcanqui,El Cenepa, Comu-
nidad de Tutino, 500 m, 20 Jul 1997 (fr), Rojas et al. 91
(MO). LoRETo: Alto Amazonas, Rfo Marant6n,Pongo de
Manseriche, Cerros Campanquiz,500-550 m, 19-21 Oct
1962 (fl d), Wurdack2339 (F, G, GH, K, NY, S, UC); Rfo
Huallaga,Shucushuyacu,250 m, 14 Sep 1981 (fr), Vasques
& Jaramillo 2501 (F, MO, NY). MADRE DE Dios: Manu,
Parque Nacional Manut,Cocha Cashu Station, Rio Manu,
250 m, 3 Sep 1989 (fl 9), Foster & Beltrdn 13050 (MO).
BRAZIL. ACRE:Brasileia, Seringal Porongaba,Colo-
cagko Sao Jose, 30 May 1991 (fl 9, fr), Daly et al. 6795
(MO, NY). AMAPA: Rio Agauaria,9 Oct 1961 (fl d), Pires
et al. 51599 (COL, G, GH, NY, US); Mun. Oiapoque, BR
156 between Cal,oene and Oiapoque, 17 km SSE of Oia-
poque, 3 Dec 1984 (fr), Mori et al. 17170
(MO).MARANHAo:Mun. Monqao, basin of Rfo Tariaqu,
Ka'apor Indian Reserve, 7 km from Urutawy, 5 Jun 1985
(fr), BalMe988 (MO).
Local names and uses. Venezuela: muneu-yek.
Brazil: mahamira, aiju'ywahu. According to W. L.
119
.7
^i --5W1z,
;E'--, r.
Balee, used as a hunting charmby the Ka'aporIndi-
ans of NE Brazil. The leaves are rubbedon the head
of a hunterto help him kill maha deer.
Endlicheriagracilis is immediatelyrecognizedby
its rotate flowers and the minutely punctulateupper
surfaceof triplinervedovate leaves thatassumea slate
grey color in the dried state. As noted by Kostermans
(1937), the androeciumis remarkablywell-developed
in pistillate flowers with stamens as large as those in
staminate flowers and anther valves often dehisced.
Further, in at least one collection, Jansen-Jacobs
3085, pollen is presentinside locelli althoughthe pis-
til is clearly fertilizedand developingtowardfruit.As
staminateflowerslack a pistillode altogether,it seems
likely that E. gracilis is androdioeciousrather than
dioecious. Still, an ITS sequence taken from Chan-
derbali et al. 250 assigns this species to the Endli-
cheria-Rhodostemonodaphneclade (Fig. 2), wherein,
as so far known, other species are dioecious. Therein
its affinitiesareunclear.The androeciumof E. gracilis
is similar to that of E. paniculata and its allies, but
120 FLORA NEOTROPICA

the slender claviform pedicels and shallow cupules flowers slightly shorter.Flowers rotate, 3 mm diam.,
with undulatemargins are reminiscentof E. sprucei sparsely grey-strigillose outside; receptacle cyathi-
and E. longicaudata, species here compared with form, 0.5 X 1 mm, densely silvery grey-tomentose
members of Rhodostemonodaphne.The appearance inside. Tepals membranaceousto chartaceous,ovate,
of remarkablysimilarleaves andfruitsin R.parvifolia 1 X 0.7 mm, spreadingat anthesis, the inner surface
also seems to supportaffinities with Rhodostemono- glabrous.Stamensof whorlsI and II stipitate,0.5 mm
daphne.Wereit not for coriaceousleaves with smooth tall, the antherstransverselyoblong, 0.3 X 0.5 mm,
uppersurfacein R. parvifolia, the two species would glabrous,the apex truncate,the connectiveslevel with
be separableonly in flower, themselves so similarin the 2 locelli, these suborbicular,introrse,the filaments
externalappearancethatit would be necessaryto note laminar,much narrowerthananthers,glabrous;whorl
differencesin stamen shape and locelli number. III stamenssessile, 0.6 mm tall, the anthersdepressed-
oblong, 0.3 X 0.4 mm, erect, locelli 2, extrorse-
latrorse,the filamentsbroaderthananthers,widening
58. Endlicheria acuminata Kosterm.,Recueil Trav. further towards base, laminar, glabrous, the basal
Bot. Neerl. 34: 553. 1937. Type. Brazil. Ama- glands sessile, globose, relatively large, filling the
zonas:Lago JuruaMiry,Jun 1901 (fl d), Ule 5581 space between filaments;whorlIV wanting;pistillode
(holotype:L; isotypes: G, HBG, K). wanting.Pistillateinflorescencewith indument,color,
and branchingas in staminateplants,the flowerssim-
Trees to 15 m. Branchletsslender,midway along ilar in size and shape; stamenssterile, smaller;ovary
flush 2-3 mm diam., angular,sparselypubescent,the glabrous, ovoid; style stout, weakly distinguished
surface barely visible to clearly exposed, the hairs from ovary; stigma broadly tri-lobed, 0.5 mm diam.
silvery-greyto pale brown, short,to 0.3 mm, straight Fruits borne on slender cylindrical pedicels of up to
to crooked,erect or appressed;terminalbuds slender, 1.3 X 0.3 cm; cupules patelliformor shallowly hem-
3 X 1 mm, densely rusty pubescent, the hairs as on ispherical,to 0.5 X 1 cm, glabrousoutside, sericeous
branchlets,ascending. Leaves alternate,widely and inside, the marginsentire;drupesellipsoid, to 3.5 x
evenly spaced along currentflush;petioles slender,to 1.5 cm.
1 X 0.2 cm, semi-terete, the indument as on bran-
chlets; laminae membranaceous to chartaceous, Distribution (Fig. 43) and ecology. Small trees
plane, ovate to ovate-elliptic, 5-15 X 2-4 cm, the of flooded forests in western Amazonia at ca. 100-
base obtuseto acute,brieflydecurrent,the apex acute, 300 m. Flowering from April to August, fruits avail-
acuminatefor up to 1.5 cm, the marginsminutelyre- able year round.
curvedthroughout,or flat aftermidlamina;uppersur-
face dull greyish green, waxy, the midriband second- Representative specimens examined. COLOMBIA.
aries immersedto sunken, the higher-ordervenation VAUPES: San Jose del Guaviare,La Libertad,cercaa Re-
raised; lower surface sparsely pubescent, the hairs serva Indfgena"Asunci6n,"CanioGrande,300 m, 10 Aug
1989 (fl d), Marulanda & Mdrquez1108 (HUA 89052),
erect or appressed,uniformlydistributed,all vein or-
Cano Nare, 280 m, 17 Aug 1989 (fl ?, fr), Marulanda &
ders raised, their prominence decreasing with rank; Ma'rquez1536 (HUA).
secondaryveins 2-4 per side, the lowermostpair su- ECUADOR. NAPO:Aguarico,ReservaFaunfsticaCuy-
bopposite shortly above the leaf base, ascending at abeno,LagunaZancudoCocha (Iripari),230 m, 28 Sep 1991
50-60?, almost straightto arcuate,otherpairsemerg- (fl 5), Palacios et al. 7804 (MO); Lagunas de Cuyabeno,
ing around or beyond midlamina, more obtuse, 300 m, 21 Aug 1981 (fl 5, juv), Brandbygeet al. 33861
arcuate, weakly loop-connected; tertiaries roughly (AAU). SUCUMBiOS:Lago Agrio, ReservaFaunfsticaCuy-
horizontal, between secondaries once-forked to abeno, 230 m, 30 Sep 1991 (fl Y), Palacios et al. 7884
straight. Staminate inflorescences evenly spaced (MO).
along currentflush in the axils of foliage leaves, to 5 PERU.LORETO:Maynas, Rfo Amazonas, Yanamono
cm long with 3 lateralbranches,branchorders2-3, Explorama Tourist Camp, halfway between Indiana and
mouth of Rfo Napo, 140 m, 26 Jul 1991 (fl 5), Vdsquez&
lateral second-orderbranchesoften reducedto a sin-
Grdndez17475 (MO); Requena,Sapuena,ArboretoJenaro
gle flower,internodesof terminalcymes reduced,the Herrera,CanioLobito, 160 m, 17 Aug 1994 (fl 5), Ortizet
flowers clusteredin pseudo-umbels,the axes sparsely al. 124 (MO). MADREDE Dios: Puerto Maldonado, Rfo
pubescent,the hairs 0.3 mm long, greyish, appressed Madre de Dios, 250 m, 22 Apr 1977 (fl Y), Gentry et al.
to ascending;bracts and bracteoles caducous by an- 19636 (MO, NY); Tambopata,Cuzco Amaz6nico, Fundo
thesis, lanceolate, the indumentas on axes; pedicels Concepci6n, Rfo Madre de Dios, 200 m, 19 May 1989 (fl
terete, to 2 mm long, those supporting secondary 5 ), Phillips & Ntiuiez52 (MO).
SYSTEMATICTREATMENT
BRAZIL. ACRE: Cruzeiro do Sul, Rio Jurua & Rio
Moa, near Uruburetama,25 May 1971 (fl d), Maas et al.
P13307 (HBG, INPA, NY); Rio Muru, Seringal Vit6ria
Velha, Colocagao BarroVermelho,20 Jun 1995 (fl d), Fi-
gueiredo et al. 908 (MO); SenadorGuiomard,basin of Rio
Purus,Rio Iquiri, 6 Mar 1997 (fr), Daly 9293 (MO). AMA-
ZONAS: Rio PurUs,between Campina and Tambaqui,Jun
1971 (fl 3), Prance et al. 13391 (HBG, INPA, NY); Lago
Preto, 2 km N of Labrea,25 Jun 1971 (fl Y), Prance et al.
13695 (HBG, INPA, K, MO, NY, US).
BOLIVIA. BENI: Marban, parque Isiboro-Secure,de
Puerto San Lorenzo, 200 m, 25 May 1992 (fr), Seidel et al.
6557 (MO);Yacuma,E of San Bora, Bosques de Chimanes,
near Rfo Mozeruma, 250 m, Nov 1988 (fr), Foster et al.
12483 (MO).
Local names. Peru:cunchimoena,muena.Brazil:
louro, louro do igapo, louro preto do igap6.
Endlicheria acuminata shares triplinervedovate
leaves with E. gracilis. In both as well, the flowers
are rotate,stipitatewhorl I and II stamenshave trans-
versely oblong antherswith truncateapices, andlarge
basal glands completely fill the space between fila-
ments of all staminalwhorls.Yetthe two arenot easily
confused because the upper leaf surface in E. acu-
minata is smooth and waxy rather than minutely
punctulate, and tertiaries are raised ratherthan im-
mersed.
As in Endlicheriaanomala and E. paniculata, ei-
ther erect or appressedhairs constitute the vestiture
of branchletsand lower leaf surfacesof E. acuminata
withoutaccompanyingdifferences.Typematerialand
a few other specimens are tomentose,but in most the
hairsare definitelyappressed.This subsericeousform
of E. acuminatais vegetativelyindistinguishablefrom
E. krukovii,where, however,the flowersareinfundib-
uliform, all anthersare ovate with acuminateapices,
and basal glands are minute and inconspicuous. As
cupules in the two are also indistinguishable,both
being shallowly hemisphericaland densely sericeous
inside, only staminodesremnanton cupule rims can
assign fruitingspecimens to species.
59. Endlicheria krukovii (A. C. Sm.) Kosterm.,Re-
cueil Trav.Bot. Neerl. 34: 531. 1937. Aniba kru-
kovii A. C. Sm., Phytologia 1(3): 117. 1935. Type.
Brazil. Amazonas: Near mouth of Rio Embira
(tributaryof Rio Taracau),26 Jun 1933 (fl Y),
Krukoff5023 (holotype:NY; isotypes: A-n.v., G,
K).
Trees, 3-22 m. Branchletsslender,midway along
flush 3-4 mm diam., angular,densely strigose, the
121
surfaceobscuredor barely visible, the hairs short, to
0.3 mm, straight,appressed,grey to light brown;ter-
minal buds slender,3 X 0.6 mm, densely pubescent,
the hairs as on branchlets,ascending. Leaves alter-
nate, widely and evenly spaced along currentflush;
petioles slender, to 2 x 0.2 cm, semi-terete,the in-
dument as on branchlets; laminae chartaceous to
membranaceous,plane, ovate to obovate, 10-25 X 5-
10 cm, the base acute, briefly decurrent,the apex
acute, acuminatefor up to 3 cm, the marginsminutely
recurvedthroughout,or flatbeyondmidlamina;upper
surface dull greyish green to olive-brown,waxy, the
midrib and secondaries sunken to immersed, the
higher-ordervenation raised; lower surface sparsely
pubescent,the hairs as on branchlets,appressed,uni-
formly distributed,all vein ordersraised,theirprom-
inence decreasingwith rank;secondaryveins 2-4 per
side, the lowermostpairsuboppositeshortlyabovethe
leaf base, ascending at 50-600, arcuate to almost
straight,other pairs emergingaroundor beyond mid-
lamina,moreobtuse, arcuate,weaklyloop-connected;
tertiaries roughly horizontal, between secondaries
once-forked to straight. Staminate inflorescences
evenly spaced along currentflush in the axils of fo-
liage leaves, to 20 cm long with 12 lateralbranches,
branchorders3-4, the highestorderdichasial,lax, the
flowers distant,the axes densely pubescent,the hairs
appressed,increasing in density and obscuring epi-
dermisdistally;bractsandbracteolescaducousby an-
thesis, lanceolate, the indumentas on axes; pedicels
terete, to 1 mm long, those supporting secondary
flowers slightly shorter.Flowers infundibuliform,2
mm diam., densely silvery-grey pubescent outside,
the hairsappressed;receptacleinfundibuliform,0.6 X
0.6 mm, glabrous inside. Tepals membranaceousto
chartaceous,broadlyovate, 0.6 x 0.6 mm, ascending
at anthesis, the inner surface glabrous, the margins
and apex inside minutely papillose. Stamens of
whorlsI andII stipitate,0.6 mm tall, the anthersovate,
0.4 x 0.3 mm, glabrous,the apex apiculate,the con-
nectives prolonged between the 2 locelli, these
obliquely hemispherical, introrse-latrorse,the fila-
ments laminar,much narrowerthananthers,glabrous;
whorl III stamens sessile, 0.6 mm tall, the anthers
ovate, 0.4 x 0.3 mm, erect,locelli 2, extrorse-latrorse,
the filaments broader than anthers, wider towards
base, glabrous, the basal glands sessile, minute,
globose-apiculate;whorl IV wanting;pistillode want-
ing. Pistillate inflorescencewith indumentand color
as in staminateplants,but shorterand with fewer lat-
eral branches, the flowers slightly deeper; stamens
sterile, smaller; ovary glabrous, ovoid; style stout,
122
weakly distinguishedfrom ovary; stigma stronglytri-
lobed, 0.6 mm diam. Fruits borne on slender cylin-
drical pedicels of up to 1 X 0.3 cm; cupules patelli-
form to shallowly hemispherical,to 0.5 X 1 cm, gla-
brous outside, sericeous inside, the margins entire;
drupesellipsoid, to 2.5 x 1.3 cm.
Distribution (Fig. 44) and ecology. Small to
medium-sizedtrees in nonfloodedforests of the east-
ern Andean foothills but also in seasonally inundated
lowlands in western Amazonia at ca. 200-1000 m.
Flowering specimens collected from February
through May, and in October and November; fruits
availableyear round.
Representative specimens examined. ECUADOR.
NAPO:Estaci6n ExperimentalINIAP-Napo,Payamino,Re-
serva Floristica "El Chuncho,"280 m, 17 May 1986 (fr),
Baker 7017 (QAME-n.v., QCNE-n.v., MO, NY); SantaCe-
cilia, 340 m, 30 Mar 1972 (fl d, juv), Dwyer & Simmons
9748 (MO). PASTAZA:Pastaza, 365 m, Oct 1-20 1990 (fl
d), Espinoza & Coba 369 (MO). SucuMsios: Gonzalo Pi-
zarro, Bosque ProtectorLos Cedros, cuenca del Rio Tigre,
4-a ~ ~~~~~~~-
FIG. 44. Distributionof Endlicheniakrukoviiand E. nilssonii.
FLORA NEOTROPICA
1000 m, 17 Mar 1992 (fl c), lipaz et al. 709 (MO); Lago
Agrio, Reserva FaunfsticaCuyabeno, 230 m, 20 Oct 1991
(fl 9), Palacios et al. 8027 (MO).
PERU. AMAZONAS: Bagua, Imaza, Comunidadde Ya-
mayakat, 600 m, 9 Jun 1997 (fl 6), Vdsquezet al. 23948
(MO); Rfo Santiago, QuebradaKusu',Bagua, 250 m, 30-31
Oct 1962 (fl 6), Wurdack2493 (K, NY, S, UC, US). Cuzco:
Rio Mapituriani,La Convenci6n, 14 Sep 1976 (fr), Was-
shausen & Encarnaci6n649 (K, MO, NY, US). HukNUco:
Pachitea, Honoria, Quebradade Macuya al oeste del Rio
Pachitea, 300-400 m, 6 May 1968 (fl d), Schunke 2570
(MO). LORETO:Alto Amazonas, Puerto Melendez, below
Pongo de Manseriche, s.d. (fl 6), Tessmann4734 (NY);
Maynas, Iquitos, km 44 carreteraIquitos-Nauta, terreneos
del Comite de Reforestaci6nIquitos (CRI), 150 m, 14 Mar
1989 (fr), Vdsquezet aL 11928 (HBG, MO). MADRE DE
DIos: Manu, ParqueNacional Manu',Cocha Cashu Station,
Rio Manu, 350 m, 1 Feb 1981 (fr), Foster & Wright8254
(MO); Tambopata,Cuzco Amaz6nico, trail to Lago San-
doval across Rio Madre de Dios, ca. 12 km E of Puerto
Maldonado,200 m, 21 Feb 1990 (fl T,juv), Gentry& Nufnez
69392 (MO). PAsco: Oxapampa,Palcazt, 300-600 m, 7
Dec 1984 (fr), Hartshorn et al. 2689 (MO, NY, U). SAN
Air
DOUBTFULNAMES AND EXCLUDEDTAXA 123

MARTiN: Huinguillo, 500-600 m, 21 Mar 1962 (fl d, juv), pairs loop-connected,the tertiariesroughly horizon-
Woytkowski7179 (HBG, MO); San Martin,5-15 km E of tal, between secondariesstraightto forked.Staminate
Shapajaon rd. to Chazuta,200-300 m, 9 Apr 1986 (fl d), andpistillateinflorescencesunknown.Fruitsborneon
Knapp& Mallet 7024(F, MO, NY, US). UCAYALI: Coronel
claviformpedicels of up to 1 X 0.4 cm; cupuleshem-
Portillo,Yarinacocha,Nueva Esperanzade Panaillo, 148 m,
ispherical,to 0.5 X 1 cm, glabrousoutside andinside,
1 Apr 1988 (fl d, juv), Vdsquez& Jaramillo10458 (F, MO,
NY). the marginsentire;drupesellipsoid, to 1 X 0.7.
BRAZIL. ACRE:Tarauaca,26 May 1977 (fl d), Mir- Distribution (Fig. 44) and ecology. Known only
anda 20 (INPA);Xapuri,Rio Acre, 10 Nov 1991 (fr), Daly
from the type material,takenfrom a tree found fruit-
et al. 7282 (MO, NY).
ing in April at ca. 1200 m in the highlandsof eastern
BOLIVIA. PANDO: Manupiri, 35 km al N de Puerto
America, 200 m, 30 Apr 1994 (fl d), Jardim594 (MO).
Venezuela.

Local names. Ecuador:ajua. Peru:moenilla, tin- Endlicheria nilssonii is a poorly understoodyet

chi, yuwich (Huambisa).Brazil:louro. distinctive species. The combinationof densely seri-
ceous branchletsbut only sparse appressedhairs on
Among species with triplinervedleaves, Endli- the leaves below is otherwisefound only occasionally
cheria krukoviiis conspicuous for its relativelymul- in E. metallica. Flowers are not known,but one stam-
tiflorousinflorescencewith densely pubescentinfun- inode found persisting in fruit shows a distinct and
dibuliform flowers within which all stamens have densely pubescentfilament and an ovate antherwith
ovate anthers.Yet, from representativesof E. acumi- a truncateapex. However, a tepal, also found on the
nata with similar appressed indument, only floral cupule margin,provides a better appreciationof the
charactersare distinguishing.AlthoughE. acuminata flowers of E. nilssonii. This tepal is ratherlarge and
has been found only in flooded habitat,and most col- fleshy but, more important,densely pubescent with
lectors have indicated terra firme conditions for E. papillose hairs covering the inner surface. As such
krukovii,any ecological distinctionseems weak since tepals only occurwith rotateflowers(e.g., E. sericea),
occasionalreports,e.g., Vdsquezet al. 23948, suggest it is most likely that flowers of E. nilssonii are rotate.
that E. krukoviialso toleratesinundation. Even if not the case, the combinationof pubescent
fleshy tepals, densely sericeousbranchlets,andsparse
appressedhairs on the leaves below is unique in En-
60. Endlicheria nilssonii C. K. Allen, Mem. New dlicheria.
YorkBot. Gard. 10(5): 66. 1964. Type.Venezuela. In the truncateantherapices, pinnatevenation,and
Bolivar: Wooded ridge, La Danta, along provi- sparseappressedindumenton leaves below, this spe-
sional road from km 125 to 127 between Leupa cies approachesto subsericeousforms of E. panicu-
and Cerro Venamo, 1200 m, 17 Apr 1960 (fr), lata, with which it is placed, with hesitation. Better
Steyermark& Nilsson 255 (holotype:NY). materialmay later more clearly indicateits affinities.
Trees to 30 m. Branchlets stout, midway along
flush 3-5 cm diam., angular,sericeous, the surface DOUBTFUL NAMES AND
concealed by the indumentcover, trichomes short,to EXCLUDED TAXA
0.1 mm, straight, appressed, light yellowish green; Endlicheria balsamea Vattimo,Rodriguesia33 (46):
terminalbuds plump, 4 X 3 mm, sericeous. Leaves 23. 1978. Type.Brazil.Rio de Janeiro:14 Jan1932
alternate, widely and evenly spaced along current (fr), Paulino & Victorios.n. R 148881 (holotype:
flush; petioles robust, to 1.5 X 0.3 cm, striate, the R; isotypes: R [7 sheets]).
indumentas on branchlets;laminaecoriaceous,plane,
ovate to elliptic, 10-15 X 4-6 cm, the base acute, The protologue describes a representative of
briefly decurrent,the apex acute, acuminatefor up to Aiouea, since it states thatnine two-locellate stamens
0.5 cm, the margins minutely recurvedthroughout; and well-developed whorl IV staminodes occur to-
upper surface deep green, waxy, the midrib and sec- gether with fruits. However,I could not find stamens
ondary veins immersed, the tertiariesprominulous; or staminodesin the ample type materialbefore me,
lower surface sparsely pubescent, the hairs as on and I thereforerefrainfrom proposinga new combi-
branchlets,appressed,uniformlydistributed,all vein nation.
ordersraised, theirprominencedecreasingwith rank;
secondary veins 3-4 per side, ? evenly spaced,
slightly more distantaroundmidlamina,ascendingat Endlicheria debilis Kosterm., Recueil Trav. Bot.
45-60? (more acutely towards apex), arcuate,distal Neerl. 34: 555. 1937. Type. Peru. Loreto: Alto
124
Amazonas, Balsa Puerto (lower Rio Huallagaba-
sin), 150-350 m, 28-30 Aug 1929 (fr), Killip &
Smith 28400 (holotype: NY; isotypes: B-n.v., F,
US) = Rhodostemonodaphnedebilis (Kosterm.)
Chanderbalicomb. nov.
Kostermans apparently overlooked the four-
locellate staminodeson the isotype depositedat F, and
found before me by V. Koch on the isotype at US.
Those from androecialwhorls I and II are narrowly
stipitatewith obovateantherswith the locelli arranged
in a very shallow, almost horizontal, arc, and the
whorl III staminodeis sessile with four locelli in two
superimposedpairs. These staminodes assign Killip
& Smith28400 to Rhodostemonodaphne,whereinR.
crenaticupulais an excellent vegetative match. The
latter differs in that whorl I and II stamens in both
staminateandpistillateplantsarelargerandmore ses-
sile and cupule marginsmore stronglylobed.
Endlicheria endlicheriopsis (Mez) Kosterm.,
Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 25:
43. 1936. Type. FrenchGuiana,Melinon 605 (ho-
lotype: P-n.v.) = Ocotea endlicheriopsisMez.
Endlicheria goeldiana Vattimo,An. XV Congr.Soc.
Bot. Bras., 169. 1964. Type. Brazil. Amazonas:
Rio Purus, 20 Jun 1903 (fl), Goeldi s.n. HAMP
3902 (holotype:R).
The type materialavailableto me consists of de-
tached leaves. As there is no sign of the flowers de-
scribed in the protologue, I cannot confirm or deny
that this materialbelongs to Endlicheria.
Endlicheria grandis Mez, Jahrb.Konigl. Bot. Gart.
Berlin 5: 124. 1889. Type. French Guiana.Meli-
non s.n. (holotype: B-n.v.; isotypes: NY-n.v. P-
n.v.) = Rhodostemonodaphne grandis (Mez) Roh-
wer.
Endlicheria impressa (Meisn.) Mez, Jahrb.Konigl.
Bot. Gart. Berlin 5: 132. 1889. Phoebe impressa Endlicheria tomentella Mez, Jahrb. Konigl. Bot.
Meisn., in DC., Prodr. 15(1): 33. 1864. Type. Gart. Berlin 5: 115. 1889. Type. Peru. Ancach:
French Guiana. Patris s.n. (holotype: G-n.v.) = Near Moro, Pearce s.n. (holotype: K-n.v.) =
Aiouea impressa(Meisn.) Kosterm. Aiouea tomentella(Mez) Renner.
Endlicheria juruensis (A. C. Sm.) Kosterm.,Recueil
Trav.Bot. Neerl. 34: 542. 1937. Aniba juruensis
A. C. Sm., Phytologia 1(3): 116. 1935. Type. Bra-
FLORANEOTROPICA
zil. Amazonas: Near mouth of Rio Embira(trib-
utary of Rio Tarauaca),Jun 1933 (fl 9, fr juv),
Krukoff4775 (holotype:NY; isotypes: A, F, G, K,
MO, U) = Rhodostemonodaphnejuruensis (A. C.
Sm.) Chanderbalicomb. nov.
Staminodesin the type materialare four-locellate
with the locelli arrangedin the shallow apical arch
typical of Rhodostemonodaphne.Othercharactersas-
sociating this species with Rhodostemonodaphnein-
clude sessile stamens and dense papillosityin the in-
ner surfaceof horizontallyspreadingtepals. After the
type collection, severalmore pistillaterepresentatives
have been found (listed below), but staminateplants
are still unknown.
Additional specimens examined. ECUADOR. CAR-
CHI: Tulcan, ParroquiChical, ReservaIndigenaAwa, 1200
m, 23-27 May 1992 (fl 9), Tipazet al. 1104 (MO), 900 m,
May 1992 (fl Y), Quelal et al. 704 (MO). ESMERALDAS:
San Lorenzo,carreteraLita-San Lorenzo,500 m, 9 Jul 1990
(fr), Rubio et al. 454 (MO). MORONA-SANTIAGO:Along
new rd. Mendez-Morona,650 m, 16 Aug 1989 (fl Y), van
der Werff& Gudinio11136 (MO).
PERU. SAN MARTiN: Mariscal Caceres, Tocache
Nuevo, Camino a Santa Rosa, Rfo Mishollo, 350-370 m, 5
Aug 1973 (fr), Schunke6726 (F, MO, NY).
BOLIVIA. LA PAZ: Prov. Franz Tamayo, Serranfade
Chepite,700 m, 3-8 Apr 1992 (fl Y, fr), Killeen3832 (MO).
Endlicheria loretensis 0. Schmidt, Repert. Spec.
Nov. Regni. Veg. 31. 178. 1933. Type. Peru. Lor-
eto: Mishuyacu,nearIquitos, 100 m, Apr 1930 (fl
J), Klug 1258 (holotype: F-n.v.; photo neg.
40486 ex F) = Ocotea or Rhodostemonodaphne.
Endlicheria maguireana C. K. Allen, Mem. New
YorkBot. Gard.10(5): 68. 1964. Type.Venezuela.
Amazonas:RioYatua,1100-1150 m, 26 Dec 1957
(fl), Maguireet al. 42521 (holotype:NY; isotypes:
K-n.v., MO, S, U, US-n.v.) = Aiouea maguireana
(C. K. Allen) Renner.
Endlicheria zapoteoides Lundell, Wrightia 1: 145.
Type. Mexico. Chiapas: Cascada near Siltepec,
1600 m, 1 Mar 1945 (fl, fr), Matuda 5153 (holo-
LITERATURECITED 125

type: TEX-n.v.; isotypes: MO, US-n.v.) =
Beilschmiediazapoteoides (Lundell) Kosterm.
Goeppertia argentea (Griseb.)Meisn., in DC., Prodr.
15(1). 174. 1864. AydendronargenteumGriseb.,
Fl. Brit.W. Ind. Isl. 1: 285. 1860. Type.Dominica,
Imray365 (holotype:GOET-n.v.) = Aniba brac-
teata (Nees) Mez.
Goeppertia caudata Meisn., in DC., Prodr. 15(1).
175. 1864. Syntypes. Brazil. Amazonas:Prov.Rio
Negro, prope Panuread Rio Vaupes,Spruce2638
(syntypes:K-n.v., M-n.v.) = Ocotea debilis Mez.
Goeppertia geminiflora Meisn., in DC. Prodr.15(1).
175. 1864. = Systemonodaphne geminiflora
(Meisn.) Mez, providedMartin s.n. from French
Guianais cited as the type. = Acrodiclidiumgem-
iniflorum(Meisn.) Mez, provided Guillemin231
from Rio de Janeiro,Brazil, is cited as the type.
ACKNOWLEDGMENTS
This study was undertakenas partof my doctoral
dissertationresearchat the Universityof Missouri-St.
Louis and the Missouri Botanical Garden.I am in-
debtedto these two institutionsfor the opportunityto
pursue graduate studies in their excellent facilities.
For providingunlimitedaccess to his expertisein the
Lauraceae, my dissertation advisor, Henk van der
Werff,is specially thanked.Grantsfromthe Mallinck-
rodt Foundation administeredby the International
Centerfor TropicalEcology, Universityof Missouri-
St. Louis, and from the SmithsonianInstitution'sBi-
ological Diversity of the GuianasProgram,are grate-
fully acknowledgedfor their generous supportof the
laboratoryand field componentsof this study,respec-
tively.
I thank Jim Solomon, Manager of the Missouri
Botanical GardenHerbarium,for requestingloans of
Endlicheriaand for providing a place in the herbar-
ium where they could be studied in relativeluxury.I
also thank the curatorsof institutionsthat provided
loans, sometimes searching for specific specimens
among theirindets, and allowing me to keep them for
the five-yeardurationof this study.
BarbaraAlongi preparedthe fine illustrationsof
the new species of Endlicheria.Mike Vieth helped to
take the SEM photographsof stamens.Henk van der
Werff correctedthe Latin diagnoses and Rosa Ortiz
correctedthe Spanishabstract.SusanneRenner,Henk
van der Werff, and the three reviewers, Mike Nee,
William Burger, and Santiago Madriinan,provided
useful comments on earlier drafts of the manu-
script.

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126
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Nees, C. G. D. 1833.Revisiolaurinarum ab Sellowioin
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NUMERICAL LIST OF TAXA
E. punctulataspecies group
1. E. punctulata (Mez) C. K. Allen
2. E. coriacea Chanderbali
Microlocellata species group
3. E. bullata Ducke
4. E. rubriflora Mez
FLORANEOTROPICA
GattungOcoteaAubl. (Lauraceae).Mitt. Inst.Allg.
Bot. Hamburg20: 3-278.
. 1993a.Nectandra.Fl. Neotrop.Monogr.60: 1-
332.
. 1993b.Lauraceae.Pp.426-437 in K. Kubitzki,
J. Rohwer& V. Bittrich(eds.),Thefamiliesandgen-
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. 2000. Towardsa phylogeneticclassificationof
the Lauraceae:evidencefrommatKsequences.Syst.
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, H. G. Richter & H. van der Werif. 1991.Two
new generaof neotropicalLauraceaeandcriticalre-
marks on the generic delimitation.Ann. Missouri
Bot. Gard.78: 388-400.
New or noteworthyplantsfromPeruandAmazonian
Brazil.Bull. TorreyBot. Club58: 87-110.
Stearn, W. T. 1992. Botanical Latin. Ed. 4. Timber
Press,Portland,OR.
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Vicentini, A., H. van der Werff & S. Nicholas. 1999.
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M. J. G. Hopkins, A. Vicentini, C. A. Sothers,
M. A. S. Costa,J. M. de Brito, M. A. D. de Souza,
L. H. Martins,L. G. Lohmann,P.A. C. L. Assuncbo,
E. da C. Pereira,C. F. da Silva, M. Mesquita,& L.
Proc6pio(eds.), Flora da ReservaDucke-Guiade
identificaodas plantasvascularesde umaflorestade
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ficationand direct sequencingof fungal ribosomal
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protocols:a guide to methodsand applications.Ac-
ademicPress,San Diego.
5. E. ferruginosa Chanderbali
6. E. longicaudata (Ducke) Kosterm.
7. E. sprucei (Meisn.) Mez
E. metallica species group
8. E. metallica Kosterm.
9. E. chrysovelutina Chanderbali
LIST OF EXSICCATAE 127

E. browniana species group E. canescens species group

10. E. browniana Mez 37. E. canescensChanderbali
11. E. jefensis van der Werff ex Chanderbali 38. E. xerampelaChanderbali
12. E. colombiana (Meisn.) Mez 39. E. citriodoravan derWerif
13. E. mishuyacensis A. C. Sm. 40. E. rubraChanderbali
14. E. pyriformis (Nees) Mez 41. E. vinotinctaC. K. Allen
15. E. formosa A. C. Sm. 42. E. oreocolaChanderbali
16. E. paradoxa Mez 43. E. szyszylowicziiMez
17. E. dysodantha (Ruiz & Pav.) Mez 44. E. duotinctaChanderbali
18. E. tomentosa Chanderbali 45. E. lorastemonChanderbali

Ampelodaphne species group E. sericea species group

19. E. aruncifiora (Meisn.) Mez 46. E. sericea Nees
20. E. arachnocome Chanderbali 47. E. bracteolata(Meisn.)C. K. Allen
21. E. arenosa Chanderbali 48. E. tschudyana(Lasser)Kosterm.
22. E. macrophylla (Meisn.) Mez 49. E. griseo-sericeaChanderbali
23. E. multiflora (Miq.) Mez 50. E. ruforamula Chanderbali
24. E. levelii C. K. Allen 51. E. aureaChanderbali
25. E. dictifarinosa C. K. Allen 52. E. lhotzkyi(Nees) Mez
26. E. melinonii Benoist 53. E. klugii0. C. Schmidt
27. E. reflectens (Nees) Mez 54. E. argenteaChanderbali
28. E. bracteata Mez 55. E. robusta(A. C. Sm.) Kosterm.
29. E. verticillata Mez
30. E. cocuirey Kosterm. E. paniculata species group
31. E. tessmannii 0. C. Schmidt 56. E. paniculata(Spreng.)J. F. Macbr.
32. E. directonervia C. K. Allen 57. E. gracilis Kosterm.
33. E. chalisea Chanderbali 58. E. acuminataKosterm.
34. E. glomerata Mez 59. E. krukovii(A. C. Sm.) Kosterm.
60. E. nilssoniiC. K. Allen
E. anomala species group
35. E. anomala (Nees) Mez
36. E. williamsii 0. C. Schmidt

LIST OF EXSICCATAE
In addition to all collections of Endlicheria examined, this index includes several specimens that were
previously annotatedas Endlicheriabut belong to other genera.
Abbott,J. R. & L. C. Isaacs,16484 (56).
Acevedo,P., 3432 (33).
Acevedo,P. & Cedeino,7416 (32).
Acevedo,P. & F. Ramfrez,9967 (cf. 59).
Acevedo, P. et al., 1370 (cf. 49); 3484 (27); 4834 (1);
4930, 6035 (14); 8646 (15).
Adalardo-Oliveira, A., 2643 (35); 2738 (22).
Agiular,M. & D. Castro,1119 (58).
Aguiar,I. J. A. et al., RUC-122(50).
Aguirre-Galviz, L. E., 1117 (13).
Albuquerque, B., 135, 136 (35).
Albuquerque, B. & L. Coelho,415 (35).
Albuquerque, B. & Elias,67-22 (43).
Albuquerque, B. & J. Lima,240 (35).
Albuquerque, B. et al., 570,742,765,778 (35);850 (13);
955, 1050 (35).
Alexiades,M. N. & C. Diaz, 849 (56).
Alexiades,M. N. & G. Nicole, 1042 (28).
Alexiades, M. N. et al., 437 (17).
Allen, C. K., 20 (48).
Allen, C. K. et al., 323 (21).
Aluisio, J., 267 (43).
Alvarez, E. et al., 482 [Ocotea cernua (Nees) Mez]; 1150
(7).
Amaral, I. L. et al., 109 (45); 157 (24); 448 (15); 488
(55); 1375 (15).
Ancuash, E., 133 (8); 179 (56); 283 (8); 404 (55); 1343
(32).
Andel, T. van et al., 293 (15).
Anderson, A., 1521, s.n. (46).
Anderson, W. R., 7759, 9604 (56); 12052 (35); 35171
(56).
Angeli, C., 587 (56).
Appun, C., 377 (23); 2006 (27).
Aquilar, M. & D. Castro, 566 (43).
Aranda, J. E. et al., 978 (48).
128
Araujo,D., 7321, s.n. R. 30974 (56).
Araujo,D. & R. de Oliveira,6779 (56).
Araujo,D. et al., 7918 (56).
Arbelaez,G. et al., 2490 (4).
Arbelaez,M. V. & F. Sueroke,711, 715 (21).
Argent,G. C. G. in P. W. Richards,6712 (52).
Argent,G. C. G. et al., 6577 (52).
Aristeguieta,L., 3012 (46); 3845, 5107 (48).
Asplund,E., 14682 (13).
Assunc,o, P.A. C. L., 128 (6).
Assun,co, P.A. C. L. & C. F. da Silva,553 (6);583 (14).
Assun,co, P. A. C. L. et al., 366 (6); 531 (33).
Aulestia,C. & M. Aulestia,975 (50); 1289 (10).
Aulestia,M., 3640 (8).
Aulestia,M. & J. Andi, 513 (56); Andi 516 (15); 841
(8).
Aulestia,M. & 0. Gonti,1755 (45); 1986 (50).
Aulestia,M. & A. Omehuat,3236 (45).
Aulestia,M. et al., 819, 1433 (15); 3593 (32).
Ayala,F., 817, 6781, 2954, 3026, 3039 (35); 3136 (cf.
59); 3491 (15); 3587 (35).
Aymard,G. & L. Delgado,6646 (47); 6752 (25); 8483
(33).
Aymard,G. et al., 372 (23); 5707, 7381 (35).
Bahia,R. P., 77 (6).
Baitello,J. B. & 0. T. Aguilar,s.n. SPSF-8148(56).
Baker,90 (6).
Baker,M. A., 7017 (59).
Balansa,2026 (56).
T. 3211
Baldwin,J. Jr., (19).
Balee, W.L., 988 (57).
Balee, W.L. & B. G. Ribeiro,1579 (6).
Balslev,H. et al., 97016 (55);97178 (15);97282 (cf. 59); Campos,J. & P. Lopez,4930 (37).
97283 (Ocoteacernua);97440, 97447 (55).
Bamps,P., 5145, 5326 (6).
Bang,A. M., 1676 (17); 1691 (43).
Barbosa,M., 866, 870 (27).
Barbour,P.J., 5742 (17).
Barclay,A. S. et al., 596 (35).
Barreto,M., 3344 (56).
Barrier,S., 2361, 2370, 2457, 3256 (46).
Barros,W.D. de, 223 (56).
Beard,J. S., 189, 228, 628 (46).
Beard,P., 1438 (46).
Beck, G., 3192 (33); 8176 (56).
Beck, G. & R. Foster,13983 (4).
Beck, G. & R. Haase,10129, 10146 (35).
Beck, G. et al., 19650 (cf. 59).
Beck, H. T. et al., 1054 (57).
Belanger,161, 979 (46).
Benitezde Rojas,C. et al., 5096 (56).
Bennett,B., et al., 4306 (57).
Benson,W., UNICAMP10847 (56).
Bentancur,J., 5404, 5405 (15); 5418 (14).
Berg,C. C., BG 770 (14).
Berg, C. C. & W.C. Steward,P 19913 (cf. 55); P19932 Cerner,116, 117 (35).
(24).
Berg,C. C. et al., P19527 (32); P19760 (35).
FLORANEOTROPICA
Berlin,B., 753 (56); 1781 (55); 1783 (32).
Bernal,R., et al., 2039a (15).
Bernardi,L., 1626 (25); 1789 (4); 18223, 18223b(56).
Bertoni,B. S., 916, 917 (56).
Bilby,R. et al., 230 (50).
Billiet, F. & B. Jadin,5966 (17).
Black,G. A., 48-2639 (24).
Black,G. A. et al., 50-9809 (43).
Blanco,C., 874 (25); 1138 (24); 1280 (47).
Bonifaz,C., 16 (14).
Boom, B. & D. Beardsley,8449 (cf. 49).
Boom, B. & M. Pacheco,8511 (7).
Boone,W., 1038 (56).
Bourdy,G., 164 (56).
Brade,A. C., 7941, 8195, 9062 (56).
Braga,P. I. S. et al., s.n. BHCB27250 (34).
Brandbyge,J. & E. Asanza,30651, 30885 (55); 31785
(17); 32271 32418 (32).
Brandbyge,J. et al., 33861 (58); 36206 (56).
Brees,E. M., 40 (37).
Breteler,F. J., 4859, 4869 (35).
Brina,A. E., s.n. BHCB39260 (56).
Brunner,D. R. et al., 917 (56).
Buchtien,O., 736 (43); 745, 747, 748 (51); 1986 (17).
Bunting,G. S. et al., 4079 (19).
Calder6n,C. E. et al., 2704 (21).
Camp,W.H.,E-1165 (45).
Campbell,D. G. et al., 435 (47); 14650 (26); P21953,
P21960 (7); P21974 (22).
Campos, J. & L. Campos,3092 (cf. 49).
Campos, J. & 0. Cano,4695 (37).
Campos,J. & S. Nuniez,4669 (42).
Campos,J. et al., 2938 (18); 4480, 4490 (16).
Capucho, P., 565 (Aiouea or Cinnamomum)
Carauta,J. P.P., 6251 (56).
Carauta,J. P.P. & E. Kautsky,3284 (56).
Carauta,J. P.P. et al., 5175 (56).
Cardenas,D. & E. Alvarez,3267 (47).
Cardenas,D. et al., 2645 (48).
Cardona,F., 2784 (25).
Cardozo,A., et al., 1822 (48).
Carnevali,G. & I. Ramirez,2706, 2707 (35).
Carpio,Del C., 2236 (29).
Carpio,Del C. & J. Ruiz, 1620 (35).
Carri6n,A. et al., 531 (24).
Castanieda, R. R., 4068 (cf. 59); 5178 (cf. 44).
Castillo,A., 13, 34 (39); 65 (32); 87 (33); 168 (Ocotea
sp.); 1281, 1282, 1300 (35); 1433, 1538 (47); 1632
(35); 2183, 2225, 2309 (47); 2437, 3653 (35).
Cavalcante,P. & M. Silva, 1782 (35).
Cavalcanti,T. B. et al., 947 (56).
Cazalet,P. C. D. & T. D. Pennington,7629 (8); 7716
(32).
Cer6n, C. E., 882 (32); 1986,2397,2411 (14); 2679 (32);
3704 (50); 7875 (17).
LIST OF EXSICCATAE
Cer6n,C. E. & M. Cer6n,2600 (50); 4520 (53); Cer6n Croat, T. B., 18609 (31); 18846 (29); 18851 (35); 19156,
4626 (57).
Cer6n,C. E. & F. Coello, 3257 (45).
Ceron,C. E. & M. Factos,7649 (32).
Cer6n,C. E. & N. Gallo,4898, 4973 (35).
Cer6n,C. E. & F. Hurtado,3882 (37); 6642 (49).
Cer6n,C. E. & C. Iguago,5562 (14).
Cer6n,C. E. et al., 2090 (48); 8781 (32); 9226 (57).
Chagas,J., s.n. INPA 1313 (47).
Chagas,J. & D. F. Coelho,s.n. INPA3695 (7).
Chanderbali,A. S. et al., 14 (33); 147, 151, 152, 153
(23); 159 (33);206 (23);208, 216 (27);250 (57);252
(33); 269 (23).
Chavez,E., 92 (53).
Churchill,H. W. et al., 3974, 3998 (10).
CidFerreira,C. A., 676 (21); 3936 (35);5563 (22);5682
(7); 7387 (14); 7485 (50); 8900, 9037 (15).
Cid Ferreira,C. A. & J. Lima, 3270 (45); 3590, 3615
(15); 3624 (50).
Cid Ferreira,C. A. et al., 203 (35); 266 (21); 291 (22);
582 (21); 690 (50); 881 (6); 1388 (35); 1563 (14);
1601 (22); 1832 (33); 1884 (28); 1888 (Ocoteasp.);
2200, 2321 (35); 5094 (45); 6033 (43); 6713 (6);
7124 (22); 7227 (15); 7257 (35); 7582 (33); 7652
(23); 7776 (7); 7803 (21); 10031 (45); 10487 (43);
10642 (32).
Clark,H. L., 7587 (35).
Clark,H. L. & G. Gomez,8047 (24).
Clark,H. L. & P. Maquirino,7910 (35); 8324 (33).
Clark,J. L., 2784 (50).
Clark,J. L., et al., 3274 (4).
Clarke,D., 2865 (14); 3695 (27); 4414 (57); 6795 (27);
7342, 7516, 7623, 7975, 8070 (14); 8397 (43).
Claussen,P., 9, 2093, 2454, 18412 (56).
Coelho,D., 29 (22); 160 (21); 3611 (8); s.n. INPA 3206,
s.n. INPA3895 (21); s.n. INPA43578 (43).
Coelho,D. & L. Coelho,26 (7).
Coelho,D. et al., 896 (21).
Coelho,L., 65 (22); 152 (47); 460 (24).
Coelho,L. & D. Coelho,39 (35).
Coelho.L. & A. Miranda,s.n. INPA36033 (6).
Cogollo,A. & J. Brand,380, 394, 415 (4).
Cogollo,A. & C. C. Estrada,290 (Rhodostemonodaphne
antioquensis).
Cogollo,A. & J. G. Ramfrez,4361 (57).
Cogollo, A. et al., 2531 (cf. 49); 3638 (47); 3735 (48);
3928 (47); 5119 (10); 5127 (47); 5128 (50); 5129,
5141 (47); 5167, 5182, 5190, 5195 (38); 6119, 6206
(47); 6556, 7061, 7062 (cf. 49).
Collella,M. et al., 1845 (35).
Command,R., s.n. G 8341/153 (46).
Cordeiro,I., 309 (35).
Cornejo,F. V. & A. Balarezo,2701 (32).
Costa,L. V., s.n. BHCB22314, s.n. BHCB 26334 (56);
s.n. BHCB32676 (34).
Cowan,R. S., 1624 (46).
Cowan,R. S. & J. J. Wurdack,31529 (47).
Cramer,81, (23).
Cremers,C., 3982 (14); 5724 (1); 6404 (14).
129
19680 (29); 19982 (31); 20680 (36); 20771 (31);
58625 (14); 62517 (17); 62737 (cf. 59); 74133 (14).
Croizat, L., 1052 (25); 919B (57).
Cuatrecasas, J. A., 9123 (32); 15292 (50); 16855 (10);
17216 (47).
Cuello, N., 637 (35).
Cunha, O., et al., 206 (24); 216 (6).
Daly, D. C., 5293 (46).
Daly, D. C. et al., 4050 (14); 5086, 5630 (15); 6194 (28);
6795 (57); 7282 (59); 7656, 8104, 9024, 9025 (50);
9293 (58); 9425 (53); 9488 (29).
Damiao, C., 2590 (35).
Daniels, A. G. H. & F. P. Jonker, 847, 990 (14).
Davidse, G., 27575 (24); 27595 (19); 27893 (35).
Davidse, G. & W. D'Arcy, 10097 (10).
Davidse, G. & A. Gonzales, 12971, 14663, 15701 (35).
Davidse, G. et al., 20840 (48).
Davidson, C. & G. Martinelli, 10041 (43).
Davis, E. W., 216 (35).
Delascio, F. & R. Liesner, 13508 (41).
Delgado, L., 582 (35); 835 (47); 868 (35).
Delprete, P. et al., 6141 (35).
Devia, W., 1052 (4).
Devia, W. & F. Prado, 2669 (38).
Diaz, C. & M. Alexiades, 3657 (17); 3661 (56).
Diaz, C. & S. Balde6n, 2304 (50).
Diaz, C. & J. Pereira, 9020 (43).
Diaz, C. et al., 783, 8144 (15); 1079 (31); 7710 (59);
8469 (55).
Diaz, W. & M. Niiio, 289 (48).
Dick, C., 6 (7).
Dik, A., 203 (5); 207 (15); 405 (49), 481 (14); 665, 767
(37).
Dik, A. & J. Andi, 1001 (45).
Dodson, C. et al., 15112 (32).
Donselaar, J. van, 1700, 1789, 2465, 2719 (23); 2037
(26).
Duarte, A. P., 7368 (29).
Ducke, A., 248 (22); 440 (35); 441 (22); 1232, 1728 (6);
1733 (47); 2083 (35); 2478 (13); 2479 (22); 6764
(29); 7185 (35); 8429 (21); 8920 (23); s.n. MG 19238
(22); s.n.MG 15252 (50); s.n. R 2480 (50); s.n. R
11373 (21); s.n. R 17542 (50); s.n. R 18656 (Licaria
sp.); s.n. R 19961, s.n. R 19962 (6); s.n. R 19966
(29); s.n. R 19970 (21); s.n. R 60261 (6).
Ducke, A. & M. Bandeira, s.n. R 162 (56).
Dudley, T. R., 10072 (8); 11493 (17).
Duivenvoorden, 738 (7).
Dus6n, P., 12129 (56).
Duss, P. 226, 575, 2217, 3624, 4093 (46).
Dutra, J., s.n. R. 30975 (56).
Dwyer, J. D. & J. E. Simmons, 9748, 9758A (59).
Edwards, K. S. & T. Roe, 423 (14).
Eggers, H. F. A. von, 332, 403, 740 (46).
Ehringhaus, C. et al., 396 (17).
Eiten, G. et al., 5899 (56).
Ek, R. C. & D. Hammond, 1033 (33).
130
Elger, W. A., 914 (29).
Encarnaci6n, F., 872 (47); 941 (28); 1241 (35); 1261
(15); 26150 (cf. 55).
Ernst, W. R., 1867 (46).
Escobar, L. de et al., 8851 (43).
Espinoza, S. & T. Coba, 369 (59); 378 (37); 588 (50).
Fanshawe, D. B., 1501 (33).
Farney, C. et al., 1744 (35); 2012 (21).
Fernandez, A. 6550 (25); 6887, 7415 (47); 7846 (24);
7968 (47).
Fernandez, J. C. & Susanna, 8414, 8460 (35).
Fernmndez,J. L., 11051 (14).
Fernandez, J. L. et al., 11101 (37).
Fernandez-Casas, J. & J. Molero, 3818, 4205, 5638 (56).
Ferrari, G. & C. E. Benitez de Rojas, 1456 (48).
Ferreira, A. R., 236 (22).
Ferreira, L. V., 237 (22).
Ferreyra, R., 19452 (59).
Feuillet, C., 10187 (1).
Figlioulo, R. & T. Lima, 125 (35).
Figueiredo, C. & I. Riveiro, 553 (cf. 59).
Figueiredo, C. et al., 908 (58).
Filho, H. F. P., 82-114 (22); 82-2 N, 82-3 N, 83-30 N,
83-49 N (50); 82-51 N, 82-55 N (29); 82-7 N (4).
Filho, H. F. P. & L. M. da Silva, 47 (33).
Florschutz, P. A. & P. J. M. Maas, 3069 (14).
Focke, 101 (23).
Folli, D. A., 1546 (56).
Fontella, P. J. et al., 476 (56).
For. Dept. Brit. Guiana, 2183 (27); G233 (57); 2931,
2949 (23).
Forero, E. et al., 2379 (50).
Foster, R. B., 4302 (13); 5406, 5475, 9394, 9688 (17);
5949 (59); 9483 (53); 11339 (56).
Foster, R. B. & B. d' Achille, 11937 (cf. 59).
Foster, R. B. & J. Arce, 11052, 11066 (17).
Foster, R. B. & S. Baldeon, 12815 (59).
Foster, R. B. & H. Beltran, 13050 (57).
Foster, R. B. & C. Janson, 6249 (17).
Foster, R. B. & J. Terborgh, 5093, 6154 (17).
Foster, R. B. & T. Wachter,7286 (53).
Foster, R. B. & P. Wright, 8254 (59); 8368 (17).
Foster, R. B. et al., 3371, 7140 (17); 8925 (55); 11605,
11783 (56); 12483 (58).
Fournet, 4205 (46).
Francisco, s.n. MG. 21114 (47).
Freire, E. & L. Santi, 3294 (50).
Freire, E. et al., 2226 (32).
Freitas, 52 (58); 53 (30).
Freitas & Mota, 467 (15).
Freitas, C. A. A. et al., 239 (32).
Froehner, C., 67 (43); 171 (56).
Fr6es, R. L. de, 20899 (50); 20960 (35); 21022 (30);
21211 (7); 21243 (47); 22620 (24); 23875 (29);
23966, 26368 (35); 28878 (6); 34834 (35).
Funk, V., 6102 (43).
FLORANEOTROPICA
Galdames,C. et al., 1362 (56).
Galeano,G. et al., 1001, 1019, 1024 (35); 1065 (36);
1120 (35).
Garcia,L. et al., 72, 103, 380 (14).
Garcia-Barriga, H., 13703 (47); 13944 (Rhodostemono-
daphne sp.).
Gardner,349, 5595 (56).
Gentry,A. H., 28964 (35);43479 (17);48990,49120 (6).
Gentry,A. H. & J. Aronson,25184 (36); 25188 (15).
Gentry,A. H. & S. Estensoro,70632 (8).
Gentry,A. H. & M. Fallen,17803 (38).
Gentry,A. H. & N. Jaramillo,58022 (15);65576 (cf. 59).
Gentry,A. H. & P. Nuiiez,69332, 69392 (59).
Gentry,A. H. & R. Ortiz,74245 (29).
Gentry,A. H. & E. Renterfa,23982, 23983 (10).
Gentry,A. H. & J. Revilla, 16514 (35); 16539 (47);
16678 (35); 20368, 20507 (15); 20841 (35); 69177
(28).
Gentry,A. H. & D. N. Smith,45169 (44).
Gentry,A. H. & J. Solomon,44524 (51).
Gentry,A. H. & B. Stein,46329 (27).
Gentry,A. H. & K. Young,31842 (15).
Gentry,A. H. et al., 2533 (15); 7695 (17); 15865 (36);
18525 (20); 19636 (58); 21944 (55); 21980 (7);
22365 (31); 24873 (20); 25797 (35); 25983 (39);
26850 (17); 27088 (59); 27145 (17); 28884 (13);
28964 (35); 31118 (Ocoteasp.); 31649 (15); 37107
(39); 37246 (cf. 55); 38124 (15); 39301 (39); 39734
(31); 42743, 42763, 42779, 42925 (15); 43074 (cf.
8); 43779, 45597, 45598, 45764, 45776, 51521 (15);
54550 (29);56205 (cf. 59); 56281 (7); 59300,59347,
59377 (56); 62990 (26); 68712 (cf. 59); 68974 (59);
76844 (14); 78108 (17).
Gillespie,L. & H. Persaud,1557 (23).
Gillespie,L. et al., 1691, 1851 (27).
Giovianni,F., 32-A (35).
Glasgo,A. & V. Slane,21 (46).
Glaziou,A., 3092 (56); 7781, 8093 (34); 12120 (56);
14210 (35); 14212 (21); 16315 (56); 18451 (34);
19795,20459, 22056, 97811 (56); s.n. (34).
Gleason,H. A., 356 (23).
Goeldi, A., s.n. HAMP 3902 (type of Endlicheria goel-
diana);s.n. MG 3911 (8).
G6mez,R. et al., 649 (35).
Gonggrijp,3692 (47).
Gonggrijp& Stahel,5607 (14).
Goodland,R., 4504 (56).
Goulding,M., 41 (35); 1379 (22).
Graham,J. & J. V. Schunke,365 (35); 1099 (17).
Grandez,C., 502 (29).
Grandez,C. & A. Chiquispama, 894 (35); 912 (15); 919
(29); 982 (50); 996 (35).
Grandez,C. & G. Criollo, 1582 (cf. 59).
Grandez,C. & N. Jaramillo,794 (13); 817 (14).
Grandez,C. et al., 537 (15); 1526(58); 1611, 1612(50);
1921 (7); 4228, 4233 (13); 5512 (45).
Granville,J.-J.de et al., 384 (14); 1106 (1); 1331 (14);
LIST OF EXSICCATAE
1448(1); 1451, 1462,2133, 3145, 3723, 3739, 3979,
4949, 5547, 7406, 7675, 7949, 8380, 9005, 9476
(14); 10538(26); 10794(14); 10816(1); 12329(14);
B.3713, B.4356, B.5496 (1).
Grayum,M. et al., 9153 (15).
Grenand,2078 (14); 2870 (1).
Grenand& M. F. Pr6vost,2089 (14).
Grewalet al., 384 (23).
Grijalva,A. et al., 243 (50); 645 (15).
Guanchez,F., 102, 166, 388, 988 (47); 1016 (25); 2154
(41).
Gudnchez,F. & A. Brown,2753 (47).
Gudiino,E. & N. Andi, 2064 (17).
Gudifio,E. & S. Papa,1902 (17).
Gudiiio,E. et al., 871, 1041 (55).
Guedes, M., s.n. R 20065, s.n. R 20081 (43).
Guerra,C., 646b (10).
Guilding,L., s.n. K H1/45/97-199(46).
Guillaumet,J. L. et al., 5707, 5716 (22).
Guillen,R. & R. Chore,2918, 3283, 3292 (35); 3814
(24).
Guillen,R. & C. Medina,3781 (35).
Guillen,R. & V. Roca, 3313 (35).
Guillen,R. et al., 3115, 4074 (35); 4454 (24).
Gurgel,15140 (56).
Hahn,L., 1068 (46).
Hahn,M., 168, 983, 3184 (46).
Hahn,W., 2088, 2591 (56).
Hahn,W. et al., 1347 (56); 5795 (33).
Halloy,S. et al., 4319 (35).
Hamilton,C. & H. Stockwell,2905 (56).
Hammel,B., 1858 (48); 13518 (10).
Hammel,B. et al., 21557 (1).
Hartshorn, G. & J. Quijano,2940 (55).
Hartshorn,G. et al., 2600 (32);2606 (4);2689 (59);2824
(53); 2827 (4); 2916 (8); 2971 (55).
Hassler,E., 8935 (56).
Hatschbach,G., 14075, 14366, 39144, 42573, 46151
(56).
Hatschbach,G. & E. Barbosa,59364 (56).
Hatschbach,G. & A. C. Cervi,48903 (56).
Hatschbach,G. & 0. Guimaraes,14678 (56).
Hatschbach,G. & R. Kummrow,52332 (56).
Hatschbach,G. & J. M. Silva,48651, 48978 (56);52169
(34); 61556 (56).
Hatschbach,G. et al., 13246, 51226 (56); 57930 (34);
58562, 61580 (56).
Haught,O., 1414 (57).
Helme,N., 351 (8).
Hemmendorff, E., 457 (34).
Henao,J. E., 9, 21 (cf. 49); 152 (15); 281 (cf. 49).
Henderson,A. et al., 342 (7); 458 (35).
Henkel,T., 4946 (14).
Henkel,T. & M. Chin,601 (23).
Henkel,T. & R. Williams,687, 2091 (23).
Henkel,T. et al., 1771 (23); 3430, 3713 (27); 5087 (23).
131
Heringer,E. P., 16026 (34); 18203 (56); 18530, 18580
(34).
Heringer,E. P. & G. Eiten, 15083, 15154 (34).
Heringer,E. P. et al., 1942, 3059, 3062, 3187, 3210,
3211, 3233, 5081, 5184, 5513, 6211, 6245, 6404,
7174 (56).
Herrera,G., 4967, 5031 (15).
Herrera,H. et al., 1057 (10).
Hetschko,s.n. R. 30983 (56).
Heyligers,P.C., 485 (26).
Hill, S. R., 13066 (35).
Hill, S. R., 24166 (46).
Hodge,W.H., 400, 1123, 1317, 2252, 3843 (46).
Hodge,W.H. & B. T. Hodge,2760 (46).
Hoehne,F. C., 3517 (56).
Hoffman,B. & D. Gopaul,365 (27).
Hoffman,B. & H. Jacobs,1072 (27).
Hoffman,B. et al., 992 (27); 1358, 2029, 2382, 2400
(23).
Hohenkerk,L. S., 837 (23).
Holm-Nielsen,L. et al., 19908, 19914,20029 (35).
Hopkins,M. J. G., 1595 (6).
Hopkins,M. J. G. et al., 633 (27); 1592 (32).
Homer,C. et al., 233 (25).
Hostmann,F. W. & A. Kappler,1163 (23).
Howard,R. A., 11145 (46).
Howard,R. A. & B. R. Howard,18005 (46).
Howard,R. A. & E. S. Howard,19541 (46).
Hoyos, S. E. & J. J. Hemrnndez, 292 (Rhodostemonoda-
phne antioquensis).
Huashikat,V. 46 (59); 76, 436 (53); 443, 781 (32); 836
(53); 1475 (cf. 59); 1488 (32).
Huber,J., s.n. MG 513, MG 9431 (6); s.n. R 18359(3).
Huber,O., 13261 (41).
Huber,0. & S. Tillett,2975 (21).
Hunt,D. R. & J. F. Ramos,6004 (52).
Hurtado,F., 2654 (17); 2979 (32).
Hurtado,F. & A. Alvarado,963 (49).
Hurtado,F. & D. Neill, 1467, 1477, 1509 (55).
Idrobo,J. & R. E. Schultes,944 (14); 1187 (33).
Imray,J., 185, 194, 199, 334 (46).
Irwin,H. S., 2688 (56).
Irwin,H. S. & L. Y. Th. Westra,47706 (1).
Irwin, H. S. et al., 9097, 13077, 23151 (56); 16777,
16993, 17122, 17291(52); 47273, 47504, 47933 (1);
48290 (47); 54744, 54747, 54946, 54975, 55132
(14); 55631 (57).
ItaipuBinacional,129, 924 (56).
Jansen-Jacobs, M. et al., 171, 261 (27); 1118 (23); 2699
(27); 3085 (57); 3311, 3394 (27); 4413 (23); 4575
(57); 4599 (23); 5101 (14); 5374 (27).
Jaramillo,J., 8349 (8); 8372 (50).
Jaramillo,J. & E. Grijalva,11582 (cf. 59).
Jaramillo,J. et al., 31147 (7).
Jaramillo,N. & S. Katip,790 (53).
Jaramillo,N. et al., 894 (55); 1084 (59); 1159 (53).
132 FLORANEOTROPICA

Jaramillo,R. et al., 7103 (cf. 45). (35); 7466 (19); 8479, 8640, 8665 (35); 18657,
Jardim,A. et al., 594 (59); 764 (50); 2576 (43). 18665, 18671, 19092 (47); 24576 (25).
Jarvenpaa, T., 27 (4). Liesner, R. L. & G. Carnevali, 22292, 22414 (41).
Jelski,165, 196 (43). Liesner, R. L. & H. Clark, 8963 (47).
Jenman,G. S. 1702 (Ocoteasp.);4127, 5321, 5823 (23). Liesner, R. L. & F. Delascio, 22008 (41).
Jervise,s.n. K H842/98-8 (11). Liesner, R. L. & B. Holst, 21854 (47).
Jimenez,B. & G. Manrn,1331 (56). Liesner, R. L. & G. Morillo, 13929 (25).
Jimenez,B. et al., 1809 (56). Lima, J., 783 (25).
Jones,J. & C. Davidson,9639 (58). Lima, J. & S. Sousa, 230 (50).
Jorge,H & R. Torres,1270 (cf. 49). Lindeman, J. C., 379 (47); 2741 (14); 4509 (47); 4590
J0rgensen,P.M., 4462 (56). (23); 4787 (26); 5272 (23); 5381 (cf. 32); 5601 (23);
Jouvin,P. P.,472 (56). 5943, 6056 (26); 6651 (23).
Lindeman, J. C. & J. H. de Haas, 747 (56).
Karsten,s.n. (4). Linden, 429 (11).
Kayap,R., 354 (8); 1374 (55). Lisb6a, P. & 0. P. Monteiro, 954 (7).
Killeen, T., 3832 (Rhodostemonodaphne juruensis); Little, E. L. Jr., 8878, 8886, 40105, 40141 (56).
3880 (50). Little, E. L. Jr. & R. G. Dixon, 21035 (15).
Killeen,T. & K. Smith,3634 (56). Little, E. L. Jr. & R. R. Little, 8259, 8308, 9565 (35).
Killip, E. P. & A. C. Smith,25975, 26077 (55); 27192 Little, E. L. Jr. et al., 221 (44).
(35); 28050 (4); 28291 (14); 28400 (Rhodostemon- LLB (Landsbosbeheer, Suriname), 12570 (37).
odaphne debilis); 29870 (13). Lleras, E. et al., P16960 (53).
Kirkbride,J. H. Jr.,2387 (56). Lohmann, L. G., 110, 343 (35).
Klein,R., 7, 21, 571 (56). Lombardi, J. A., 1987 (34).
Klug,G., 26, 161 (31); 204 (13); 272, 273 (7); 411, 621, Lopes, M. et al., 124 (21).
703 (13);728 (35); 1264, 1403(20); 1884, 1904(53); L6pez, A. et al., 8635 (35).
2253 (30); 2313 (14); 2833 (56); 2958 (14); 3187, Lorea, F. G., 5581 (56).
3745 (28). Lorea, F. G. & M. Gorgulho, 5579, 5580 (56).
Knab-Vispo,C. & G. Rodriguez,479 (25). Loureiro, A. et al., 742, 1320 (35); 5739 (47); s.n. INPA
Knapp,S. & P. Alcom, 7778 (49). 35784 (7); s.n. INPA 37822, s.n. INPA 37846 (22);
Knapp,S. & J. Mallet,653 (14); 2829 (27); 7024 (59); s.n. INPA 37949 (7); s.n. INPA 38000, s.n. INPA
7110, 8557 (14). 38870, s.n. INPA 38878 (35); s.n. INPA 39526 (22);
Knapp,S. et al., 4496 (56). s.n. INPA 47996 (21, 22); s.n. INPA 48137 (47).
KrollSaldania, B., 134, 326 (8); 560, 689, 795 (56). Lugo, H. S., 3906, 3925, 4125 (32); 4188, 4241, 4302,
Krukoff,B. A., 1565 (15); 1933 (14); 4714 (15); 4717, 4394, 4450 (45).
4767 (17); 4775 (Rhodostemonodaphne juruensis); Luna, M. L., 112 (44).
4855 (17);4932 (8); 4943 (53);4959 (13);5023 (59); Luteyn, J. L. et al., 8541 (59); 8586 (cf. 59); 8680, 9004,
5030 (14); 5156 (15); 5204 (17); 5279 (50); 5281 9019 (17).
(15); 5780 (3); 6116 (50);6300 (58);6406 (50);7206
(32); 7265 (22); 8096 (24); 8293 (29); 10787 (17); Maas, P. J. M. & J. A. Tawjoeran, s.n. LLB 10928 (23).
11103 (43); 11262 (51). Maas, P. J. M. & L. Y. Th. Westra, 3815 (23).
Kubitzki,K., 75-34 (29); 87-22 (22). Maas, P. J. M. et al., 4599, 6637 (35); 6798 (32); 7136,
Kubitzki,K. & H. Poppendieck,79-29 (21). 7166, 7215, 7308 (27); P13058 (40); P13307 (58).
Kubitzki,K. et al., 79-172 (19). Machado Nunes, G., 318 (34).
Kuhlmann,J. G., 171 (35); 3376 (27); s.n. R 11966, R Maciel, U. N. & C. S. Rosairio, 1559 (24).
136580 (56); s.n. R 18360 (50); s.n. R 20028 (53); Mackenzie, C. A. et al., INPA/WWF 2107.388 (2).
s.n. R 20036 (29). Madriiain, S., 710 (35).
Kurtz,B. C. et al., 80, 87 (56). Madrifian, S. & C. Barbosa, 936, 948, 953 (41).
Kvist,L. P. & L. FreitasA. 706, 736, 808, 921 (35). Maguire, B., 24538 (14); 24712 (37, 47); 24898a (37);
40775, 40790 (14).
Labroy,s.n. (35). Maguire, B. & D. B. Fanshawe, 23481 (23); 32463 (47).
Lanjuow,J. & J. Lindeman,2367 (14). Maguire, B. & L. Politi, 27384, 28159, 28380 (47);
Larpin, D., 851 (14). 28428 (32).
Lasser,T., 1881 (57); 2036 (48). Maguire, B. & J. J. Wurdack, 34727 (25); 34859 (35).
Lasser,T. & C. K. Allen, 13 (48). Maguire, B. et al., 29436 (57); 30601 (21); 36314 (47);
Leng,H., 128 (23). 42244 (41); 42521 (Aiouea maguireana); 46713 (23);
Lepschda Cunha,N. M. et al., 391 (2). 53519 (47); 56091 (52); 56624 (22); 60121 (47).
Level, S., 127 (24). Maia, L. A. et al., 208 (35); 344 (22); 430, 438, 562 (24).
Lewis, G. P., 1488 (25). Mallorguim, R., 16 (56).
Liesner, R. L., 3961, 4183, 6237 (35); 6657 (32); 7137 Malme, G., 755 (56).
LIST OF EXSICCATAE
Manara, B., s.n. VEN 172708 (48).
Manso & Lhotzky, 84 (52).
Marcano-Berti, L., 361 (26); 2528 (25); 2581 (47).
March, W. T.?,s.n. (23).
Marin, E., 438 (19); 981 (24).
Marfn, G. & B. Jimenez, 322 (56).
Martinelli, G., 151 (56).
Martinelli, G. & D. Sucre, 200 (56).
Martinelli, G. et al., 23 (34).
Marulanda, 0. & S. Marquez, 1108-HUA 88921 (35);
1108-HUA 89052, 1536 (58).
Mathias, M. E. & D. Taylor, 5514 (29); 5532 (58); 5957
(17).
Matos, F. et al., 207 (50).
Matuda, E., 5153 (Beilschmiedia zapoteoides).
McDaniel, F. & L. Santiago, 2539 (58).
McDaniel, F. & T. Wilkes, 2483 (29).
McDaniel, S. & M. Rimachi, 18838 (58); 20744 (36).
McDowell, T., 2663 (47).
McDowell, T. & S. Tiwari, 1861 (27).
McPherson, G., 8493 (12); 10000, 10444, 11310, 11582,
11936 (48).
McPherson, G. & M. Merello, 8131 (15).
Mecenas, V. V. & F. Q. Leite, 92 (56).
Meier, W. & 0. Kunert, 4632 (56).
Meier, W. & K. Walter-Weisbeck, 2586 (48).
Meier, W. et al., 2756 (25).
Melinon, 204, 216, 227, 551 (1).
Mello, F., 24 (35); s.n. INPA 78 (47); s.n. INPA 3526
(6).
Mello, F. & L. F. Coelho, s.n. INPA 4167 (47).
Mello, F. & M. Emmerich, 2934 (56).
Mello, F. & J. Ribamar, s.n. INPA 58316 (43).
Mendez, P. et al., 22 (15).
Mendonqa, R. C. & F. C. Silva, 61 (56).
Mennaga, A. M. W., 86 (14).
Mexia,Y., 5025, 5091, 5138 (34); 6268a (53); 6329 (56).
Meyer, G., 146 (56).
Miers, J., 4270 (56).
Miller, J. S. & G. Davidse, 1649 (24).
Miller, J. S. et al., 638 (55).
Miller, R., 524 (35).
Milliken, W. & S. Bowles, 304 (25).
Milliken, W. et al., 142, 439 (27); 524 (35).
Miralha, J. M. S., 78 (24).
Miralha, J. M. S. et al., s.n., Araca Inventory BO-6-611
(24).
Miranda, S. I., 20 (59); 1000 (27).
Molas, L., 771, 820 (56).
Molino, J.-F., 1663 (14).
Monsalve, M., 1975 (38).
Monteiro, 0. P., 1246 (21); 1252 (32).
Monteiro, 0. P. & J. Lima, 145 (7).
Moore, S., 518 (52).
Moraes, P. L. R. de, 122, 123, 403, 561 (56).
Morales, J. F. et al., 2083 (15).
Morawetz, W. & B. Wallnofer, 11-12888 (28); 13-9288
(48).
Moretti, C., 1196 (14); 1443 (8).
133
Mori,S., 7708 (56); 23935 (14).
Mori,S. & C. Gracie,21867, 21962, 22473, 22492 (35).
Mori, S. & J. Kallunki,3472 (10).
Mori,S. & T. Pennington,18041 (1).
Mori,S. et al., 6476 (10); 9097 (53); 9178 (Ocoteasp.);
14910, 14995 (1); 15059, 15649 (14); 17170 (57);
20614 (7); 20927 (26); 22268 (1); 22979 (14).
Morillo,G. & R. Liesner,8818, 8936 (25).
Moscheta,I. & D. dos Saldovino,3 (56).
Mota, C. D. A., 685 (28); s.n. INPA 60364 (55); s.n.
INPA60608 (21).
Mota,C. D. A. & 0. P. Monteiro,s.n. INPA61281 (21).
Muinera, M. L. E., 13 (cf. 49).
Mutchnick,P., 687, 1243, 1303 (23); 1464 (33).
Nadruz,M. et al., 573 (56).
NascimentoJ. R. & E. C. da Pereira,606 (cf. 53).
Nascimento,0. C., 386 (6); 685 (32); 826 (19).
Nee, M., 34692 (43); 39040 (56); 39834 (17).
Nee, M. & L. Bohs, 49533 (17).
Neill, D., 7152 (53); 7164 (cf. 59); 7266 (14);7514 (15);
8730 (53); 9842 (14); 9886 (32); 10095 (17); 11028
(53).
Neill, D. & F. Hurtado,8807 (55).
Neill, D. & W. Palacios,9618 (45).
Neill, D. & W. Rojas,9950 (17).
Neill, D. et al., 8094 (49); 8475 (48); 8953 (49); 10168
(15); 12614, 12615 (cf. 49); 12616 (42).
Nelson,B. W. & J. F. Lima,P21108 (21).
Nevers,G. de & D. Cavagnaro,4795 (56).
Nevers,G. de & H. Herrera,4186 (10).
Nevers,G. de et al., 5339 (10).
Nicholls,H. A., s.n. (46).
Nicholson,D. H., 4237 (46).
Nunhez,P., 5907, 6202, 6260 (17); 11195 (43); 13986
(15).
Nunfez,P. & C. Munioz,5324 (14).
Nufiez,P.et al., 8017 (17); 10804(4); 14261(17); 14373
(cf. 59); 14429 (56).
Oldeman,R. A. A., 79, 2448, B. 2993, B.3580, T-35, T-
55 (47); 1251, 2866, B. 1881, B. 817, B. 3996, B
4034 (14); T-23 (1).
Oldenburger et al., 1221 (cf. 33).
Oliveira,F. C. A. et al., 770 (34).
Oliveira,P.I., 256, 926 (56).
0llgaard,B. et al., 34616 (17); 57630, 57623A (10).
Ortiz,R. & J. Vargas,206 (53).
Ortiz,R. et al., 122, 124 (58).
Palacios,W., 1005(14); 1370(53); 1423(59);3260 (14);
3496, 3496 (55); 4044 (15); 4212, 4386 (57); 4397
(28); 4413, 4473 (4); 4747 (49); 5482 (15); 5597,
5663 (53); 6732 (51); 7500 (28); 10294 (33); 11001
(49); 11268 (17); 13750 (10); 13897, 13906 (15);
13936 (33).
Palacios,W. & E. Freire,5119 (8).
Palacios,W. & C. Iguago,4541 (49); 4696 (8).
Palacios,W. & D. Neill, 1289 (50); 1516 (53); 6522,
6531 (55); 6572, 6577, 6669, 6688 (45).
134
Palacios,W. & G. Tipaz,11074 (45).
Palacios,W. & M. Tirado,11165 (15); 13298, 13302
(57); 13310 (45); 13342 (51); 13425, 13439, 13459
(42).
Palacios,W.et al., 1030(56); 1183(53);4936, 7726 (8);
7804, 7884 (58); 7879 (15); 8027, 8042 (59); 8142,
8144 (35); 8164, 8165, 8399 (37); 8492 (45); 8585
(57); 8644 (15); 8840 (57); 8845 (7); 8938, 8940
(48); 8963, 9492 (15).
Paniagua,N. & R. Foster,690 (35).
Pariona,W., 399 (32).
Pariona,W. & J. Ruiz,974 (32).
Pariona,W. & A. Sebastian,25 (32); 46 (4).
Patris, s.n. G8341107, G8341109, G8341110,
G8341111,G8341112,G8341113,G8341117(26).
PaulinoR. & F.Victorio,s.n. R 2322 (typeof Endlicheria
balsamea).
Pavon,J., 506 (17); s.n. G 8341 (17).
Pearce,R., s.n. K 52, MO 1611827(51);s.n. K 59, K 60
(16).
Peckolt,115 (56).
Peixoto,A. et al., 3466, 3504 (56).
Pennington,R. T. & J. Ramos,P22768 (43).
Pennington,R. T. et al., 80 (50).
Pereira,E., 5356 (56).
Perry,A., 999 (43).
Persaud,A. C., 49, 59 (23).
Pesha,V., 68 (50).
Pfrommer,A., 6 (56).
Philipson,W.R. & J. M. Idrobo,2013 (32).
Philipson,W.R. et al., 2192 (15).
Phillips,0. & P. Nuinez,52 (58).
Phillips,0. et al., 468 (15).
Pierre,L. L. Jr.& V. Slane,357 (46).
PilarFranco,R. et al., 5337 (37); 5499 (51).
Pinkus,263 (47).
Pinto,F. & P. Q. Bernal,1624, 1642 (cf. 59).
Pipoly,J. J., 10316 (47).
Pipoly,J. J. & G. Samuels,6868 (24).
Pipoly,J. J. et al., 10607 (33); 12202 (13); 12633 (7);
12727 (20); 13554 (31); 14321 (28); 16280 (15);
16545, 16568, 16718, 17080(47); 17315, 17453(cf.
49).
Pires,J. M. & R. P. Belem, 12405 (14).
Pires,J. M. & G. A. Black,112(6); 1055(13); 1229(24).
PiresJ. M. & P.B. Cavalcante,52649 (47); 52676 (14).
Pires,J. M. & N. T. Silva, 10647, 10658 (6).
Pires, J. M. et al., 9235 (56); 13887 (35); 16821 (25);
50825 (14); 51168 (1); 51599 (57).
Pires,0. & D. Coelho, 175 (2).
Pittier,H., 8017 (56).
Pizziolo,W., 280 (56).
Plotkin,M., 159 (47).
Plowman,T. & E. W. Davis, 5098 (17).
Plowman,T. & H. C. de Lima, 12895 (56).
Plowman,T. & J. V. Schunke,11575 (14).
Plowman,T. et al., 6693, 7013 (36).
Poeppig,1520, 2298 (56); 2552 (35).
Pohl, 5611 (56).
FLORANEOTROPICA
Poiteau,M., s.n. G 122, G 123, G 124, G 128, LE 2811,
NY 99497, P 162 (14).
Polanco,R., 222 (56).
Pollard,79 (27).
Poncy,O., 89 (14).
Poole, J. M., 1717 (35); 1972 (19).
Prance,G. T., 30162 (22).
Prance,G .T. & D. F. Coelho, 17581 (7).
Prance,G. T. & T. D. Pennington,2004 (cf. Rhodoste-
monodaphne tumucumaquensis Madriniain).
Prance,G. T. & A. E. Prance,14774 (35).
Prance,G. T. & N. T. Silva, 59385 (24).
Prance,G. T. et al., 1343(OcoteadiffusavanderWerff);
1563 (14); 2269, 2688 (47); 2972 (35); 3008 (7);
3180 (47); 3563 (29); 3817 (21); 3886 (6); 3919 (7);
4211 (27); 4282 (25); 4678 (21); 5290, 5295 (35);
5327 (22); 6182 (29); 6834 (35); 7744 (29); 7810
(17); 8343, 8892 (6); 9961 (Ocoteasp.); 11069(25);
11511 (24); 11568, 11596, 11737 (35); 12102 (7);
12161 (cf. 59); 12214 (17); 12233 (55); 12436 (7);
12501 (17); 12514 (cf. 59); 12563 (45); 12633 (13);
13391, 13442, 13485, 13695(58); 13888(32); 13987
(14); 14013 (50); 14150 (40); 14498 (4); 14616 (E.
metallicaaff sp. 1); 14671 (24); 15035, 15725 (47);
15817(6); 16527(50); 16695(58); 17742(7); 17785
(22); 18032, 18037 (7); 24626 (35); 29998 (22).
Prevost,M. F. & Grenand,1958 (14).
Proctor,G. R., 18205, 21575, 21585 (46).
Proenqa,C., 863 (56).
Pulle, A., 223 (14).
Qazarin,P.R., 11 (29).
Quelal, C. et al., 704 (Rhodostemonodaphnejuruensis).
Quentin,R. P., 1091 (46).
Quifiones,L., 1419 (14).
Raimondi,4098 (43).
Ramage,G. A., s.n. (46).
Rambo,B., 43408, 45269, 63615 (56).
Ramfrez,J. G. & D. Cardenas,1016 (8).
Ramfrez,J. G. et al., 4886 (32).
Ramirez,R. C., 1068 (29).
Ramos,J., 450 (45); s.n. INPA54118 (29); s.n. INPA
62184 (47).
Ramos,J. & D. Coelho,693 (32).
Ramos,J. & G. Mota, 192, 332 (7); 350 (45).
Ramos,J. & R. Sousa,68 (52).
Ramos,J. et al., 684 (Ocoteasp.);s.n. INPA62154 (21).
Ratter,J. A. et al., 840, 1278, 1944, 5590, 5634, 5713
(25).
Ravedutti,C. et al., 7 (56).
Regnell,A. F., s.n. R. 30958, R. 30976 (56).
Reif, 1785 (56).
Reitz,P.R., 3098, 4423, C153 (56).
Reitz,P.R. & R. Klein,286, 931, 1140, 1676, 1833(56).
Renterfa,E. et al., 2821 (8).
Restrepo,D. & A. Matapi,376, 466 (7).
Revilla,J., 210 (29); 410 (35); 460 (29); 625 (35); 636
(13);675, 695 (35);798 (29); 885 (35); 889 (29);945
LIST OF EXSICCATAE
(7); 1153(29); 1224(28);2111 (35); 3627 (47);3772
(29).
Revilla,J. & J. Forero,4175, INPA82158 (22).
Revilla,J. et al., 2472 (35).
Reynel, C. & E. Meneses, 5089 (50); 5115 (8); 5125
(50).
Reynel,C. et al., 5244 (28).
Ribeira,R. & I. Silva, 2061 (56).
Ribeiro,J. E. L. S. et al., 885 (47).
Ribeiro,R., 544 (56).
Ribeiro,R. & W. L. Araujo,130 (56).
Ribeiro,R. et al., 1021 (cf. 56).
Richards,P. W., 6871 (52).
Riedel,L., 1142 (56); 1374, 1451 (35).
Rimachi,M. Y, 2533 (29); 2590 (58); 3226 (31); 3506
(36); 4286 (35); 4397 (55); 4644 (7); 7739, 9113
(20); 9906 (29); 10391 (28); 10402, 10532 (20);
11199 (13); 11305, 12263 (31); 12301 (50); 12302
(29).
Ritter,N., 1706 (17).
Robert,A., 4856 (56).
Robertson,K. R. & D. F. Austin,90, 100 (35).
Rodrfgues,E. & P.M. Reyes, 1807 (44).
Rodrigues,H., 2545 (24).
Rodrigues,R. S., s.n. MG 8265, s.n. MG 8811 (43).
Rodrigues,W., 374 (47); 508 (21); 552 (53); 574 (47);
1064 (32); 1596 (35); 1972 (22); 5389 (32); 6715
(22); 7028 (28); 7977 (cf. Ocotea rufovestita);8780
(6); 8781 (7); 8835 (35).
Rodrigues,W. & J. Chagas,1259 (21); 2350 (47).
Rodrigues,W. & D. Coelho,461 (21); 2473 (22); 2588,
2743, 2783, 3033, 4558A (21); 5278 (35).
Rodrigues,W. & L. Coelho,2564 (21); 5306 (7).
Rodrigues,W. & J. Lima,2605 (35); 4124 (21).
Rodrigues,W. & A. Loureiro,5926 (33); 7075 (47);
9454 (2).
Rodrigues,W. & F. Mello, 5355 (35); 7760 (29).
Rodrigues,W.et al., 114 (56);941 (55); 1980(21);8523
(2); 10494 (21).
Rojas,R. et al., 91 (57); 172 (55); 506 (32).
Rollet,B., 1672 (46).
Romero,G. A. & E. Melgueiro,2010 (35); 2129 (21).
Romoleroux,K., 2122 (cf. 59).
Romoleroux,K. & R. Foster,1908 (15).
Romoleroux,K. et al., 2567 (48); 3502 (50).
Rosales,J. et al., 1245 (25).
Rosario,C. S. et al., 1032 (6).
Rosas,A. et al., 216 (43).
Rubio,D., 8 (45); 11, 12, 40 (55); 115 (13); 306 (14).
Rubio,D. & A. Alvarado,2395 (49).
Rubio,D. & E. Gudiiio,199 (13).
Rubio,D. et al., 454 (Rhodostemonodaphnejuruensis);
920 (15).
Rudas,A. et al., 2116 (50); 2152 (35); 2262, 2288 (50);
2290 (29); 2504 (15); 2588 (50); 2610, 2642, 2648
(37); 3601 (15); 4057 (57); 5817 (15).
Rueda,R., 445 (29); 1116 (57).
Rueda,R. & J. C. Ruiz, 679 (35).
Ruiz, H. & J. Pav6n,s.n. (17).
135
Rufz,J. C., 1436, 1466a, 1515 (35).
Ruiz,J. C. & H. Murphy,228 (13); 230 (31).
Ruiz,J. C. et al., 6285 (29).
Rusby,H. H., 572, 2671 (17).
Ruzz,J., 1409 (15).
Sabatier,D. G., 1087 (47); 2365 (26); 3501 (14).
Sabatier,D. G. & M. F. Prevost,3195 (33); 3819, 4073
(26).
Salino,A., 3274 (56).
Samaniego,A. V. & A. C. Vivar,90 (44).
Sampaio,A., 2223 (56).
Sanchez,I. V. & M. Dillon, 8687 (43); 8916, 8925 (28).
Sanchez,I. V. et al., 9345 (E. metallicaaff. sp. 2); 9575
(56); 29363 (8).
Sanchez,M. et al., 1870 (22).
Sanoja,E., 2522 (25).
Santoset al., 39 (50).
Santos,M. R., 563 (22).
Saraiva,R. S. & L. de Lima, 1523 (43).
Sastre,C ., 1488 (1); 1619 (14); 1644 (1); 1651 (14).
Sastre,C. & F. Sastre,2046 (46).
Schinini,A. & G. C. Marmori,27009 (56).
Schomburgk, R., 171(240)(23); 475/801(27); 784 (35).
Schrainer,s.n. R 61160 (56).
Schultes,R. E., 3337 (cf. 59); 7149 (35).
Schultes,R. E. & I. Cabrera,12423 (32); 12658, 12664
(35); 12735 (32); 13067, 13073, 13076, 13236,
13241 (35); 13566 (13); 13849 (24); 14125 (13);
14356 (21); 14606 (47); 14903 (45); 15566 (35);
16230(13); 17033, 17898,17898, 18421(35); 19540
(39); 20003 (21); 22578, 24200 (35).
Schultes,R. E. & F. L6pez,9954 (24).
Schultes,R. et al., 24174, 24200 (35).
Schunke,J. V., 1587, 1604, 1617 (56); 2570 (59); 2924
(56); 3329 (17); 3474, 5679 (14); 4532, 4766 (17);
5060 (28); 6231 (15); 6726 (Rhodostemonodaphne
juruensis);6957 (28); 7053 (17); 7245, 8069 (28);
9242 (56); 10304 (28); 10657 (56); 12481 (17).
Schwacke,C. A. W., 95 (56); 351 (22); 617 (35); s.n. R
30973 (34); s.n. R 61142 (56).
Seemann,B. C., 1094 (10).
SEF,8740 (50); 9247 (32).
Seidel, R. et al., 6557 (58).
Sellow,433, 1166, 2244, B 383, c 418 (56).
Serato,A., 305 (56).
Setz, E., F779 (43).
Shiki,D., RBAE359(14).
Shillingford,C. A., 517 (46).
Sidney,1308 (52).
Sieber,F. W., 175 (46).
Silva, A., 135, 282 (6); 309 (43).
Silva, A. S. L. da et al., 635 (14).
Silva, E. S., 185 (24).
Silva,J. A., 181 (24).
Silva,J. M. & L. M. Abe, 2883 (56).
Silva,J. M. & E. Barbosa,2435 (56).
Silva, J. M. et al., s.n. INPA/WWF2303.4840(28).
Silva, M. F., 788, 823, 824 (35); 960, 976 (22).
136
Silva, M. F. & R. Souza, 2546 (OcoteanigrescensVi-
centini);2630 (Ocoteadiffusavan derWerff).
Silva, M. F. et al., 381, 680 (35); 808 (47); 1707 (19);
1752 (35).
Silva,M. G., 1042 (6); 6059, 6078 (50); 7148 (7).
Silva,N. T. da, 38 (6); 968 (43); 57857 (6).
Silveira,M. et al., 468 (29);943 (28); 1499(cf. 59); 1555
(15); 1565 (50).
Simonis,J. E. et al., 121 (56).
Simpson,D. R., 782 (20).
Slane,V. & P. Acevedo,287 (46).
Slane,V. & B. Rollet,514 (46).
Smith,A. C., 2221, 3129, 3376 (27); 10356 (46).
Smith,D. N., 4491 (44); 5318 (56); 6764 (50).
Smith,D. N. & S. Vasquez,4775 (44).
Smith,D. N. et al., 1748, 8670 (56); 13908 (33); 14052
(56).
Smith,H. H. & G. W. Smith,240, 323, 353, 1840 (46).
Smith,S. F. et al., 1503 (8); 1599 (17).
Sneidern,1191 (7).
Snethlage,E.(?),s.n. MG 9411 = R18362 (6).
Soares,E., 189 (47); 462 (21).
Soejarto,D., 580 (32).
Soejarto,D. et al., 4198 (35).
Solomon,J. C., 9255 (51); 12948 (56).
Solomon,J. C. et al., 7140 (56).
Soria,N., 3419 (56).
Sothers,C. A., 727 (6).
Sousa,J., 76 (28).
Souza,M. A. D. & P.A. C .L. Assunqao,491 (28).
Souza,M. A. D. et al., 396 (28).
Sperling,C. R. et al., 6157 (14).
Spichiger,R. & F. Encarnaci6n,1043, 1122 (35).
Spichiger,R. & P.A. Loizeau,4108, 4161 (cf. 55);4758,
4759 (28).
Spruce,R., 1453, 1453 bis (22); 1757 (35); 2769 (7);
3061 (19); 3092 (47);Nectandra6 (35).
Stahel,G., 43 (14); 236 (23).
StehIl, H., 387, 919, 1168 (46).
Stehle,H. & M. StehIl,6337 (46).
Stein,B. & A. Gentry,1660 (41).
Steinbach,J., 3274, 5291, 5369, 6985, 7264 (56); 7264
(35).
Steinbach,R. F., 765 (56).
Stergios,B. & G. Aymard,4171, 7371, 7386, 7400 (35);
7524 (19); 9006, 9104 (35).
Stergios,B. & P. Stergios,11427 (47).
Stergios,B. & J. Velazco,14495 (33).
Stergios,B. et al., 8187, 8280, 9607, 9610, 9706, 9839,
13259 (35); 13295 (21).
Steward,W.C. et al., P20357 (21).
Steyermark, J. A., 57935 (47); 59001 (cf. Rhodostemon-
odaphnegrandis);60428 (57); 75564 (47); 90315
(35); 90316 (47); 90535 (25); 91524 (48); 97760
(47); 107391(25); 107466(47); 111387(37).
Steyermark, J. A. & G. Agostini,7 (48).
Steyermark, J. A. & G. Davidse,116876, 116957(46).
Steyermark, J. A. & V. C. Espinoza,110275(56).
Steyermark, J. A. & R. Liesner,118687(56).
FLORANEOTROPICA
Steyermark, J. A. & S. Nilsson, 255 (60).
Steyermark, J. A. & P. Redmond,117081(35).
Steyermark, J. A. & C. Steyermark, 95347 (48).
Steyermark, J. A. et al., 100298(48); 101408(4); 111746
(56); 119895(46); 126162(24); 126285(57); 131425
(27); 131500(35).
Strier,K. B., 992 (34).
Sturrock,583 (46).
Stutzde Ortega,L. C., 1238, 2175, 2223 (56).
Sucre,D, & P. I. S. Braga1404, 1825 (56).
SurinameForestBureau,2936, 3197 (37); 3494 (23);
5884, 6884 (37).
Sytsma,K., 994 (10).
Talbot, H. F., s.n. (56).
TameiraoNeto, E., 87, 2653 (56).
TameiraoNeto, E. & M. S. Werneck,1244, 1245 (56).
Teixeira,E. M. & A. E. Brina,s.n. BHCB36260 (56).
Teixeira,L. 0. A. et al., 852 (28); 1025, 1163 (7); 1221
(22).
Tello,560 (56).
Tessmann,G., 3439 (35); 3058a, 3999, 4248 (17); 4734
(59); 4736 (14); 5126 (35); 5146 (31); 5288 (29).
Thomas,W.et al., 4059, 5370 (43);5089 (22);6775 (29).
Thomsen,K., 595 (15).
Tillett,S. S. & C. L. Tillett,45705 (47); 45789 (37).
Tillett,S. S. et al., 43990, 45163 (33).
Timana,M. & N. Jaramillo,2466 (59); 2476 (50); 2538
(59).
Timana,M. & 0. Phillips,1868, 1878, 1882 (15).
Tipaz,G., 2516, 2632 (15); 2730 (17).
Tipaz,G. et al., 581 (17);709 (59); 1104(Rhodostemon-
odaphnejuruensis).
Tirado,M., 1010 (8).
Tirado,M. et al., 462 (15); 586 (10).
Toledo, F. R. N. & J. A. Lombardi,s.n. BHCB 46022
(56).
Torres,J., 156, 300 (35); 3092 (cf. 59).
Triana,J. 1032 (4); 1033 (11); 1059 (4); 2040-2 (11);
2060 (4).
Tunqui,S., 215 (32); 316, 589, 649 (cf. 59).
Ule, E., 5 (56); 5581 (58); 5584 (29); 5710 (17); 6296
(14); 8125 (27); 8848 (21); s.n. R. 61128 (56).
Urrego,L. E. et al., 1012 (45); 1218 (cf. 51).
Usteri,P. A., 3026 (56).
Valcarcel,J. H., 383-I/F(29).
Valcarcel,J. H. & M. Chota,6/619 (cf. 55); 7/75 (28); I/
98 (33).
Valdespino,I. A. et al., 278 (48).
Valle,M. H., 5 (56).
Valverde,L. & I. Penia,1094 (24).
Varejao,de JesusM., s.n. INPA140522(22).
Vargas,H., 1152 (59).
Vargas,H. & P. Grefa,764 (32).
Vargas,I. G. et al., 120 (15); 1098, 1137, 2478 (17).
Vargas,R., 2 (56); 1096(35); 3319 (9); 4193 (32); 6611,
10989 (35); 12287 (58); 12296 (29).
Vasquez,R. & G. Criollo,1785 (13).
INDEX OF LOCALNAMES
Vasquez,R. & C. Grandez,17475 (58); 17487 (15).
Vasquez,R. & N. Jaramillo,40 (cf. 59); 203 (56); 1256
(35); 2425 (53); 2441 (50); 2501 (57); 3157 (20);
3165 (47); 5253 (20); 5258 (35); 5277 (15); 5457
(20); 5697 (8); 6085, 7174 (15); 7620 (20); 7633,
7644 (47); 7647 (20); 8114, 8215 (31); 8270 (15);
8398 (35); 8414, 8766 (20); 9123, 9125 (29); 9180,
9183 (9); 9593 (39); 9606 (32); 10228 (15); 10458
(59); 10749 (13); 11442 (55); 11467 (cf. 59); 11577
(35); 13609 (31); 13637 (28); 13711, 13945 (31);
14922 (39); 20300 (32).
Vasquez,R. & R. Jong,12397 (54).
Vasquez,R. & R. Rojas,21920 (cf. 49).
Vasquez,R. & T. Soto, 12350 (28).
Vasquez,R. & H. Valderrama, 1363 (31).
Vasquez,R. & A. Vdsquez,20969 (8).
Vasquez,R. et al., 111 (35); 648 (13); 651 (31); 4778
(35); 5307 (28); 5434 (20); 6393 (58); 6665 (15);
6666 (29); 6675 (15); 6742 (7); 10934 (15); 10939
(36); 11928(59); 11954(50); 12391(32); 12435(8);
14393 (31); 17706 (39); 18095 (53); 19459 (8);
21426, 21528, 21535 (53); 23419 (50); 23530 (13);
23873 (32); 23906 (8); 23948, 24025 (59); 24341
(55); 24898 (8); 25231 (39); 25237 (31).
Velazco,J., 587 (35); 859 (47); 1032 (33).
Velloso,H., 170, 205, 209, 398 (56).
Vester,H. & R. Roman,865 (45).
Vicentini,A. & E. da C. Pereira,903 (8).
Vicentini,A. & E. M. Venticinque,1000 (32).
Vicentini,A. et al., 635 (cf. 53); 993 (cf. 55); 1224 (7);
1317A, 1359, 1394 (21).
Vidal,J., 1342, 1710, 2020 (56).
Wallnofer,B., 17-31588 (17); 111-8488 (53); 14-
221287 (32).
Walter,B. M. T. et al., 3517 (56).
Wasshausen,D. C. & F. Encarnaci6n,649 (59).
Weberbauer,4680 (28).
Wedel,H. von, 2257 (10).
Werff,H. van der, 12953, 12964 (14).
INDEXOF LOCALNAMES
aguacat6n,45 canela, 74, 76
aiju'ywahu, 119 canela amarella,117
ajua, 123 canela frade, 117
amarillo,79 canela garuva, 117
anis moena, 82 canela paluda, 117
ant tree, 67 canelao, 74
azywa'yw-pihun,31 canoe tree, 42
casha moena, 45
bastardsilverballi,59 cedre, 97
bois marble,95 chaviaco negro, 76
137
Werff,H. van der & E. Gudinlo,11136 (Rhodostemono-
daphnejuruensis).
Werff,H. van der & W. Palacios,9246 (cf. 49); 10385
(49).
Werff,H. vanderet al., 8340, 8342 (43);9776 (39);9796
(13); 9815 (9); 9991 (39); 10045 (28); 10048 (58);
10053 (29); 10084 (8); 10187 (39); 10207 (20);
10237 (50); 10242 (39); 13116, 13147 (15); 15436
(40); 15736 (50); 15738 (8).
WesselsBoer,J. G., 1043 (14); 1381 (23).
Whitefoord,C., 3533, 4267, 4490, 5349 (46).
Wijninga,V., 580 (56).
Wilde,L., 6-88 (47); 14-88 (22).
Williams,L., 609 (35); 1002, 1003, 1004, 1193, 1203
(36); 1494, 1877(35); 11348(47); 14370(32); 14788
(47); 15179 (25); 15367 (35).
Wilson-Browne,Fr. G., 104, 178 (27); 454 (57); 470
(27).
Wolfe, 12194 (17).
Worbes,M., 1012 (22).
Woytkowski,F., 5829 (17); 5864 (8); 6304 (29); 6315
(35); 7179 (59); 7593 (17); 8323 (43).
Wurdack,J. J., 2339 (57); 2362 (37); 2493 (59).
Wurdack,J. J. & L. S. Adderley,43064 (35);43153 (19);
43169A,43169B (35);43435 (19);43592 (4);43647
(25).
Wurdack,J. J. & J. V. Monachino,39775 (35).
Young,K. & G. Sullivan,663 (33).
Zak,V., 4093 (55).
Zak,V. & S. Espinoza,4671 (8); 4835 (50).
Zardini,E. M., 7710, 7845, 7911 (56).
Zardini,E. M. & C. Benftez,3159, 3182, 3182, 3205,
3205, 3236, 3429 (56).
Zardini,E. M. & L. Guerrero,47737, 47746 (56).
Zarucchi,J. L., 3031, 3054 (35).
Zarucchi,J. L. & C. E. Barbosa,3679 (35).
Zarucchi,J. L. et al., 1830 (35); 3093 (22); 7120 (11).
Zaruma,J., 695 (50); 810 (8).
Zaruma,J. & A. Arguello,453 (8).
Zuleta,J., 42 (50); 45 (57); 50 (50); 142 (59).
cunchi moena, 121
cunshi moena, 45
dimukuima,97
harige pisie, 63
ha-ro, 70
hioma cocuir-ey,68
honcatohue,105
138 FLORA NEOTROPICA

inchaquito,105 louro cururu,70 palta moena, 70

isma moena, 76 louro de folha cinzenta, 105 pampamuena, 78
isma muena, 78 louro de f6lha larga, 72 pee-shee', 76
jigua, 78, 97 louro de f6lha peluda, 90 peroua-yek,62
jigua de mierda,27 louro do igap6, 76, 121 pisie, 63
louro dorado, 109 plata gigante, 105
ko-ma-nee-nee-ko,76 louro dourado,109 pucacurocaspi, 67
ko-mwa-kb-ree,42 louro flMr,76 puspo moena, 68
louro imbauba,72
las brisas, 70 lourojambo, 67 roble, 117
laurel, 70, 117 louro pichuri, 105 roble anis amarillo,70
laurel aguacate, 117 Louro pirarucu,24 roble fusi, 90
laurel babosa, 72 louro preto do igap6, 121 roble palta, 27
laurel blanco, 76, 92, 97 louro roxo, 109 roblecillo puchirfn,27
laurelblanco de orilla del rfo, 76 louro seda, 105
laurel bobo, 98 louro tambaqui,76 sacha muena, 117
laurel bongo, 98 louro vermelho,90 sacha palta, 102
laurel carutillo,72 sanango, 76
laurel de babilla, 61 mahamira,119 shiringochy, 117
laurel de revalta,76 mantaga,34 shisho moena, 66
laurel del rio, 76 mautaga,34 shisho moenita, 66
laurel fino, 76 moena, 66, 67, 90 siroewaballioenilebobandikoro,59
laurel hediondo, 70 moena amarilla,69 sweetwood, 95
laurel hormiguero,62 moena blanca, 45, 114
laurel moroti, 117 moena callhuangiaamarilla,34 temachi,45
laurel negro, 34, 63 moena de altura,45 tikis, 70, 114
laurel oriyera, 76 moena de bajo, 76 tinchi, 114, 117, 123
laurel plateado,97 moena de hoja grande,72 tinci, 70
laurel rebalsero,76, 97 moena hoja ancha, 105 tindu, 114
laurelito,76 moena negra, 34 tunchi, 114
laurierbord de mer, 95 moena rosada, 117
lauriergris, 95 moenilla, 114, 117, 123 uchitinchi,70
lauriergwa gwen, 95 moenito amarillo,48 und yuwich, 70
lauriermarbre,95 moroman,62 urcuayua,105
laurierpete, 95 muena, 32,.42, 69, 70
limon moena, 82 muena amarilla,67 wakamei, 62
louro, 23, 57, 58, 66, 70, 121, 123 muenablanca, 76 wau yuwich, 70
louro abacate,45, 67 muneu-yek, 119 wild pear, 72
louro amarello, 109
louro branco, 105 ocatoe, 70, 94 yacu muena, 76
louro canela, 76 ocatue, 105 yahuemomo,94
louro cedinha, 97 yellow silverballi,72
louro cedro, 67 palometamicuna,48 yiyiwa-ageyi, 45
louro comum, 79 palta amarilla,27 yuwich, 110, 123

INDEX OF SCIENTIFIC NAMES

New namesand combinationsarein boldfaceandsynonymsarein italics. Page numbersin boldfaceindicate

primarypage references.

Acrodiclidium Ampelodaphne,3, 11, 16

geminiflorum,125 arunciflora,51
Aiouea, 5, 16 dasyantha,60
impressa, 124 macrophylla,16, 57
maguireana,124, 132 Anaueria,7
tomentella, 124 Aniba, 13, 72, 106
INDEX OF SCIENTIFICNAMES 139

Aniba (continued) Endlicheria(continued)

bracteata,125 bullata,3, 8, 16, 21, 23*, 26, 28, 127
flexuosa, 42, 44 canescens, 4, 6, 7, 13, 16, 22, 67, 78*, 79, 80, 127
juruensis, 124 canescens species group, 13, 14, 82, 92, 127
krukovii,121 chalisea, 5, 13, 16, 19, 66, 71, 72, 73*, 127
reticulata,26, 28 chrysovelutina, 4, 5, 11, 14, 17, 34*, 35*, 36, 126
tschudyana,97,98 citriodora,4, 13, 22, 31, 80, 82, 83*, 127
Aspidostemon,7 cocuirey, 3, 13, 18, 62, 66, 67*, 68, 127
Aydendron, colombiana,20, 38, 40*, 41, 127
argenteum,125 coriacea, 5, 8, 19, 23*, 24, 25*, 126
macrophyllum,36 debilis, 123
dictifarinosa,3, 13, 18, 59*, 61, 62, 68, 127
Beilschmiedia,7 directonervia,13, 18, 57, 69, 70, 71*, 127
zapoteoides, 125, 133 duotincta, 4, 13, 16, 78, 89*, 90, 91*, 92, 94, 127
dysodantha,3, 4, 19, 42, 47, 48, 49*, 127
Caryodaphnopsis,7 endlicheriopsis,7, 79, 124
Cassytha,2, 7 ferruginosa, 8, 22, 26, 28, 29*, 30*, 79, 126
Chlorocardium,7 formosa, 4, 16, 19, 33, 44, 45, 46*, 114, 127
Cinnamomeae,7 glaberrima,42, 44
Cinnamomum,5, glomerata,3, 4, 7, 13, 16, 19, 26, 72, 73*, 74
Citrosma goeldiana, 124, 130
paniculata, 184 gracilis, 3, 5, 14, 16, 20, 21, 32, 65, 117, 119*, 121,
Cryptocarya,7, 127
hirsuta, 2, 114 grandis, 124
pyriformis,42 grisea, 118
robusta, 112 griseo-sericea, 14, 17, 95, 99*, 100, 101*, 102, 127
Cryptocaryeae,7 guadaloupensis,94, 95
hirsuta, 16, 114
Dicypellium, 31 impressa,3, 124
jefensis, 11, 19, 37, 38, 39*, 40*, 127
Endlichera,3, juruensis, 124
Endlicheria klugii, 4, 6, 14, 17, 20, 108*, 109, 110, 112, 127
sect. Macrolocellata,3, 5, 16 krukovii,3, 14, 20, 121, 122*, 123, 127
sect. Microlocellata,3, 5, 8, 16, levelii, 13, 18, 59*, 60, 61, 62, 127
subgen. Ampelodaphne,3,16, 32, lhotzkyi, 7, 17, 105, 108*, 109, 127
subgen. Anosphaeria,3, 16 longicaudata,3, 5, 8, 21, 27, 30*, 31, 119, 126
subgen. Euendlicheria,3, 16 longifolia, 114, 117
subgen. Hemiajouea,3, 16, 47 lorastemon, 5, 13, 22, 77, 89*, 92, 93*, 94, 127
subgen. Ocoteopsis, 3, 16 loretensis, 124
acuminata,3, 4, 14, 20, 65, 82, 117, 119*, 120, 121, macrophylla,13, 18, 56*, 57, 58, 59, 127
123, 127 maguireana,124
Ampelodaphnespecies group,3, 4, 5, 11, 13, 14, 33, 51, melinonii, 3, 13, 18, 59, 62, 63*, 64, 127
57, 58, 59, 60, 64, 66, 69, 74, 78, 127 metallica,4, 5, 6, 11, 14, 16, 17, 19, 20, 33, 34*, 36.
bracteatasub-group,13, 51, 66 105, 123, 126
multiflorasub-group,13 metallica aff. sp. 1, 11, 17, 134
anomala,3, 4, 5, 6. 13, 16, 17, 21, 74, 76*, 78, 92, 94, metallica aff sp. 2, 11, 19, 135
109, 121, 127 metallica species group, 4, 11, 14, 126
anomalaspecies group, 13, 14, 127 Microlocellataspecies group, 8, 14, 31, 126
arachnocome, 4, 18, 49*, 51, 52, 53*, 54, 67, 127 mishuyacensis,5, 11, 19, 37, 40*, 41, 42, 127
arenosa, 13, 18, 54, 55*, 56*, 57, 67, 127 multiflora,18, 58, 59*, 60, 62, 63, 65, 127
argentea, 17, 110, 111*, 112, 113*, 127 nilssonii, 14, 20, 122*, 123, 127
arunciflora,3, 4, 6, 18, 49*, 51, 52, 54, 67, 127 oreocola, 13, 21, 83*, 86, 87*, 88, 127
aurea, 18, 106, 107*, 108*, 109, 127 panicularis, 114
balsamea, 123, 134 paniculata,2, 3, 4, 5, 6, 7, 14,16 20, 21, 48, 77, 114,
boliviensis, 114,117 116*, 117, 119, 121, 123, 127
bracteata,3, 19, 61, 62, 63*, 65, 66, 67, 68, 127 paniculataspecies group, 14, 127
bracteolata,3, 16, 17, 64, 95, 96*, 97, 127 paradoxa,3, 16, 19, 45, 46*, 114, 127
browniana,5, 6, 11, 16, 20, 34*, 36, 37, 38, 127 poeppigii, 114, 117
brownianaspecies group, 11, 14, 41, 45, 48, 51, 114, punctulata,3, 4, 5, 8, 19, 22, 23*, 24, 126
127 punctulataspecies group, 2, 8, 14, 126
140 FLORANEOTROPICA

Endlicheria(continued) Licaria, 13, 31, 128

pyriformis,4, 16, 19, 27, 31, 37, 42, 43*, 44, 127 cannella, 24
racemosa, 114, 117 Lindera,5
reflectens,3, 4, 13, 18, 32, 59, 63*, 64, 65, 127
robusta,17, 20, 45, 112, 113*, 114, 127 Mespilodaphne
rubra, 3, 13, 14, 22, 82, 83*, 84*, 127 pyriformis,42
rubriflora,6, 8, 16, 19, 21, 23*, 26, 27, 28, 126 Mezilaurus,7
ruforamula, 3, 4, 14, 16, 17, 92, 102, 103*, 104*,
Nectandra,4, 15
105, 109, 110, 127
lucida, 114
sericea, 3, 7, 16, 17, 18, 94, 95, 96*, 97, 105, 109,
Neocinnamomum,7
123, 127
Neolitsea, 7
sericea species group,4, 13, 14, 38, 45, 77, 92, 94,
100, 102, 105, 109, 127
Ocotea, 2, 5, 7, 8, 13, 14, 24, 124, 127, 132, 133, 134
sprucei, 8, 13, 14, 22, 30*, 31, 32, 119, 126 cernua,24, 128
szyszylowiczii, 13, 16, 21, 88, 89*, 90, 127 cernua species group, 8, 23
tessmannii,7, 13, 18, 67*, 68, 69, 71, 127 debilis, 125
tomentella, 124
diffusa, 134, 136
tomentosa, 4, 21, 48, 49*, 50*, 127 discrepens, 13
trianae, 26, 28
endlicheriopsis, 13, 79, 124
tschudyana,18, 97, 99*, 100, 105, 114, 127 helicterifoliaspecies group, 15
verticillata,3, 4, 7, 13, 18, 51, 62, 66, 67*, 68, 127 indirectinervia,13
villosa, 58, 60 lhotzkyi,108
vinotincta,3, 5, 13, 14, 21, 32, 83*, 85, 86, 88, 127 nigrescens, 136
williamsii, 4, 5, 13, 17, 77, 78*, 90, 92, 127
pauciflora,8, 24
wurdackiana,26, 28 punctulata,22, 23
xerampela, 4, 13, 21, 78*, 80, 81*, 127 rufovestita,13, 135
zapoteoides, 124 simulans, 74, 77
Ocotea complex, 8 15
Goeppertia,3, 16, Ocotea s.str, 7, 8, 15
anomala, 74 Oreodaphne
argentea, 125 colombiana,40
cantagallana,114
caudata, 125 Persea, 5, 7
chrysophylla, 109 Phoebe
dysodantha,47 impressa, 124
geminiflora,125 Pleurothyrium,5, 15
hirsuta, 16 Phyllostemonodaphne,31
var. coriacea, 114
var.glabrata, 114 Rhodostemonodaphne,2, 7, 8, 11, 13, 14, 15, 28, 33, 34,
var. hirsutior, 114 36, 51, 84, 86, 105, 119, 124, 130
var. latifolia, 114 antioquensis,8, 28, 129, 131
var. robusta, 114 celiana, 13, 86
longifolia, 114 crenaticupula,13, 33, 124
multiflora,3, 58, 60 cyclops, 11
panicularis, 114 debilis, 124, 132
polyantha, 74, 77 elephantopus,8, 24
grandis, 11, 34, 36, 105, 124, 136
reflectens,64
sericea, 94 juruensis, 124, 132, 134, 136, 137
var. bracteolata,95 kunthiana,13
sprucei, 31 laxa, 11, 14, 51
mirecolorata,13
negrensis,21
Huberodaphne,3, 16, 31 parvifolia,31, 119
longicaudata, 16, 30 peneia, 11, 34
Hypodaphnis,7 praeclara,11, 34
recurva, 11, 13, 20
Laureae,7 revolutifolia,13, 22, 28, 30
Laurus,5, 7 rufovirgata,13
dysodantha,47 sauilensis,11, 34
INDEX OF SCIENTIFICNAMES 141

Rhodostemonodaphne(continued) Strychnodaphne
scandens, 26, 31 lhotzkyi,108
steyermarkiana,13, 86 Systemonodaphne
tumucumaquensis,134 geminiflora,125
Rubiaceae,3
Umbellularia,5
Sassafras,7
Schauera,3, 16
Sextonia, 7 Williamodendron,7