SERIES OF REVISIONS OF APOCYNACEAE
PART XXIVA REVISION OF THE TRIBEAMBELANIEAE(APOCYNACEAE-PLUMERIOIDEAE)

JAMES L. ZARUCCHI
Missouri Botanical Garden,P.O. Box 299,
St. Louis, Missouri 63166, USA
Received29Oct.1986
DateofPublication 16Febr.1988

Wageningen

mm A g r i c u l t u r a l U n i v e r s i t y0000 0262 8796\/

ISBN90-6754-100-1© AgriculturalUniversity,Wageningen,theNetherlands,1988.
No part of thispublication, apart from abstract, bibliographic data and brief
quotationsembodiedincriticalreviews,maybereproduced,recordedorpublished in any form including print, photocopy, microform, electronic orelectromagnetic record without written permission from the publisher Agricultural
University,P.O.Box1901,6700HBWageningen,theNetherlands.
PrintedintheNetherlandsbyDrukkerijVeenmanb.v.,Wageningen

INTRODUCTION
The present paper is a monographic revision of the six genera and seventeen
species comprising the wholly neotropical tribe Ambelanieae, with two of thegenera described herein as new. The genera of Ambelanieae with the number
ofspeciesineach are: Ambelania Aublet (3), Mucoa Zarucchi (2), Spongiosperma
Zarucchi (6),Molongum Pichon (3),Rhigospira Miers(1),and Neocouma Pierre(2). In addition to the two new genera, two new species are described and nine
newcombinations aremade.Thisrevision isbased upon the study of specimensfrom more than fifty herbaria, and upon field studies made by the author in
Colombia and Braziloneightspeciesinfiveofthesixgenera.

TAXONOMIC HISTORY
The first description and illustration ofaspeciesbelonging tothetribe Ambelanieae appeared in AUBLET (1775), Histoire des plantes de la Guianefrançoise
under 'Pentandria, Monogynia' oftheLinnaean sexualsystem.Thespecies, Ambelaniaacida,was based on a collection made in thevicinity of Cayenne during
Aublet's residence in the French colony (1762-64). The name Ambelania, aswith most of Aublet's generic names, was formed by the Latinization of one
ofthelocalGalibi namesfor theplant, 'Ambelani'.
Until 1860,Ambelania included A. acidaasthe only species.Five new species
and one variety from Brazil, and one species from French Guiana, were added
by MUELLER (1860a, 1860b).These species were:Ambelania cuneata, A. macrophylla, A. quadrangularis, A. tenuiflora with var. tenuiramea, and A. sagotii.
MUELLER (1860a) also made the combination A. laxa, based on Tabernaemontanalaxa Bentham.
MIERS (1878) described a new species, A. cucumerina, which is shown in thepresent work to consist of material of two distinct species - Ambelania acida
and Spongiosperma macrophyllum. MIERS excluded two of MUELLER'S species
from Ambelania {A. quadrangularis and A. macrophylld), placing them in his
new genus Rhigospira (R. quadrangularis and R. venulosa,respectively). RhigospirawasincludedbyMIERSinthetribeTabernaemontaneae, andincluded sevenspecies previously treated by MUELLER (1860a) under Tabernaemontana sect.
Stenocephalium and under Ambelania. Four of the species of Rhigospira recognized by MIERS are referable to Macoubea, and the remaining three represent
speciesofNeocouma, Spongiosperma, and Rhigospira sensu stricto.
Rhigospira wasnotaccepted byseveralsubsequent reviewersofthe Apocynaceae. BÂILLON (1891) in Histoire des Plantes and SCHUMANN (1895) in Engler
and Prantl,DieNatürlichen Pflanzenfamilien included thegenus,alongwith numerous other generic segregates, in synonymy under Tabernaemontana. SCHUMANN listed Ambelania as containing six species, yet he named only A. acida,A. laxa, and A. macrophylla.
Agricuniv. WageningenPapers87-1 (1987)

1

PIERRE(1898)described Neocouma, based on Tabernaemontana ternstroemiaceaMuell.-Arg.
HUBER (1902) described Ambelania grandiflora with the common name 'Molongó' from near Belém at the mouth of the Amazon River. In the protologue,
hecompared thenewspeciestoA. macrophylla from theupper Rio Negro.
In 1910, GLAZIOU noted Ambelania camporum [nomen nudum] in his list ofplantsfrom centralBrazil. MONACHINO(1945)questioned itsdetermination andplacement in Ambelania, and PICHON (1948) listed it as referable to Secondatia
densifloraA. De Candolle.
In the 1930's,severalnew speciesinAmbelania and Neocouma were describedfrom Amazonian Brazil. MARKGRAF (1932b) described Neocouma duckei, thesecond species in that genus. This species was later transferred to Ambelania
by MONACHINO (1945)under thenewname A. markgrafiana, sincethe binomial
A. duckeiv/as alreadyinuse.TwoadditionalspeciesofAmbelania weredescribed
byMARKGRAF(1935):A. zschokkeiformis andA. duckei,both from central Amazonia.Inthecurrentrevision,theyareplacedinthegeneraMolongum and Ambelaniarespectively.
MARKGRAF (1941) provided a brief review of Ambelania, dividing the genus
into two sectionsbased on inflorescence position. He alsomade thenew combination 'Ambelania lucida (Humb. & Bonpl.) Mgf.', a species which had been
included in'Lacmellia' by MIERS(1878).
MONACHINO (1945)included eleven speciesin his 'Revision ofAmbelania, inclusive of Neocouma', proposing four sections: 'Acidae', 'Neocoumae', 'Quadrangulariae', and 'Laxae'. Sincethesesectionswerenot accompanied by Latin
diagnoses,theywerenotvalidly published.
PICHON (1948)proposed a new tribe, Ambelanieae ('Ambélaniées') with two
subtribes (Ambelaniinae and Macoubeinae) to include five American genera.
The five genera were: Ambelania, Molongum, Rhigospira, Neocouma, and Macoubea.PICHONconsidered Ambelania toconsist ofthree speciesintwo sections,'Euambelania' and 'Neocoumopsis'. These names are not validly published.
'Euambelania'contained twospecies,A. acidaandA. duckei,withA. cucumerina
treated asa synonym ofA. acida.'Neocoumopsis' consisted ofonly onespecies,
A. markgrafiana. Molongum Pichon was described as a new genus, equivalent
to'Ambelania sect.Laxae'of MONACHINO (1945),withthetypespecies designatedasM. laxum. Pichon divided thegenusinto three sections(Trichosiphon, Coilanthera,and Oligoon)withatotalofsixspecies.Thesection Trichosiphonserves
as the basis for the genus Spongiosperma, described herein. PICHON (1948) also
emended the genus Rhigospira to include a single species, R. quadrangularis,and retained Neocouma asamonotypic genus.
ROBERT WOODSON, in SCHULTES(1951),described Ambelania lopeziifrom theColombian-Brazilian border; A. lopeziiishereplaced in synonymy under Spongiosperma macrophyllum (basionym:Ambelania macrophylla).
MONACHINO (1958)described Ambelania oleifolia(as 'oleaefolia') from Venezuela,and in 1961added anewvariety, A. 'oleaefolid var. riparia.Both of thesetaxa aretreated hereinasdistinct speciesof Spongiosperma.
2Agric univ. WageningenPapers 87-1 (1987)

In 1971, MARKGRAF described two new speciesof Ambelania from Venezuela
and Brazil. Ambelania longiloba was described from the vicinity of Manaus(MARKGRAF in PRANCE, 1971b), and A. lanceolata from the Orinoco basin of
Venezuela (MARKGRAF, 1971).Both of these new species are referable to Spongiosperma inthepresent treatment.
Ambelaniapantchenkoana Markgrafwasdescribedin 1975from theVenezuela-Brazil border, and is referable to the genus Mucoa. Ambelania parvifiora,
placed herein as the second known species of Neocouma, was further added by
MARKGRAF in 1983.
In 1975,BOITEAU and SASTREdiscussed the systematic position of Macoubea,
removing it from the tribe Ambelanieae; it had been placed by PICHON (1948)

in its own subtribe. The new tribe Macoubeae was described and placed withfour additional tribes in the subfamily Tabernaemontanoideae. BOITEAU, ALT
LORGE, and SASTRE (1978) validated the tribal name Ambelanieae, attributing
itto PICHON.WiththeexceptionoftheremovalofMacoubea from the Ambelanieae, they accepted the treatment by PICHON (1948) recognizing four genera:
Ambelania, Molongum, Rhigospira, and Neocouma.

MORPHOLOGY AND ANATOMY
Habit
The seventeen known species in the Ambelanieae consist of woody shrubs
to small or medium-sized trees which under normal conditions have a single
erect trunk. Spongiosperma oleifolium, endemic to the base of Cerro Yapacana
in Venezuela, is the most diminutive species, with fertile collections gatheredfrom individuals less than 0.5 m in height, the tallest individual reported is acollectionof Rhigospiraquadt-angularis{Berlin3519)from thePeruvian Amazon
which measured 30 m. Most species in the tribe have individuals which rarelyattain a height of over 15m. In addition to R. quadrangularis only three otherspecies, Ambelania acida, Mucoa duckei, and Neocouma ternstroemiacea, are
known from individuals greaterthan 15m tall.
Treesof Ambelania, Mucoa, Rhigospira, and Neocouma areknown only fromterra firma forest where they usually form part of the lower story below the
canopy.Treesand shrubsof Spongiosperma andMolongum arefound in riparian
habitats, with M. lucidum known from both riparian and savanna areas of theupper Rio Negro.
Localconditions affect thehabit ofthetreeswithin theconstraint of the individual growth pattern for the species. Availability of sufficient light and nutrients, and the degree of crowding from adjacent trees all affect the eventual
habit of the tree. Those species of terra firma forest have branches restricted
to theupper portion of the treeand havea straight bole.The sharply ascending
branches (due to the crowding of other trees) may become horizontal and thebranchlets pendulous when laden with mature fruit. Individuals of riparian orsavannahabitats(i.e.,speciesofSpongiosperma andMolongum) aremoreirreguAgricuniv. WageningenPapers87-1 (1987)3

larly branched, although the basic architectural plan is one of dichotomous
branching following a terminal inflorescence. Riparian trees ofMolongum havetrunkswhichmay beswollenat the base.
The architecture of Molongum lucidum, which has terminal inflorescences,appearstofitScarrone's Model.Accordingtothismodel,thereisa rhythmically
active orthotropic terminal meristem with the trunk bearing branches in tiers;
theseorthotropicbranchesarethen sympodiallybranched duetoterminal flowering (HALLE, OLDEMAN & TOMLINSON, 1978). Some individuals of M. lucidum
whichIobservedpersonallyintheColombianVaupés(e.g.Zarucchi 2139,2483)
appear to adhere more closely to Leeuwenberg's Model, in which the apicalmeristem ofthemain trunk isaborted earlyindevelopment, producing dichotomously branchingmodules,thegrowth ofwhichisterminated by inflorescences.
HALLE (1978) has shown that a species which corresponds to Leeuwenberg's
Model when grown in open areas may follow Scarrone's Model when grownin the shade. In this case the difference between the two architectural models
isdetermined by theearly abortion or continued activity of the apical meristem
respectively. Sincethe change from Scarrone's Model to Leeuwenberg's Model
involves primarily the abortion of the apical meristem (which may happen at
any time, even by the accidental destruction of the growing apex), the lattermodelmay beconsidered asderived from the former.
Thenormalgrowth ofRhigospira quadrangularis,whichisencountered interrafirma forest and notinopenareas,fitsScarrone'sModel.Individuals ofSpongiosperma, likethoseof Molongum and Rhigospira, haveterminal inflorescences
whichcausethevegetativeapextobecomedichotomously branched. Depending
on other factors, as discussed above for M. lucidum, either Scarrone's Modelor Leeuwenberg's Modelwould apply to treesof Spongiosperma.
InAmbelania andMucoa theinflorescences areaxillary.InMucoa theinflorescences develop in a manner similar to that found in Couma of the Carisseae,
one ofthemany similaritieswhich suggested the anagrammatic name of Mucoa
forthenewgenusdescribedinthefollowing pages:theinflorescences areaxillary
to and develop simultaneously with the terminal vegetative axis. In Couma theleaves and branches occur normally in whorls of three or four. According to
HALLE, OLDEMAN and TOMLINSON (1978), Couma guianensis represents Rauh's
Model. Rauh's Model is characterized by a monopodial trunk which grows
rhythmically, thusproducing tiersofbranches;thetrunk and branchesare morphologically identical. Inflorescences are always axillary and do not affect theshoot system. This model may also be represented by Mucoa, according to myown observations of a mature tree of M. duckei {Zarucchi et al. 1852) 15 m
tall.
Bark
The bark of Rhigospira quadrangularis and Spongiosperma macrophyllum isthin and white to greyish white. In Mucoa duckei the bark is rather thick (ca.
8 mm), brownish grey when fresh turning to chocolate brown when dried, andexudes a moderate amount of white latex which is not sticky. In some of the4

Agric univ. WageningenPapers87-1 (1987)

other species of the tribe,including R. quadt-angularisand S. macrophyllum, thelatexisabundant and sticky.
Bark samples of Mucoa duckei show short (1-2 cm) longitudinal fissures
which appear aseruptions from below the surface. The bark of Rhigospira quadrangularis,however,iscovered with numerous fine cracks oriented transversely,aswellasirregular wavyverticalconnecting fissures.
The bark of Molongum lucidum isgrey, relatively thin, and has a flaking appearance,especially on older trunks.
Wood
Detailedinformation onwoodstructureofrepresentativespeciesofAmbelanieaeispresented inTable 1.
In the first reported study of the wood structure of the tribe, WEBBER (1945)
considered theminute structure of Molongum laxum (under thename Ambelania)based on a single voucher specimen, Ducke 163,from the upper Rio Negroof Brazil. She noted the extreme lightness of the wood, referring to the notes
published by BENTHAM (1841), and attributed this character to the thin-walled
cellswhichmakeup thewood. RECORD and HESS(1943)reported that the wood
hasaweight of9poundspercubicfoot, and WEBBER(1945)mentioned itspossibleusefulness wherelightweight woodsare desired.
ThewoodstructureofMolongum laxum isdiffuse-porous, with inconspicuous
growth rings. The dominant cell types in the wood are imperforate tracheary
elements.The pores areminute tomedium-sized, and inmost woods examined,
their frequency of occurrence ranged from solitary pores to radial multiples ofseveral pores. Radial multiples of ten or more pores are common in species of
Molongum. The vessels in wood of M. laxum are difficult to distinguish fromother cell types because of similarities in cell diameter and wall thickness. The
vessel element perforations are simple with wide rims at the ends or at somedistance from the ends of the vessel element. The perforations are horizontal
to oblique in orientation, and more than two perforations per vessel element
may be present. Wall pitting is minute and distinctly bordered, with no spiral
thickenings.Theparenchyma isgenerallyapotracheal-diffuse withfew scatteredcellsseenincross-section, although inseveralcases,shortwavytangential bandsof cells (metatracheal parenchyma) may be present. The wood fibers are notseptate, and no crystals have been observed in the wood. Rays are uniseriate,biseriate,rarelytriseriate,and heterogeneous.Squareanduprightcells normallyoccur at the ends of the rays. Radial channels are encounted in wood samples,
which RECORD and HESS(1943)and WEBBER (1945)attributed to distintegrated
leaf traces.
Stems and Nodes
The stems of plants in the Ambelanieae are usually terete although some arelaterallycompressed;inthecaseofRhigospira theyoungstemsaresharply quadrangular. Theyoung stemsinMolongum arethethinnest inthetribe,being only
about 2mmindiameter;plantsofNeocouma havethethickest stems, measuring
Agricuniv. WageningenPapers87-1 (1987)

5

es

e.

a) G
3 '~

S >»
l-l

'•3
e

t P

+l

2

•H 2

00 S-•od

Q -1. 8

CA

)

1—

?.!„

iffuse-3, us
attere»•3S.6ON ^ - ^çd

V * ""> T3+1?

\ £> S © s

O 00

oo°ï
r*^ —/

S? Û .

d in
i-H ooOO rt.

< g äK .3 H 2 S

ë
•3

eu
en

CA

"3

a

O
eu

i

•0

3

CA

O eu
h4
a "cd
0

<

"cd
'G

eu
OJQ
O

S

+iT
50

0

CU cd
"eu ' C

CA

K

eu

c
3

CD

CA

-D

<S

CL)

X)

J3

CU

"ss

a

.a

3 '«^
o j>*

3
tX)
c
,0
0

CIN S

K

CU

aai

?

-4-*

13

o 'M
D- 3 -o
« 3 2

Crt

'5
eu

"cd <uc
3 • euO CA ntCDC £ <u
CU
00 0
O
CÓ*

—v

-s

Q
eu
C
0

•S

t-i

T3
C

a

PO

§ 15
06 vp

,3

Q

CA +Z

t- y

00

'5 g v o
&

CA

CD

1-1

O 03
ON ^ *( - S o ,.3 Jr^euê« 5 w"la-4-*

CU CÖ > 1u •c S(N \©
<N ( S

-H7

r-^
•

a scd

eu " C

CA ~ -

+1^

1

i^

CU
_N cd

^

'B 'C

eu •4-»eu"û- cd

B
S
ci
C4-I

CA

t-T

CA Cd n-t

3

3 ^

o s e

v- 3 eu

o.

(N

ta

à fa
J X CO

5 'St l m .S

1s

-H' I

S3 °°

(N —

CN O

-H 2

+1"

<N vi

11

M

•a &j 8s — 81 «s•^ ^au .-H« S 2S

0

eu c
•S 0
eu
eu • 0
X! 3
O
O
0
ON 0

>

^

O

s

-^ 0

•8-3

§I

•9 s
3

•g
•s
%

<5

o

'S

as
Agricuniv. WageningenPapers87-1 (1987)

morethan 1 cmindiameterinmanyspecimens.Inmostspeciestheteretestemsof the internodal regions becomecompressed when approaching a node frombelow. The direction of compression below the node is perpendicular to theplane of the opposite leaves at the node. In many cases theyoung stemshave
conspicuous longitudinal grooves which run along opposite sides of thecompressedportionofthestem.Thesegroovesareveryapparentinelongatingvegetative shoots of Mucoaduckei; in this instance the groove is so deep that the
young stemappears astwofused teretestems.Theleaf arrangement alongthe
stemisdecussate and thedirection ofcompression ofthestemabovethe nodeisalwaysperpendiculartothefoliarplaneatthepreviousnode.
TheyoungstemsoîRhigospiraprovideaclearvegetativecharacterforidentification.Theseangledstemsaregreenandsquareorrectangularincross-section;
withageandsecondarygrowththeybecomebrownishgreyandterete.
SpeciesintheAmbelanieaehavedecussateleaveswithunilacunar,singletrace
nodes;thetracedepartsthesteleatorjustbelowtheleveloftheleafattachment
tothestem.Thetracesaresimpleandcontinuous,lackribtraces,andareusually
U-shaped at thenodalgap.Upon entering thepetiole thetrace forms aclosed
siphonostelewhichremainsinthatforminthelowerportionofthepetioleuntil
itonceagainformsaU-shapedtracethatentersthemidriboftheblade.
One special feature of the nodes of the Ambelanieae is the presence of aninterpetiolar line which may be confused with a stipular scar. Due to such a
confusion Molongum lucidum was first assigned to the rubiaceous Psychotria.
Thisinterpetiolarlineorridgeisathinbandoftissuewhichconnectsthepetiole
basesoftheoppositeleavesandmaycontainconspicuousorhiddenglandscapableofproducingaresinousexudate.TheseglandsgivetheflowersandinflorescencesofAmbelaniaandMucoa(thetwogenerainthetribewithaxillaryinflorescences)astickyandresinousappearance.InRhigospirathedriedexudateprotects the terminal bud from damage, presumably from insect herbivores. The
petioles at the point of attachment contain many flattened glands which aretightly appressed in the axils and are visible even on older leaves.The glands
ofMucoa areclavate and produce aresinous exudatewhich not only protects
thedevelopingvegetativeshootbutalsothesimultaneouslyelongatinginflorescenceaxes.
The leaf axils of Neocouma ternstroemiacea contain a single small saucershaped gland ontheadaxialsurface ofthepetiole. CoumaintheCarisseaehas
asimilartypeofglandinthesameposition.
LeavesThe leaves of all members of theAmbelanieae are decussate and uniformly
spacedwithdiscreteinternodalregionsalongtheultimatebranchesoftheshrub
ortree.Leaf shapesmaybelanceolate,elliptic,ovate,oblong,orobovate,and
theymayrangefrom about5toover40cminlength.Leavesonsterilebranches
or older leaves some distance from the inflorescence usually far exceed theselimitsinsize.Theleafbasesandapicesarealsohighlyvariable.
The petiole ranges from lessthan 5mm to more than 40mmin length andAgricuniv. WageningenPapers87-1 (1987)1

is usually conspicuously canaliculate, especially near the point of attachment
to the stem. Glands are frequently present in large numbers and are packed
together tightlyintheleafaxils.
Following the terminology proposed by HICKEY (1979) for the architectureof dicotyledonous leaves, all plants of the Ambelanieae have camptodromous
leaves, as the secondary veins do not terminate at the leaf margin. Illustrations
ofcleared leavesfrom sixrepresentative species,one from each ofthesixgenerain the tribe, are presented in Photographs 1and 2. Fine structure of the leaf
venation (at 15 x or 30 x ) is also presented for the same leaf samples. All
genera except Rhigospira have brochidodromous-type venation. In many cases
the leaf margin isor becomes revolute upon drying and may somewhat obscuretheconnectingintramarginalvein.TheleafarchitectureofRhigospiraiseucamptodromous. In Rhigospira the secondaries are connected to the superadjacentsecondaries by a number of cross veins; see Photograph 2C. The leaf venation
ofnearlyallspeciesofthetribeiscloselyreticulate,although leavesin individuals
ofMolongum zschokkeiforme havearather open and irregular reticulation.
Thethickleavesof Neocouma ternstroemiaceanotonlyhaveafinereticulationof interwoven veins and veinlets but also contain sclerenchymous cells within
the leaf tissue. The sclereids of Neocouma were first observed and mentioned
inthegenericprotologueby PIERRE(1898);Photograph 2Fshowsa30 x magnifiedsection of cleared leaf tissuefrom Fróes475which illustrates the very compactarrangement ofthevascular systemwith sclereids.Thethreeknown speciesof Ambelania also contain sclereids in the leaf tissue. In both Ambelania and
Neocouma branched and unbranched filiform sclereids are present, sometimes
with one to several short spicule-likeprocesses. In A. duckei (and other species
ofAmbelania) unbranched sclereidspredominate, whereasintheobserved samplesofNeocouma both branched and unbranched sclereidsoccurred in approximately equal numbers.
Inflorescence
The inflorescence typically found in the tribe Ambelanieae is a many-flowered, axillary or terminal cyme. According to WOODSON (1935), in his study of
inflorescence types within the Apocynaceae, such a dichasial cyme represents
the'primitive'typeofinflorescence for thefamily. IntheAmbelanieaethe inflorescencesmay beopen and lax,or compact.
The inflorescences contain usually small and scale-like bracts and bracteoles.
InNeocouma ternstroemiacea, however,thebractsarelarge,foliaceousand calycine,growingupto 1 cminlength.Thecommon pedunclemay beeither severely
reduced, in Ambelania and Mucoa, or rather long, usually over 5cm in Rhigospiraquadrangularis,Molongum laxum, and M. lucidum.
The axillary inflorescences of Ambelania are few-flowered, with rarely morethan eight flowers and buds visible at any one time. The peduncle is reduced
orabsent. InAmbelania, theinflorescences areusually borneseveralnodes fromthe growing apex of the stem, and they may continue to produce flowers wellafter thesubtending leaveshave fallen.
8

Agricuniv. WageningenPapers87-1 (1987)

-•v,rwj'wwv**-vrnmm

V.' .

PHOT. 1.Cleared and Stained Leaves of Representative Species of Ambelania , Mucoa, and Spongiosperma. A - B . Ambelania acida (Frees 1956); C - D Mucoa auckei (Zarucchi et al. 1852); E - F .
Spongiosperma riparinm(Schultes &Lopez 10132).(A,C,E. 1.5 x ;B,D. 15 x ;F.30 x).
Agricuniv. WageningenPapers87-1 (1987)

PHOT. 2.Cleared and Stained Leavesof Representative Species of Mologum, Rhigospira, and Neocouma.A - B .Molongum lucidum(Zarucchi 2139);C - D . Rhigospira quadrangularis(Zarucchi 1570);
E-F. Neocouma ternstroemiacea(Fróes 415). (A, C,E. 1.5 x ; B , D . 15 x ; F . 3 0 x ) .

10Agric univ. WageningenPapers87-1 (1987)

In Mucoa the development of theaxillary inflorescences issimultaneous withthe elongation of the terminal vegetative shoot. The peduncle isreduced or absent, and the total number of flowers within an individual inflorescence (twopernode,oneineachoftheoppositeleafaxils)ismuch smallerthanin Ambelania.
The major branching in the inflorescence reflects the decussate phyllotaxy
ofthevegetativeportionsoftheplants.Thesemajor brancheswithintheinflorescenceareoppositeand subtended bybracts.Subsequent branchingwhich mightoccuralongtheprimaryinflorescence axis,suchasinMolongum and Rhigospira,are perpendicular to the plane of the first pair of branches. Lateral branches
withinaninflorescence arealsodeterminate;theterminalflowersofthebranches
open onlyafter theterminal flower onthemain axisofthe inflorescence.
The flowers and inflorescence axes in individuals of Ambelania and Mucoa
may have a sticky, resinous appearance due to the exudate produced by the
glands in the leaf axils. The other four genera in the tribe do not exhibit thisresinousappearance orproperty eventhough theytoohave axillary glands.
Spongiosperma, Molongum, Rhigospira, and Neocouma have strictly terminal
inflorescences which usually bear many white flowers. In thecaseof Neocouma,
terminal inflorescences may occur at the apex of short side branches. In SpongiospermaandNeocouma theinflorescence iscompact,withtheflowers radiatingfrom a point atop the peduncle. The compact aspect of these inflorescences isduetotherelatively short lateral branchesand pedicels.
The pedicels in individuals of the Ambelanieae measure usually less than 1
cm in length and contain two small scale-like bracteoles.Within each bracteole
is a bud capable of producing several additional flowers from the bracteoles
on its own pedicel. The bracteoles may occur at any point along the pedicel,
but inmost casesthey arelocated near themiddle. In thismanner those flowers
which develop later tend to overtop the earlier flowers from the bracteoles ofwhich they have developed. In congested inflorescences not only are there few
inflorescence branchlets, but also the pedicels are relatively short and/or thebracteoles arelocated primarily at thebaseofthepedicels.
Flowers
PICHON (1948) used two floral characters to differentiate this tribe from theother berry-fruited genera in the tribe Carisseae: the ring of tissue at the baseoftheclavuncleand thecaudate anthers.
The flowers of theAmbelanieae arepedicellate with acup-shaped, five-lobed
calyx containing usually a variable number of glandular structures at the basewithin. The flowers arepentamerous, with the salverform corolla ranging fromwhite to orange and frequently fragrant. The five corolla lobes are contorted
in bud and twisted to the left at anthesis. The epipetalous stamens are inserted
in the lower part of the corolla tube, with the filaments adnate to the tube. The
anther cellsopen by slits,and haveapicescomposed ofsteriletissue.Thegynoecium includes a syncarpous superior ovary with axile placentation. The ovules
arenumerous and areattached to theplacental septum. Severaldifferent modes
Agric univ. WageningenPapers87-1 (1987)11

of development can occur giving rise either to a one- or two-celled fruit. The
clavuncle is borne at the top of the style and positioned near the middle of thecorollatube,atthesamelevelasthestamensbutnotaffixed tothem.Theclavuncle has a ring of tissue at the base, sometimes flange-like, and is usually fiveangled at the apex.
Flower size varies from those in Molongum, with the corolla tube less than
1cminlength, tothoseinSpongiosperma, with thetube surpassing 3cm.
Calyx
The calyx consists of five sepals which areconnate near the base, completely
glabrous, and tightly clasped to the base of the corolla tube. The sepals varyin length from about 1 to 14 mm in length and are quincuncially imbricate.
Insomecasesthesepalsmay vary slightlyinsizewithin a singlecalyx.
The smallest calicesoccur inspeciesofMolongum and inRhigospira quadrangularis, where they may be as small as 1mm in length. The largest sepals by
far are found in Neocouma ternstroemiacea, where they measure up to 14 mmlong and 7mm wide.The calyx ispersistent through mature fruit in all species
ofthetribeexceptfor N. ternstroemiacea, wherethey arecaducousearly in fruit
development. The sepals do not enlarge appreciably during fruit development,even though they may become thickened and slightly or markedly distended
by the expansion of the fruit and attached pedicel. In the fruit of Rhigospira
the sepals become hidden or barely perceptible at the articulation between thepedicel and the fruit. Thepedicelundergoes anon-uniform increasein diameter
along its length; the end attached to the fruit expands more, thus giving the
fruit pedicel theshape ofafrustum or truncated cone.
CorollaThecorollaissympetalous,withfivelobescontorted inbud,andis salverformwith a narrow tube, very constricted at the orifice, swelling near or below the
middle of the tube where the epipetalous stamens are inserted. The corolla isusually white or cream-colored, but may be yellow or somewhat orange in Molongum.
The smallest flowers are found in Molongum, where the corolla tube may
be shorter than 4mm, and the largest flowers are those of Spongiosperma, withcorolla tubesup to 30mm in length.
Pubescence occurs generally only within thecorolla tube in the area between
the insertion of the stamens and the small orifice. The portions of the adaxial
surface ofthecorollalobeadjacent totheopeningmayalsobyslightlytodensely
pilose.In Molongum, however, theflowers arecompletely glabrous.
The corolla lobes are twisted in bud. After anthesis the corolla lobes becomereflexed; because oftheoverlap of thelobeswheretheyareattached tothe tube,
thelobesappear to spiralto theleft whenviewed from above.
Androecium
Thefiveantherswiththeir filaments areadnatetothecorollatubeator slight12Agricuniv. WageningenPapers87-1 (1987)ly below the middle of the tube. The position of the stamens is evident froman external view,duetothe slighttomarked dilatation ofthetube at that point.
Theanthersarelanceolatewithauriculatetosagittatebases,sessileorsubsessile,from about 2to 10mm in length. The stamens are positioned at the same levelwithin the flower as the clavuncle, but they are not attached to this organ by
an exudate asisthecaseinsomeothermembers ofthe Apocynaceae.
The introrse anthers are attached to the filaments near the base, and dehisce
bytwolongitudinal slits.Theapicalportion oftheantheriscaudateand consistsof sterile tissue. In most of the genera, the suprastaminal part of the corolla
tube is pilose, with some hairs found at the point of attachment of the anthertothe filament/corolla tube.
Gynoecium
The ovary is syncarpous, glabrous, superior, bicarpellate, and bilocular withtheplacentation axile.Theovaryisfrom 1 to5mminheight,conical,cylindrical,
or ovoid in shape, and with the apex pointed, rounded, or truncate. The ovules
are numerous in each of the two locules. The terminal style is from less than
1mm toalmost 10mm inlength, and surmounted bytheclavuncle.
In all six genera of the tribe, the clavuncle has a thickened ring of tissue at
its base, a structure which both MONACHINO (1945, 1946) and PICHON (1948)
used to distinguish species contained in the Ambelanieae from members of the
Carisseae. Illustrations of the clavuncle for representative species in Spongiosperma, Molongum, and Rhigospira using the scannning electron microscope
are presented inPhotograph 3,with an illustration of a highly similar clavuncleofMacoubea. Thethickened ring oftissueat thebasemaybesimpleand slightly
undulate as in Spongiosperma or flange-like as in Rhigospira and Molongum.
The distal portion of the clavuncle is usually 5-lobed or 5-angled, sometimes
with short, dense hairs on the lobes (see Photograph 3A). The placement of
the clavuncle in relation to the anthers is such that the basal thickening of theclavuncleisat thelevelofanther attachment.
Fruit
The fruit in the Ambelanieae is a berry. The baccae vary in structure andappearance among thesixgenera, yetinallcasesonefinds a syncarpous, indéhiscentfruit which develops from abilocular ovary with axileplacentation. Fruit
variation within the individual genera is not significant. Characteristics whichareuseful indistinguishingthegeneraare:thenatureofthepericarp,the number
and arrangement of the seeds, the presence and type of pulp, and the one- or
two-celled mature fruit.
Variation in the structure of the pericarp is extensive. It can be thick andleathery as in Ambelania and Spongiosperma, relatively thin and delicate as in
species of Molongum, or hard and somewhat bony with a granular consistency
asinMucoa, Rhigospira, and Neocouma. The hard, granular surface ofthe fruitin Mucoa and Rhigospira is several millimeters thick, whereas in Neocouma it
iscrustaceous, about 1 mm thick, and somewhat brittle.
Agricuniv. WageningenPapers87-1 (1987)

13

PHOT.3.ScanningElectronMicrographsofGynoeciaof:A.Spongiosperma macrophyllum (Zarucchi
2435,37.5 x);B.Molongum lucidum(Zarucchi 2139,40X);C.Rhigospiraquadrangularis(Zarucchi
2428,37.5 x);D.Macoubea sprucei(Zarucchiet al.2554,32 x).

14

Agricuniv. WageningenPapers87-1 (1987)

The baccae ofAmbelania (sensu stricto)mature sothat the partition between
the two fused carpels of the syncarpous ovary remains intact and develops intoa fleshy dissepiment onwhichmany seedsareattached byfleshy funicles. Ambelaniaisuniqueinthat thetwocellsremain asopen cavitiesintowhich thedeveloping seeds project. In the other cases where the septum apparently remains
intact {Mucoa and Molongum), the seeds are completely embedded in the fruit
pulp, and are oriented more or less in the same direction. In Spongiosperma
the dissepiment tears during fruit maturation, leaving a one-celled fruit; thespongy seeds are attached to the fruit wall by low fleshy projections into thecavityfrom regionsadjacent totheremnantsoftheseptum.Theseedsarepacked
tightly into the cavity. In Spongiosperma, as in Mucoa, the seeds are numerous(usually morethan 50per fruit) and areacropetally imbricate. The seedsin Molongum are few (usually 2 to 4) and their orientation is either acropetally appressedorimbricate,dependingonthenumberofdevelopedseeds.In Rhigospira
they occur embedded inthefruit pulp and apparently lack anyparticular orientation.Thismay bepresumed tooccurinNeocouma aswell,although only dried
fruits of'Neocouma havebeenavailablefor study.Maturefruits ofR. quadrangularisusuallyhave30-60seedswhereasinN. ternstroemiaceathenumberisabout
15-20.
In somecollections ofRhigospira quadrangularissmall,apparently parthenocarpic fruits without seeds are known. These aberrant fruits reach only 2 - 3
cminlength, areovoidwith an acuteapex,and havea scarred surface.
Fruit shape varies appreciably in the several genera of the tribe. The fruitsin Ambelania are lanceovoid or ellipsoid to ovoid or globose. In cross-section
theymayberound,broadlyellipticduetobeingslightlycompressed, orpentagonal.Thedriedfruits ofAmbelania acidausuallyhavefiveprominentribs runninglongitudinally. Thefruit ofA. duckeiisovoidwithapointed apexand numerous
low,roundedribsrunningthelengthofthefruit. InMucoa, globoseand ellipsoid
fruits occur. In Spongiosperma, they are fusiform, ellipsoid, oblong, ovoid, or
globose. The few-seeded fruits of Molongum are ellipsoid to lanceovoid. Rhigospira fruits may become stipitate at the base when mature and are ellipsoid
to obovoid. Neocouma has ellipsoid or globose fruits. The size of fruit ranges
from about 1cm in Molongum to the large and elongate fruits of Ambelania
whichmay reach 15cmormorein length.
At maturity, fruit in most species is yellow or yellowish green. Both the exocarp and thepulp of Mucoa duckei are orange.
Seeds
Seeds are oblong, elliptic, or ovate in outline with the ventral surface usuallycompressed and the dorsal face rounded or angular. The seed coat ranges fromsmooth and shiny as in Ambelania to a tortuous, spongy and pitted surface inSpongiosperma. The seeds measure from about 5 mm in length in Molongum
to 16mm inNeocouma ternstroemiacea.
In the Ambelanieae the embryo is almost as long as the seed and is straightand embedded in an oily endosperm close to the usually planar ventral surface
Agricuniv. WageningenPapers87-1 (1987)15

of the seed. PICHON (1948) used the small (1-1.5 mm in length), flat, ovate tocircular cotyledons as one of the key characters to separate the Ambelanieae
from the Carisseae. In the Ambelanieae the straight, cylindrical radicle is atleast three times as long asthecotyledons, whereas in the Carisseae, the cotyledons areequal to or longer than theradicle.The hilumislocated on the ventral
surface, positioned approximately overthecenteroftheradicle,about one-third
the distance from the base of the seed. The circular or broadly elliptic hilum
is slightly depressed or.flush with the surface of the testa in all genera except
Spongiosperma, wherethere isa swellingof thespongy testa into a toroid shapearound thehilum.Ontheventralsurface oftheseeds,thereisaverylow,narrow,rounded ridge running from the hilum to the cotyledonal end of the seed; it
ismostpronounced inspeciesof Spongiosperma.
Seedlings
Seedling development has been observed in Spongiosperma grandiflorum by
the author and reported in Ambelania acidaby BOITEAU, ALLORGE, and SASTRE(1978).Inbothcasesthegermination wasepigeal;theshootapexand cotyledons
wereborne upwards bytheelongation ofthe hypocotyl.

GEOGRAPHIC DISTRIBUTION
Members of the tribe Ambelanieae are restricted to northern South America
withonlyRhigospiraknownfrom westoftheAndescordillera.GenericdistributionsofsixgeneraareillustratedinMaps 1-6. Distributionsofindividual species
areillustrated inthe Systematic Treatment.

CYTOLOGY
The author obtained the first approximate chromosome counts for the tribe
Ambelanieae. These meiotic counts, taken from pollen mother cells,are as follows:
SpeciesSpongiosperma macrophyllum
Molongum lucidum
Rhigospira quadrangularis

16

VoucherCount

Zarucchi 2504
Zarucchi 2483
Zarucchi 2428

n = ca.22
n = ca. 11
n = ca. 11

Agricuniv. WageningenPapers87-1 (1987)

Map 1
Ambelania

Map2
Mucoa

Map 3
Spongiosperma

Map 4
Molongum

Map 5
Rhigospira

Map6
Neocouma

Maps 1 - 6.Knowngenericdistributionsfor theAmbelanicae 1)Ambelania Aublet;2)Mucoa Zarucchi;3)Spongiosperma Zarucchi;4)Molongum Pichion; 5)Rhigospira Miers;6)Neocouma Pierre.

PALYNOLOGYPollencharacteristicsfor representativespeciesofAmbelanieaearesummarizedinTable2,andillustratedinPhotographs4and5.
The pollen of species in the Ambelanieae show differences not only among
theseveralgenerabutalsowithinthesamplesstudied.Mostofthepollensamplescontained grainswith avariable number of apertures, ranging from three
tofive,sometimeswithinthesamesample.Allofthespeciesexaminedhadmedium-sizedgrains(25-50urnindiameter),colporate,globose-oblateinequatorial
view,andpolygonalinpolarview.
IntheAmbelanieaethecolpiarerelativelyshortandextend usuallynomore
thanone-third ofthedistancetoeitherpoleofthegrain.Theaperturesinmost
speciessampled aresurrounded byanannulusorthickening ofthepollenwall
layers.Thesemaybequiteconspicuous,asinMucoaand Spongiosperma,making the grains appear polygonal in polar view.The pores within thecolpi areconspicuous and up to ca. 5-6 umin diameter. Thepollen grains ofallofthe
generaexceptNeocoumahaveapittedsurface.Thesefoveaeareusuallyscatteredand of varying densities. Pollen of Neocouma have a rugulate surface. Pollen
ofallmembersofthetribelacktectalprocesses.

TABLE2.Summary ofPollenData for Representative Speciesinthe Ambelanieae.
Species/Voucher (Origin)b

Ambelania acida
Irwinetal.55676 (Surinam)
Mucoa duckei
Cuatrecasas 7129(Colombia-Vaupés)
Spongiosperma grandiflorum
Pires&Black 553(Brazil-Parâ)Spongiosperma macrophyllumb
Zarucchi &Balick 1763 (Colombia-Vaupés)
Spongiosperma riparium
Schuttes &Lopez 10132 (Colombia-Guainia)
Molongum laxum
Ducke 44(Brazil-Amazonas)
Molongum lucidum
Zarucchi 2139 (Colombia-Vaupés)
Rhigospira quadrangularisb
Maas etal.P12717 (Brazil-Acre)
Zarucchi 2428 (Colombia-Vaupés)
Neocouma ternstroemiaceab
Ducke 220 (Brazil-Amazonas)

Pollen Size(um) a

Aperture

30.6-(33.9±1.8)-36.7

3(-4)-colporate

foveolate

32.9-(37.7+2.3}~42.7[N= 50](3-)4(-5)-colporate foveolate
4-5-colporate

foveolate(4-)5-colporate

foveolate

4-5-colporate

foveolate(4-)5-colporate

foveolate3(-4)-colporate

foveolate

3-colporate
3-colporate

foveolatefoveolate3(-4)-colporate

rugulate

37.8-(44.3+ 3.6)-52.4
39.0-(44.7+ 3.0)-50.0
36.6-(40.4±1.6)-43.9
32.9-(36.8+2.5)-41-5
39.0-(43.0±2.5)-48-8
29.3-(33.1±1.8)-36.7
26.8-(30.4±2.0)-35-4
43.9-(48.3±2.5)-53-6[N= 50]a

The size represents the equatorial diameter of 25 (or 50, if so noted) pollen grains mounted in glycerine jellusing light microscopy; the size range, mean, and standard deviation are given to the nearest 0.1 (im for eac
sample.
b
Illustrated bySEM micrographs inthepresent work.

18

Agric univ. WageningenPapers87-1 (1987)

PHOT. 4. Representative Pollen Grains of: A - B . Ambelania acida (Irwin et al. 55676; A. 1500 x ,B. 3750 x); C - D . Mucoa duckei (Cuatrecasas 7129;C. 1500 x ,D. 7500 x); E - F . Spongiosperma
macrophyllum (Zarucchi &Balick 1763;E. 1500 x , F. 7500 x).
Agricuniv. WageningenPapers87-1 (1987)

19

PHOT. 5. Representative Pollen Grains of: A - B . Molongum laxum (Ducke 44;A. 1500 x , B. 7500
x );C D . Rhigospira quadrangularis(Maas etal.P12717;C. 1500 x ,D. 7500 x);E - F . Neocouma
lemstroemiacea (Ducke 220;E. 1500 x , F.7500 x).

20

Agric univ. WageningenPapers87-1 (1987)

Theprimarilytetra-andpentacoloporatestructureofthepollengrainsfoundinMolongumandSpongiospermaindicatesthatthesetwogeneraaremorecloselyrelatedtoeachotherthantoanyofthefourothergeneraoftheAmbelanieae.
Ambelania, RhigospiraandNeocoumaallhaveprimarilytricolporategrains,yet
the structures of the colpi, ora, and exine make each genus distinct from theothers.Thepollen of Mucoa, likethat of Molongum and Spongiosperma, possessesa variable number ofapertures (4 ± 1),acharacteristic which supports
itsremovalfromAmbelania(sensustricto).Theregulatepollensurface structureofNeocoumagivesevidenceofitsdistinctionfrom theothergeneraofthetribe.
AMBELANIEAEPichonexBoiteau,Allorge&Sastre,Adansonia,sér.2,18(2):276.1978.
AmbelanieaePichon,Mém.Mus.Natl.Hist.Nat.24:165.1948sub'Ambélaniées',nom.nud.;WagenitzinMelchior,A.Engler'sSyllabusPflanzenfam. 412.
1964,nom.nud.
Smalllactescent shrubsorsmallormedium-sized treesupto30mtall;stems
teretetosubtereteorsharplyquadrangular.Nodesunilacunarwithasinglevascular trace;interpetiolar line orridgeconspicuous, appearing as a stipule-likescarwithglandsfound usuallyinabundanceandproducingaresinousexudate
withinandinaxilsofyounger leaves.Leavesdecussate,shorttolongpetiolate,glabrous,subcoriaceous to thick-coriaceous, sclereidseitherpresent or absent,
venationcamptodromous,marginsentireorcrenulateandusuallydistinctlyrevolute.Inflorescencesterminaloraxillary,few-tomany-flowered;pedunclevery
short to long, to ca 6cm; bracts and bracteoles small and scale-like'to large
andcalycine,pedicelsbibracteolate.Flowerspentamerous;calyxlobesquincunciallyimbricate,basallyconnateandeitherwithorwithoutglandswithin,persistentorcaducous, 1-14mmlong,glabrousorciliateonlyatapex.Corollasympetalous,actinomorphic,salverform,whiteorcreamtogoldenyellowororange,
tube narrow, pilose or glabrous within; lobes glabrous or pilose only at baseandadjacenttocorollaorifice,convoluteandoverlappingtoleftinbud,becomingreflexedatanthesis,equallingormuchshorterthantube.Stamens5,epipetalous,included;filamentsadnatetocorollatube;anthersintrorse,basifixed,bilocular,sessileor subsessile,basallobesauriculate or sagittate,apicallycaudate,openingbylongitudinal slits;pollenmedium-sized, 3-to 5-colporate,exinefoveolateor rugulate. Ovary glabrous,syncarpous, superior, bilocular, placentation axilewith many ovulesin each locule,basally nectariferous without, 1-5
mmlong.Styleterete,glabrous,shorttolong,surmountedbyarelativelylarge
stigmoidheadorclavuncle.Clavuncleapically5-lobedandbasallywithathickenedringorflange oftissue,glabrousorwithshort,densepubescenceonapicallobes; stigmoid lobes 2, short and blunt to long and terete, sometimesjoined
for most of their length.Fruit a few-to many-seeded, one-or two-celled berrywith seedseither embedded ina fluffy or stringy, aromatic pulp, or projecting
intoanopencavity,attachedbyfleshyfunicles;pericarpeitherleatheryorindurAgricuniv. WageningenPapers87-1 (1987)21

ated with a granular texture;fruit surface smooth to verrucose, sometimes withlongitudinal ribs,glands frequently conspicuous. Seeds generally plano-convex
and oblong, elliptic, or ovate in outline, 5-16 mm long; testa smooth, shiny
to dull, minutely reticulate or nearly so, to tortuous, spongy, pitted. Embryostraight, almost equalto seedinlength,embedded inan oilyendosperm; cotyledons small, plane, thin, ovate or circular; radicle cylindrical, terete, 3-8 times
aslongascotyledons.Germination epigeal.X = 11.
Typegenus.Ambelania Aublet, Hist. PI.Guian. 1:265-268,pi. 104.1775.
Genera. Ambelania Aublet, Mucoa Zarucchi, Spongiosperma Zarucchi, MolongumPichon, Rhigospira Miers,and Neocouma Pierre.
The tribe Ambelanieae was first described by PICHON (1948),comprising thegenera Ambelania, Molongum, Rhigospira, Neocouma and Macoubea, which heseparated from theCarisseae.He treated thesefivegeneraintwosubtribes:Ambelaniinae, with four genera, and Macoubeinae, with only Macoubea. These
tribaland subtribalnameslacked Latindiagnoses,andweretherefore illegitimatelypublished {nominanuda). Few botanists accepted thetaxonomiejudgmentsof PICHON regarding these genera and other aspects of apocynaceous classification;hisuntimely death in 1953may havecontributed tothis disregard.
The name Ambelanieae has been published according to the International
Rules,incomparative studiesof theTabernaemontanoideae and the Plumerioideae by BOITEAU & SASTRE (1975) and BOITEAU, ALLORGE & SASTRE (1978).
Thesetwoarticlesdealprincipallywiththecircumscriptionofthe Tabernaemontanoideae,intowhichtheyplaceMacoubea initsowntribe;four additional tribes
are recognized in the subfamily. In the second paper, BOITEAU et al.(1978) discuss characters which distinguish the Ambelanieae (Plumerioideae), as represented byAmbelania and Rhigospira, from the Macoubeeae (Tabernaemontanoideae), as represented by Macoubea. The Latin diagnosis for Ambelanieae
points out the following characters: 'Semina arillo destituta. Fructus carnosus
bilocularis sine strate indurato. Stamina sine textura movente sed caudis rudimentariisinstructa.' Thisdescription, however,isincomplete,and doesnot fully
describe the variation found within the tribe. The fruit iseither bilocular, witha fleshy dissepiment, or unilocular, wherein the septum tearsduring fruit maturation (e.g. Spongiosperma). The pericarp is indurated, with a granular consistency, in several of thegenera. Furthermore, theanthers are apically caudate
and the clavuncle has a basal ring or flange of tissue. These latter characterswerenoted byPICHON(1948)asbeingdiagnosticfor separatingthe Ambelanieae
from therelated tribe Carisseae.

KEYTOTHEGENERAOFAMBELANIEAE
1. Inflorescences axillary
Inflorescences terminal
2. Sepalseglandular at basewithin;young stems tereteto subterete;leaveswith
tertiary veinsobscureabove,lamina withmany filiform sclereids; inflores2223Agricuniv. WageningenPapers87-1 (1987)

cences usually developing several nodes from stem apex; fruit ovoid toelongate, sometimesribbed,pericarp leathery;seedsprojecting into open
cellsattachedbyfleshyfunicles
1. Ambelania
Sepalsmultiglandularinseveralseriesatbasewithin;youngstemscompressed;leaveswithtertiaryveinsvisibleabove,laminawithoutfiliformsclereids;
inflorescencesdevelopingaxillarytoandsimultaneouslywithterminalvegetativeshoots;fruitglobosetoellipsoid,lackingribs,pericarpinduratedwith
agranularcomposition;seedsembeddedinastringypulp . . . . 2. Mucoa
3. Youngstemssharplyquadrangular;leafvenationeucamptodromous;sepals
eglandular at basewithin;corollalobeslessthan one-half length oftube;
fruit pedicelfrustumic
5. RhigospiraYoungstemstereteorsubterete;leafvenationbrochidodromous;sepalswith
fewtomanyglandsatbasewithin;corollalobestwo-thirdstomuchgreater
thanlengthoftube;fruit pedicelcylindrical4
4. Corolla tube completely glabrous within, short, 3-9 mm;fruit few-seeded,
withthinpericarp,seedsembeddedinwhite,fluffy pulp . .4. Molongum
Corolla tube pilose within, short to long, 3.5-30 mm; fruit few- to manyseededwiththickandleatheryorcrustaceouspericarp
55. Bracteoles not calycine, borne usually on lower portion of pedicels; calyx
withbasalglandsinsingleserieswithin;corollatube10-30mmlong;leaves
withlateralveins 15-35perside;fruit one-celled,many-seeded;seedsnot
embeddedinpulp,attachedtofruit wallbylowfleshyfunicles
3. Spongiosperma
Bracteolescalycine, at or above middle ofpedicels;calyxwith basal glands
in several serieswithin; corolla tube 3.5-12mmlong;leaveswith lateralveins7-13perside;fruit two-celled,few-tomany-seeded;seedsembedded
inpulp
6. Neocouma
1. AmbelaniaAublet, Hist. PI.Guian. 1: 265-268,/?/. 104. 1775;Jussieu, Gen.
PI. 148. 1789; Lamarck, Tabl. Encycl. 1: 125-126, t. 169. 1791;Roemer &Schultes,Syst.Veg.4:545.1819;G.Don,Gen.Hist.Dichlam.PI.4: 102.1837
as'Ambellanid';Endlicher,Gen.PI.578.1838;Meisner,PI.Vase.Gen.262[parsprior], 171[parsaltera]. 1840;Oken,Allg.Naturgesch. 3(2):1049. 1841;A.De
Candolle,Prodr.8:328.1844;J.S.Presl,Wseob.Rostlin.2:1065.1846;MuellerArgoviensis in Martius, Fl. Bras.6(1): 15-18,pi.3-4. 1860;Bentham inBentham& Hooker, Gen.PI.2:694. 1876;Miers,Apocyn. S.Amer. 13-14,/»/. IB.
1878as'Ambellanid1; Bâillon,Hist. PI. 10:172-173. 1891[1889];Schumann inEngler&Prantl,Nat. Pflanzenfam. 4(2):123-124. 1895; Markgraf inPulle,Fl.
Surinam4(1):5-7. 1932;Markgraf,Notizbl.Bot.Gart.Berlin-Dahlem 15: 620.
1941;Monachino, Lloydia 8(2): 109-130. 1945;9(4):302. 1946;Pichon, Mem.
Mus.Natl.Hist.Nat.24:166-167,/»/.IV-11-13,21-23, 32-34. 1948; Lemée,Fl. Guy. Franc. 3:296. 1953;Macbride, Fl.Peru,Publ.Field Mus.Nat. Hist.,
Bot.Ser.13(5/1):373-374.1959;Schultes,J.Ethnopharm. 1:166-167.1979.
Willughbeja Scopoli ex Schreber in Linné, Gen. PL,ed. 8, 1: 162. 1789;J. F.
Agricuniv. WageningenPapers87-1 (1987)

23

Gmelin, in Linnaeus, Syst. Nat., ed. 13, 2(1): 434. 1791 as 'IVillughbeia' (nonRoxb.);Dietrich,SynopsisPlantarum 1:714.1839as'Willughbeid'(non Roxb.).
BenthecaNecker, Elementa Botanica 2:54.1790nonBenteca ÇBenteka') Adanson.
Smalltomedium-sized treeupto25mtall,lactescent,withyoung stemstereteor subterete;interpetiolar lineorridgeconspicuous,glandular within, becoming
resinouswithexudate.Leaves decussate,petiolate,glabrous,coriaceousto stiffcoriaceous, venation brochidodromous, margins entire and revolute, blade decurrent on canaliculate petiole; midrib plane to deeply impressed above, raised
to very prominent below; lateral veins inconspicuous and plane to deeply impressed above,plane toprominent below, especially near base of blade; tertiaryveins inconspicuous; foliar filiform sclereids abundant; small resin-producing
glands in leaf axils. Inflorescences axillary, cymose, few- to many-flowered; pedunclesmostly reduced; bracts ovate, small.Flowerssubsessile to short pedicellate;bracteoles small,ovate,near baseofpedicel;calyxlobes 5,persistent, quincuncially imbricate, basally lacking glands within; sepals free nearly to base,eciliate,ovate,apically rounded to obtuse;corollasympetalous, actinomorphic,
salverform, whitetoyellowish,thick;tubenarrow,straight,somewhatlong,withsuprastaminal region pilose, becoming dense near orifice, dilated at insertionof stamens; corolla lobes 5, convolute in bud, overlapping to left, becoming
reflexed atanthesis,approximating tubeinlength. Stamens 5,insertednear middle of tube; anthers lanceolate, subsessile, basally sagittate, apically caudate.
Ovary superior, conical-ovoid, glabrous, apically acute, bicarpellate, placentation axilewith ovules numerous in each locule;style terete, short; clavuncle cylindrical to ovoid or ellipsoid, with basal ring of tissue pentagonal in cross-section, apically five-angled, star-shaped incross-section, glabrous; stigmoid lobes
terete, thin, either subequal to or longer than clavuncle. Fruit a many-seeded,
two-celled berry,broadly ovoidtolanceovoid withseedsattached tothickdissepiment by fleshy funicles, projecting into two open cavities; pericarp leathery,
thick, surface smooth orwithlowtoprominent ribs. Seedsplano-convex, somewhat naviculate,ellipticto ovate inoutline;testa shiny todull,reticulate to very
lowandinconspicuouslyrugulose;hilumsmall,ovatetocircular.Embryo almost
aslongasseed,with small,thin,plane,ovatecotyledons;radiclestraight, terete,
about 5-8 timesaslongascotyledons.
Type species. Ambelania acida Aublet, Hist. PI. Guian. 1:265-268,;?/. 104.
1775.
E t y m o l o g y . From thenative Galibi name for thetree, 'Ambelani'.
D i s t r i b u t i o n . Widespread in Venezuela, the Guianas (Guyana, Surinam,
French Guiana) and Amazonia ofBrazil,Colombia, and Peru; Map 1.
AUBLET (1775) described Ambelania under 'Pentandria, Monogynia' of the
Linnaean sexual system, with the single species A. acida based on a collection
24

Agricuniv. WageningenPapers87-1 (1987)

MAP I.Ambelania

whichhemadeinFrench Guiana. Intheprotologue, AUBLETpresented along
discussion not only of the plant's appearance and salient structures, but alsoof itsuseby local peoplein the French colony. He said that thefruit isacidic
andediblewhendecorticated;thecortexactsasapurgative.
Ambelania and Mucoaare easily separated from the four remaining genera
intheAmbelanieae bytheiraxillary inflorescences andusually reduced peduncles.CharacterswhichareusedinthegenerickeyallowonetodistinguishAmbelaniafromMucoa,evenwhenonlysterilematerialisavailable.SomeofthemostinterestingcharacteristicsofAmbelaniaarethefoliar filiform sclereids,thetwocelled fruit with open cavities, and the seeds which project into these cavities
andareattachedtothefruit wallbyfleshy funicles.
All known collections ofAmbelania comefrom terrafirmaforest, including
individualsofthethreespeciestreatedinthepresentwork.
Agricuniv. WageningenPapers87-1 (1987)

25

KEY TO THE SPECIES OFAMBELANIA

1. Fruits ovoid, 8—12(—15) cm long, 6—9(—10) cm in diameter, with blunt apicalpoint; mature flower buds 17-20mm long; leaves narrowing toward base
(Peru;Rondônia and regionofManaus, Amazonas, Brazil)
1-2. A.duckei
Fruits elongate, narrowly elliptic to ovoid or lanceovoid, 4-12 cm long and2-4cmindiameter;matureflower buds24—27mmlong;leavesnot narrowingtoward base2
2. Leaf midrib plane to impressed above, raised below; lateral veins 15-25 perside;inflorescences few- to many-flowered (Widespread in Guianas, Eastern and Central Amazonia)
1-1.A.acida
Leaf midrib deeply impressed above, especially near base of blade, above
and very prominent below; lateral veins9—13(—16) per side; inflorescences
few-flowered (Western Amazonia ofBrazil,Colombia &Peru)
1-3. A.occidentalis

1-1.AmbelaniaacidaAublet,Hist.PI.Guian. 1:265-268,pi.104.1775; Lamarck,
Tabl. Encycl. 1: 125-126, t. 169. 1791;Roemer & Schultes, Syst. Veg. 4: 545.
1819; Descourtilz, Fl. Med. Antilles 7: 123-126, pi. 480. 1829; G. Don, Gen.
Hist. Dichlam. PI. 4: 102. 1837as 'Ambellanicr;A. De Candolle, Prodr. 8:328.
1844;J. S. Presl, Wseob. Rostlin. 2: 1065. 1846; Schumann in Engler & Prantl,
Nat. Pflanzenfam. 4(2): 124. 1895; Pulle, Ree. Trav. Bot. Néerl. 6: 285. 1909;
Markgraf in Pulle, Fl. Surinam 4(1): 6-7. 1932;Markgraf, Notizbl. Bot. Gart.
Berlin-Dahlem 15: 620. 1941;Monachino, Lloydia 8(2): 117-119. 1945; 9(4):302. 1946; Pichon, Mem. Mus. Natl. Hist. Nat. 24: 166-167, pi. I V - 11, 21,
32. 1948;Lemée,Fl. Guy. Franc. 3:296.1953.
Type: French Guiana: 'In sylvis Caiennae &Guianae.' Sep 1762-1764 (fl fr),
Aublet sn(BM,holotype;isotypes:LINN-SmithHerb.,P-RousseauHerb.).Homotypic synonyms: Willughbejaacida(Aublet) G. F. Gmelin, Systema Naturae
2(1):434. 1791;Willdenow, Linné Sp.Plant.,ed.4,1: 1231.1797;Smithin Rees,
Cyclopaedia Vol. 38. 1819;Sprengelin Linnaeus, Syst.Veg.,ed. 16,1: 672. 1825
as 'Willughbeia;'' Dietrich, Synopsis Plantarum 1: 714. 1839 as 'Willughbeia.''
Willughbeiaacida(Aublet) Oken,Allg.Naturgesch. 3(2): 1049.1841.
Heterotypic synonyms: Ambelania sagotii Mueller-Argoviensis, Linnaea 30:
389-390.1860as'A. sagotï; Markgraf inPulle,Fl.Surinam4(1):7.1932; Markgraf, Notizbl. Bot. Gart. Berlin-Dahlem 15:620. 1941.Types: French Guiana:
Acarouany, 1854(fl fr), Sagot 393(W,lectotype;isotypes:BM, BR, G [F photo
26844; F, GH, NY, US], K, MPU, P, S, U), and French Guiana: Cayenne.
1819-1821 (fl),Poiteau sn(paratypes:G, K, LE,W).
Ambelania tenuifloraMueller-Argoviensis in Martius, Flora Brasiliensis 6(1):16-17,pi.3.1860;Huber, Bol.Mus.ParaenseHist.Nat. 3:444.1902; Markgraf,
Notizbl.Bot.Gart. Berlin-Dahlem 15:620.1941.Types:Brazil:'insylvisprimaevis prov. Paraensis,' Sep [no year] (fl), Martius sn (lectotype: M; isotype: L).
26

Agricuniv. WageningenPapers87-1 (1987)

M 104.

jfmSe/aîuAAca/ii
FIG. 1.Reproduction ofOriginalIllustration oîAmbelania acidaAubletfrom hisHistoire desplantes
de la Guianefrançoise (1775): 1. corolla; 2. open corolla showing stamens; 3. separate stamen; 4.
calyx;5.gynoecium;6.ovary,stigma, style;7.fruit; 8.open fruit, partition, andseeds.

Brazil:Amazonas: 'incampisadVillainNovamda Rainha s. Topinambarana
prov. do Alto Amazonas,' Sep-Oct [no year] (fl), Martius sn (M, lectotype;
isotype: L). Brazil: Amazonas: Borba, Aug 1828 (fl), Riedel1368 (paratypes:
LE[2sheets,oneassn]).
Ambelania tenuifloravar. tenuiramea Mueller-Argoviensis in Martius, Flora
Brasiliensis 6(1): 17. 1860. Type: Brazil: Amazonas: Ega [= Tefé], Sep 1831
(fl fr), Poeppig 2604(W[Fphoto 34363;F, GH,NY, US],holotype;isotypes:
F872668,LE,NY,P).
AmbelaniacucumerinaMiers,Apocyn.S.Amer.13-14,pi.1B.1878as'Ambellania'inpart;Monachino,Lloydia8(2):120-122.1945.Type:Brazil:Amazonas:
Barra[= Manaus],Mar 1851 (fronly),Sprucesn(K,inMuseumsSpiritCollection, lectotype [here chosen]).The syntype, Spruce 2483 (K - Hook., cited as'2413'byMiers)isanisotypeofAmbelaniamacrophylla[= Spongiospermamacrophyllum].
Smallto medium-sized treeupto 25mtallwithleaf-bearing branches 2.5-5mm in diameter. Leaf blades elliptic to narrowly ovate (10-)12-20(-25) x
Agricuniv. WageningenPapers87-1 (1987)

27

3.5—8(—9)cm,glabrous,coriaceousto stiff-coriaceous, with(11—)15—25 pairsof
lateral veins; bases rounded to subacute and decurrent on petiole; apicesrounded to subacute becoming remotely acuminate; margins entire, revolute;
midrib plane to impressed above, raised below; lateral veins plane to slightly
impressed above, slightly raised below. Petioles 7—15(—18) mm long,canaliculate.Inflorescencesaxillary,cymose,(2-)4-l5(-50)-flowered witholderinflorescencesdeveloping a largemass ofclustered pedicels,with only several flowers
expanded at any one time; peduncles reduced, up to 8mm long; bracts andbracteolesovate,small,ca1x 0.8-1mm; pedicels1-3 mm long.Flowers:maturebud 24-27 mm long; sepals ovate, 1.8-2.8 x 1.5-2 mm, apically acute androunded.Corollawhitetoyellowwithtube10—12(—15)mmlong;lobesnarrowly
elliptic to lanceolate, 10-12 x 2-3 mm. Stamensinserted 4-5 mm from baseof corolla tube; anthers 2.5-3 mm long, subsessile with filament 0.2-0.4 mm
long, basally sagittate, apically caudate with acumen 0.2-0.3 mm long. Ovary
conical-ovoid, 1.8-2 mmlong;style 1-2 mmlong;clavuncle0.6-0.7 mmlong;stigmoidlobes0.8-1.8mmlong.Fruitnarrowlyellipsoidtoovoid,4~8(-12)cm
long,2-4 cmindiameter, round to ellipticor pentagonal incross-section, apicallyand basally rounded to subacute. Seedsnumerous,plano-convex, elliptic
toovateinoutline,8-11 x 5-7 x 2-3mm.
Etymology. Latinacida = acid,inreferencetotheedible,acidic fruit.
Distribution. Widely distributed from southeastern Venezuela and the
Guianassouth to the Rio MaracassuméinBrazilian Maranhäoand ineasternandcentralAmazonia;Map7.
Specimens examined:
VENEZUELA. Bolivar: 0-4 km N of El Pauji on trail to Uaipaur, 4° 30' N, 61° 35'W, alt 800-^900
m,4Nov 1985(fr),Liesner 19460(MO).
GUYANA.Waramuri Mission, Monika River, Pomeroon District, 23-27 Oct 1922(fl), Cruz 3068
(NY); Kaieteur Falls, Potaro River, 23 Oct-3 Nov 1923 (fr), Cruz 4423 (F, GH, MO, NY, US);
Mazaruni Station, 30 Jul 1940 (fl fr), Forest Dept. Br. Guiana3265 [F529J (K, NY); Waranama,
intermediate savannas Berbice River, 16Jun 1958(fr), Harrison 1147(K); 108/109 miles on Cattle
Trail, Liroppa Creek, Essequibo River, 5Jul 1920(fr), Hohenkerk 68A (K);Demerara River, May1888(fl),Jenman 4929(K);Hyde Park, Sep 1924(fr), Persaud 150(K, NY); Demerara River, Bootooba, Sand Hill, May 1923(fr), Persaud277 (F);Essequibo River, Moraballi Creek, near Bartica,
altnearsea-level, 16 Oct 1929(flbuds,fr.),Sandwith 465(K,NY, RB,U),30Oct 1929(fl), Sandwith537 (K, LE, NY, P, RB, S, U, US); basin of Kuyuwini River (Essequibo tributary), ca 150 miles
from mouth, 21-26 Nov 1937 (fl, yng fr), A. C. Smith 2581 (A, F, G, K, MAD, MO, NY, P, S,
U, US); Upper Mazaruni River Basin, Partang River, Merume Mountains, trail along SE side of
Merume Mtn., alt 1140 m, 5 Jul 1960 (fl fr), Tillen et al. 44822 (VEN); Upper Mazaruni River
Basin, Mt. Ayanganna, forest below talus of cliffs along NE side,alt 700-800 m, 4 & 17Aug 1960(flfr), Tillen et al. 45033(VEN).
SURINAM. Forest Reserve Zanderij I, tree no. 51, 1Dec 1915 (st), Boschwezen 1556(U), 23 Apr
1920(fr), Boschwezen 4660 (U); Forest Reserve Sectie0, tree no. 536, 5May 1916(fr), Boschwezen
1882 (U); Forest Reserve Wahamiri, tree no. 1653, 28 Oct 1919 (fl), Boschwezen 4441 (BR, U),
17 Feb 1920 (fl, yng fr), Boschwezen 4552 (BR, RB, U), 3 Jul 1920 (fr), Boschwezen 4713 (NY,U); Forest Reserve Kaboerie, tree no. 686, 6Nov 1920(fl), Boschwezen 4907 (BR, L, NY, U, US),
28 Jul 1922 (fr), Boschwezen 5932 (RB, US); Forest Reserve Brownsberg, tree no. 1176, 12 Oct
1923(fl), Boschwezen 6295 (K), 17Apr 1925 (fr), Boschwezen 6863 (U); Wilhelmina Gebergte, 3.5
28

Agricuniv. WageningenPapers87-1 (1987)

MAP 7.Ambelania acidaAublet.

km SSE of Juliana Top, 11.5 km N of Lucie River, alt 450 m, 8Aug 1963 (fr), Irwin et al. 54587(NY); Wilhelmina Gebergte, 2 km below confluence of Oost River, alt 225 m, 13 Sep 1963 (fl),Irwin el al.55676 (COL, F, GH, K, NY, S, U, US, VEN); Cottica River, mouth of the Patamacca
River, 10Aug 1933 (fr), Lanjouw 416 (U); S of road from Afobakka to Brownsweg at a pt. 8 km
from Afobakka, 24 Feb 1965 (fl buds), Lems 5253 (NA); Forests of Zanderij, 31 May 1916 (fr),Samuels sn (US); Sectie 0,Nov 1944(fl), Stahel 265 (GH, IAN, K, MAD, NY, S, U, WAG, WIS);
Zanderij I, May 1945 (fr), Stahel 265a (DS, GH, IAN, K, MAD, NY, U, UC, WAG, WIS); fluv.
Marowyne, Aug 1904 (fr), Versteeg 626 (RB, U); without specific locality, no date (fl), Hostmann
1317{G,K),nodate (fl) Weigeltsn(W).
FRENCH GUIANA. Camp Lorrain Chantier de la SOFOG terrain sain, plateau, 8 Jun 1953 (fr),
BAFOG 25M (P, U); Crique Balaté, 25 Jan 1951 (fr), BAFOG 5169 (P); Crique Sparouine Rive
droite (Maroni), 26 Mar 1952 (yng fr), BAFOG 6137 (P); Fleuve Ouaqui = Grigel, 4 Jul 1973(fr), Granville B.4882 (P); Haut Oyapock, Parcelle Zidock Paul, Itusâsây tap 1, 10 Feb 1975 (fr),Grenand750(P); SE of Saül, 3°37'N., 53° 12'W., ait 200 m, 11Feb 1978(st),Leeuwenberg 11779(NY); Cayenne, no date (fl), Martin sn (K); Bords de la Rivière du Maroni, 1861 (fr), Mélinon
14(P); Montfinery, May 1840(st),Mélinon 82(P); Mana, no date (fl buds), Mélinon sn (P); Route
Agricuniv. WageningenPapers 87-1 (1987)

29

du Gallion, après, pont sur tour de l'île, Feb 1975 (yng fr), Moretti 133 (P); Piste de St Elie, 16
Jun 1976 (fr), Moretti 481 (P); Saül, Monts La Fumée, 3° 37' N., 53° 12'W., ait 200-400 m, Moriet al. 15105(MO); Cacao, env. 60km Sde Cayenne, 11Jun 1965(fr), Oldeman 1351A (P); Grandcircuit Montagne Pelée, au km 1,400, 18 Aug 1971 (fr), Oldeman B.4025 (P); Saül, sur le circuitdelamontagneGrand Fossé,aukm0.900,28Oct 1971 (st),OldemanB.4161(P);Haute Approuague,dans le layon de la crique Parépou, 10Oct 1968 (fl), Oldeman T-205(P); St. Laurent du Maroni,
Sep 1910 (fl), Santini 4170 (L); Haut Maroni (Itany), Antecum-Pata (Malavate) rivière Marouini,
24Apr 1975(fr),Sastre etal.3931(MO,P);FleuveMaroniOukoumeMont Foret,entre Maripasoula et Antecum-Pata (Malavate), 25Apr 1975(fr), Sastre et al.3955 (MO, P, WAG); Rivière Inini,
afluent du Moyen-Maroni (Lawa) en amont de Maripasoula, 26 Apr 1975 (fr), Sastre et al. 3972
(MO, P); Fleuve Maroni, crique Charvein, 5 May 1975 (fr), Sastre et al. 4164 (MO, P); Village
Bellevue, 8 km d'Iracoubo vers St. Laurent, 13May 1975 (st), Sastre et al. 4192 (MO, P); Fleuve
Oyapock Saut Maripa, 6 Mar 1976 (fr), Sastre 4341 (P); Sinnamary, route de Ste. Elie, camp no.
2, 14Jul 1977 (fl fr), Sastre 5449 (P); Godebert, no date (fr), Wachenheim 0.17 (P), no date (oldfl), Wachenheim 202 (P), no date (fl), Wachenheim 403 (P); without specific locality, 1802 (fl, yngfr), Gabriel sn (F, G), no date (fr), Herb. Maire sn (P), no date (yng fr), Leprieur sn (P), no date(fr), Herb. L. C.Richardsn (P).
BRAZIL. Amapâ: Oiapoque, campo de aviaçào, 29 Sep 1949 (fl), Black 49-8192 (IAN, P); Rio
Jari, Cachoeira Mucurû, 0° 32' N, 53° 8' W, alt ca 140 m, 20 Aug 1961 (fl buds, yng fr), Egler&Irwin 46545 (IAN, K, MG, NY, S,US);Oiapoque, 30Jan 1950(fr), Fróes25720 (IAN); Cunani,
14Oct 1895 (fl), Huber MG 1044a (MG); Rio Oiapoque, ca 2.5 km SE of Clevelandia, 3° 48' N,
51° 53' W, 8 Oct 1960 (st), Irwin et al. 48654 (G, NY); Rio Araguari, 1° 40' N, 51° 56' W, alt
135 m, 29 Aug 1961 (fl), Pires et al. 50572 (F, IAN, K, MG, NY, S, US); Rio Araguari, Camp
13,1°45'N, 52°W, 9Oct 1961(fl),Pires etal.51643(IAN, MG, NY);Estrada de Cutias-Macapâ,16Jul 1980 (fr), Rabelo 516 (NY); Contagem entre Porto Platon e Serra do Navio, 10Oct-15 Dec
1976 (st), Rosa 1218 (MG). Amazonas: Tefé, 21 Apr 1970 (fr), Albuquerque &Lima 345 (INPA),
29 Sep 1904 (fl), Ducke MG 6829 (BM, G, MG), 11Jul 1973 (fr), Lieras et al. P16600 (F, INPA,
MG, NY, Z),28Nov 1959(fr), W. Rodrigues &Coêlho1425(INPA), 5Jan 1961(fr), W. Rodrigues
2059 (INPA); Manaus, Est. Manaus-Itacoatiara, km 12, 14Jul 1967 (fr), Cezärio & Vieira INPA
20612 (INPA); Manaus, km 10 da BR-17, Campos Sales, 30 Aug 1954 (fl), Chagas INPA 1 (F,INPA, MO); Manaus, Réserva Florestal Ducke, 14Mar 1957 (fl), L. Coêlho INPA 5157 (INPA,
MG, ÜB), 11Aug 1976 (st),A. OlivieraINPA 60537 (INPA), 19Jul 1963(fl buds), W. Rodrigues
5380 (INPA), 8Apr 1965 (fl), W. Rodrigues &Monteiro 6893 (GH, INPA), 11Sep 1979 (fl buds),
Zarucchi et al. 2534 (GH, INPA, NY); along the Rio Negro between Manaus and Sâo Gabriel,Acânga, 1° 10' S, 62° 30'W, 27Jun 1979(fr), Alencar et al. 92(NY);Mun. deTefé, Rio Solimôes,margemdireita, Estrada do Dendê, 18Oct 1982(flbuds),Amoral etal.121(MO);Mun.deAlvarâs,Rio Solimôes, margen direita, Lago de Alvarâs, 3° 13' S, 64° 50' W, 29 Nov 1982 (fr), Amaral
et al. 659 (MO); km 139 da estrada Manaus-Itacoatiara, 22 Jun 1972 (fr), L. Coêlho & Miranda
INPA 36021 (GH, INPA); Rio Negro, Barcellos, 11 Jun 1905 (fr), Ducke MG 7156 (MG); Rio
Madeira, Borba, 5 Jul 1936 (fl), Ducke 212 (A, F, K, MO, NY, S, US); Sâo Paulo de Olivenca,
18Apr 1945 (fl fr), Fróes 20751 (IAN, K, NY, US); Tapuruquara, 8 Apr 1947 (fl), Fróes 22113(IAN,NY);Rio Demeni,PostoIndigena Genipapo, 17Oct 1952(fl),Fróes&Addison 28989(IAN);
Manaus, Agricultural Experiment Station, alt 25 m, 13 Oct 1929 (fl), Killip &Smith 30007 (NY,US); Barro Alto, Parana do Ramos, 16 Mar 1924 (fl fr), J. G. Kuhlmann RB 21875 (RB, S, U,US);Environs de Manaus, 1906(fl),Labroy sn(P);Parana Autaz-Mirim, LocalJucâ, 25Mar 1973(fr), Loureiro etal.INPA 37638 (GH, INPA); Est. Torquato Tapajós, km 182,proximo ao igarapé
do Péreu, 4 Apr 1975 (fr), Loureiro et al. INPA 48424 (INPA); AM-1, km 139, 16 Jun 1972 (fr),Monteiro &Lima 141(GH, INPA), 16Jun 1972(old fr), Monteiro &Lima 144(INPA); km 65-70
daRodovia Manaus-Itacoatiara, 21 Oct 1963(fr), R. Oliveira2736(IAN,UB);Manaus andvicinity,
Tarumà road, 18km N of Manaus, 14 Mar 1967 (fr), Prance el al. 4628 (F, GH, INPA, K, MG,
NY, S,U, US);Estrada Manaus-Caracarai, km 39,Réserva Experimental de Silvicultura Tropical,
8 Sep 1977 (fl), Ribamar & Ramos 140 (INPA); Vila Iauareté, margem do Rio Uaupés, fronteira
Brasil-Colômbia,21May 1975(fr), Ribeiro 997(IAN, MG);Manaus, km9da BR-17,Ig.do Buiâo,
2 Oct 1959 (fr), W. Rodrigues & Coêlho 1333 (INPA, MO), 5 Oct 1962 (fl fr), W. Rodrigues &Chagas 4684 (GH, INPA), 14May 1963 (fr), W. Rodrigues & Chagas 5200 (INPA); Manaus, Ig.
30

Agricuniv. WageningenPapers87-1 (1987)

do Bindü, 5May 1961 (st), W. Rodrigues &Chagas 2402 (F); Manaus, Est. do Ig. do Passarinho,
23 Apr 1962 (fr), W. Rodrigues & Chagas 4402 (INPA); Est. Manaus-Itacoatiara, km 30, 2 Sep
1970 (fl), W. Rodrigues 8931 (GH, INPA); Manaus, Ponta Negra, Hotel Tropical, ârv. no. 91,19
Jul 1977 (fl), W. Rodrigues 9967 (INPA); Rio Negro, Rio Jauaperi, Estirào Tacuera, 18Feb 1977(fr), Santos 76(MG, NY); vicinity of Manaus, Cachoeira Baixa de Tarumâ, 11-14 Apr 1972 (fr),
Schuhes &Rodrigues 26136A (ECON, GH, INPA); Rio Negro, Manâos, Aug 1900 (fl), Ule5177(G, HBG, K, L, MG); Manaus and vicinity, Rosa de Maio, 12Sep 1979 (st), Zarucchi &Coêlho
2538 (GH, INPA, NY); Mun. de Borba, lower Rio Canumä, 2 km N of Vila de Canumä, Lat.
4° 02' S, Long. 59° 03' W, 29 Jun 1983 (fr), Zarucchi et al. 2903 (INPA, NY, US). Maranhâo:
EstradadaFeitoria,assuregion,23Apr 1933(fl),Fróes34(US);MaracassuméRiverregion, Estrada
da Feitoria, 20 Oct 1932 (fl, yng fr), Fróes 1956 (A, BM, F, G, K, MICH, MO, NY, P, S, US);Rio Maracassumé, Jul 1958 (fr), Fróes 34386 (IAN); Rio Alto Turiaçu, Nova Esperança, 2° 55'
S, 45° 45' W, ait 0-100 m, 29 Nov 1978 (fr), Jangoux &Bahia 41 (NY); Maracassumé, Oct 1932(st), Moses 55 (BM); Turiaçu km 6da BR-106, Maracaçumé Santa Helena, fazenda Maracaçumé
Agro Industrial, 28 Nov 1978 (yng fr), Rosa & Vilar2710 (F, NY). Para: Belém, grounds of IAN[= IPEAN & EMBRAPA], 30 Oct 1942 (fl, yng fr), Archer 7766 (IAN, NY, US), 18 Dec 1942(fr), Archer 8009 (IAN, MO, US),2Jan 1943(fr), Archer 8106(F,IAN, K), 8Feb 1943(fr), Archer
8234(IAN, MO), 15Feb 1952(fr), Black 52-14159(GH,IAN, INPA), 10Jan 1950(flfr), T. Guedes
246 (IAN, MO), 27 Dec 1962 (fr), E. Oliveira2367 (IAN), 19Jan 1947 (fl fr), Pires &Black 1269
(MO), 1Nov 1960 (yng fr), Pires 48899 (K, NY, US), 7Apr 1977(fr), Rosa 1757(MG, NY), Dec
1965 (fl), Schubert 2206 (IAN, US); Ruropolis Presidente Medici, 55° 04' W, 4° 00' S, alt 200 m,
6 Feb 1976 (fr), Bamps 5324 (BR, K, NY, U); Mun. Igarapé-Açû, casa de campo do Sr. Natan,
12Dec 1978(fl buds, fr), Bastos et al. 134(MG, NY, WAG); Rio Trombetas, km 109,access roadfrom Cachoeira Porteira to Perimetral do Norte, 27May 1974(fr), Campbellet al.P22293 (INPA,
NY, US,Z); Rio Trombetas, 3km Sof Cachoeira Porteira, 8Jun 1974(fr), Campbellet al. P22573(INPA, NY, Z);Benevides,estrada p/ Mosqueiro, 10Feb 1966(fr), Cavalcante&Elias 1447(MG);Mun. de Oriximinä a 200 m da estrada da cachoeira Porteira, paralelo ao Rio Trombetas km 6,
18Jun 1980 (fr), Cid &Ramos 1056(NY); Belém, Murutucü, 30 May 1902 (st), Ducke MG 2640
(MG); Obidos, 11 Jan 1905 (fr), Ducke MG 6959 (MG, RB); Faro, Castanhalda da Bôa Vista,
1 Feb 1910 (fr), Ducke MG 10629 (G, MG); Rio Cuminâ, mattas a E do Lago Salgado, 30 Aug
1910 (fl), Ducke MG 10899 (BM, G, MG, RB, US); Obidos, Uchyzal grande, 29 Sep 1915 (fl),Ducke MG 15760(MG); Rio Trombetas, Lago do Moura, 25Oct 1919(fl), Ducke RB 11397(RB);Rio Tapajoz, Villa Braga, 23 Sep 1922 (fl), Ducke RB 17458 (RB); matta da Pirelli, 12Mar 1966(fr), Elias de Paula 216 (MG); Rio Capim, 28 Mar 1949 (fr), Fróes & Pires 24171 (IAN, MO);
Castanhal, Colonia 3deOutubro, 8Dec 1949(fl),Fróes24886(IAN, INPA, MO, UC); Currupirû,localidade Gato, 22 Aug 1954 (fl), Fróes 31046 (MO); Regiâo do Anapü, Rio Flexal, Portel, 23
Jul 1956 (fr), Fróes 32901 (IAN); Belém, Marco, 20 Oct 1896 (fl), Huber MG 642 (MG); Belém,
estrada do Quinze,20Oct 1981(fl), Monteiro INPA 99007(INPA); Estrada de Ferro de Bragança,
Marituba, Belém,4Mar 1965(fr), E. Oliveira3248(IAN);Mosqueiro, 11Mar 1971(st),E. Oliveira
5501 (IAN); Belém, 29 Nov 1945 (fl fr), Pires &Black 732 (GH, IAN); FAO, no date (st), Pires
7408(IAN);vicinityof Belém,Sep-Oct 1961(st),Pires51874(NY,UB,US);W bank of Rio Pacaja,
opposite Ilha de Breu, 14 Oct 1965 (fr), Prance et al. 1622 (NY); 14 Oct 1965 (fl), Prance et al.
1624(F, IAN, K, NY, S, US); Road BR-22, Capanema to Maranhâo, vicinity of Cachoeira, km
96, 27 Oct 1965 (fl), Prance & Pennington 1717 (F, GH, IAN, K, NY, S, US); km 873 CuiabâSantarém, SerradoCachimbo,6Nov 1977(fr), Pranceetal.24941(MG);Ilhado Marajó, Curralinho, 28 Sep 1971 (fl), Ribeiro 169 (IAN); Igarapé-assü, 11Feb 1903(fr), R. S. Rodrigues MG 3377
(MG); Plantation de Paricatuba, route de Belém-Mosqueiro, 11 Mar 1968 (fl buds, fr), Sastre &Sastre 94(IAN,P,U);Acarâ,Jacaraquara, Tapera,21Feb 1966(fr), M. Silva540(MG); Oriximina,
beira estrada para Obidos, 26 Aug 1968 (fl fr), M. Silva 1735(MG); Santarém, km 35 da estrada
do Palhäo, 29 Aug 1969 (fl fr), M. Silva & Souza 2438 (F, K, MG, NY, S, U, US, Z); Belém,
Bosque Municipal, 18Jan 1948(fr), N. T. Silva 70(IAN);without preciselocality, 1915(fr), Curran
9 (US). Roraima: BoaVista-Caracarai road (BR 174),68km Sof BoaVista, 3Jan 1969(fr), Prance
et al.9532 (INPA, K, MG, NY, US); Rio Branco, margem estrada do Parana do Mararâ, 4 Mar
1977(fr), Santos 147(MG,NY).Without preciselocality:no date (st),A. R. Ferreirasn(P); 'Quinta

Agricuniv. WageningenPapers87-1 (1987)31

da BoaVista,SäoChristoväo, Rio-Jan.,Jan-Feb (fl), Glaziou 14078"(C)[probably apirated collection;seediscussionbelow].

Local names. GUYANA. Machieraro (Jenman 4929), Makurero (Arawak language,Hohenkerk 68A), Makuriro(ForestDept. Br.Guiana3265[F529],Makurriro {Sandwith 465, 537), Wild Cucumber (Persaud 277). SURINAM. Bat batti(Lanjouw 416),Batibati (Stahel 265), Makoriro (Stahel 265a),Wokorira (Stahel265). FRENCH GUIANA. Akusiwalapulu (Wayampi language, Grenand 750,
Sastre 4341), Ambalani (Galibi language, Sastre et al.4164),Ambelani (Galibi
language, Aublet sn), Graine biche (Créole language, Oldeman 1351A), Mapa
(BAFOG 25M, 5169, 6137, Sastre et al.3955, 3972), Papaya biche (Créole language, Granville B.4882, Oldeman T-205), Paraveris (Galibi language, Aubletsn), Quienbiendent (Créole language, Aublet sn), Taki-taki (Sastre et al. 3972).
BRAZIL. Molongó sem espinho (E. Oliveira 2367), Pau de colher (R. Oliveira2736), Pepino (Ducke MG 15760, Pires 51874, W. Rodrigues & Chagas 4684,
Schuhes &Rodrigues26136A, N. T.Silva 70, Zarucchi etal.2534),Pepino bravo(A. Oliveira INPA 60537, W. Rodrigues & Coêlho 1425, Rosa 1218, 1757, M.
Silva &Sousa2438),Pepino domato [Pipino,and other spellingvariants] (Albuquerque&Lima 345, Archer 7766,8009, 8106, Ducke MG 2640, MG 6959, MG
10629,RB 11397, RB 17458, 212, Elias 216, Fróes 1956, Loureiro et al. INPA
37638,Monteiro INPA 99007, Moses 55,E. Oliveira3248, Rabelo 516, W.Rodrigues &Coêlho 1333, W. Rodrigues 2059, W. Rodrigues &Chagas 2402, 5200,
M. Silva540,Zarucchi &Coêlho2538,Zarucchi etal. 2903),Pepinodoce (Amoralet al. 659, W. Rodrigues &Chagas 4402), Pepino verde (Cid &Ramos 1056),
Pipim(Archer 8234),Pipino (Prance&Pennington 1717).
Individuals of Ambelania acida are known from widespread areas from the
Guianas south to the Brazilian state of Maranhäo and westward into the Amazon basin asfar asTefé. Theyaresmall to medium-sized trees ofthelower storyof terra firma forest, and individuals of the species occur apparently in low frequency throughout its geographic range. The species is morphologically variable, sothat the study of only a fewcollectionsmight lead oneto interpret themas representing two or more species. The differences, however, in characterssuch as leaf shape, branch thickness, floral size,and fruit shape, are apparently
not correlated with each other. Some difficulty exists in the interpretation of
specimens of A. acida, as well as for other species in this tribe that have large
fleshy berries for fruits. Most fruiting collections have one or a few obviously
immature fruits. Thisbiasincollecting,whichisprobably due to thefruits' size,
abundance of thick and sticky latex-,and difficulty of preservation, poses problems for the interpretation of dried carpological material. The availability ofalcohol-preserved fruits and personal field studies aided greatly in the present
study.
One of the collections cited above, Glaziou 14078, was cited previously by
GLAZIOU (1910)asrepresentingAmbelania macrophylla. Thisspecimen contains
erroneous collection data, claiming to havebeen collected in thevicinity of Rio
32

Agricuniv. WageningenPapers87-1 (1987)

deJaneiro. WURDACK (1970)hasdiscussedtheincorrectinformation sometimes
associated with Glaziou-numbered specimens. Another Glaziou specimen (No.
14081) is cited under Spongiosperma macrophyllum in the present work, andmay actually represent a pirated specimen of Spruce 2483, the type collectionof the basionym: Ambelania macrophylla. Additional confusion between thesetwopresently-recognized taxa {Ambelania acidaand Spongiosperma macrophyllum) was caused by Ambelania cucumerina Miers (1878), since the species description was based on elements of both species. Ambelania cucumerina Miers
isherelectotypified bythespirit-preserved fruits intheKewMuseums collection.
The paratype (Kew-Hooker sheet of Spruce 2483;somefragments of this specimen are also attached to the BM duplicate with the'pencilled notation 'Ambellania cucumerina' in Miers' hand) is also an isolectotype of Ambelania macrophylla; seediscussion under Spongiosperma macrophyllum.

1-2. Ambelania duckei Markgraf, Notizbl. Bot. Gart. Berlin-Dahlem 12:
296-297. 1935; 15:620. 1941;Monachino, Lloydia 8(2): 119-120. 1945;Pichon,Mem. Mus.Natl. Hist.Nat. 24: 167,pi.I V - 12,22, 33.1948.
Type: Brazil: Amazonas: Manâos, Estrada do Aleixo, 2 Nov 1929 (fl fr),
Ducke RB22422(B,holotype,lost;isotypes:INPA 16429,RB nv).
Small tree up to 10 m tall with leaf-bearing branches 3-6 mm in diameter.
Leaf bladeselliptic to narrowly ovate or obovate, usually narrowing near base,(7-)10-20(-22) x (3-)4-8 cm, glabrous, coriaceous, becoming yellowish to orangewhendry,with 13-20pairsoflateralveins;basesrounded toobtuse,slightly
decurrent on petiole; apices obtuse to subacute.becoming remotely acuminate;margins entire, revolute; midrib slightly impressed above, raised below; lateralveinsmostlyinconspicuous,usuallynotraised.Petioles7-15mmlong, canaliculate. Inflorescences axillary, cymose, (2-)5-10(-25)-flowered, with only a few
flowers expanded at any one time and forming pedicellular masses at nodes;
peduncles reduced, up to 5mm long; bracts and bracteoles ovate, small, ca 1
x 0.8-1mm;pedicels2-4mm long.Flowers: mature bud 17-20mmlong;sepals
ovate, 1.5-2 x 1.4-2 mm, apically rounded to obtuse. Corolla yellowish withtube 9-12 mm long; lobes narrowly ovate to lanceolate, 8-10 x 2.5-3.5 mm.
Stamens insertedca3.5mmfrom baseofcorollatube;anthers3mmlong,subsessilewith filament ca0.3mmlong,basallysagittate,apicallycaudatewith acumen
0.5-0.6 mm long. Ovary conical-ovoid, 1.1-1.4mm long; style 1mm long;clavuncle 0.6-0.7 mm long; stigmoid lobes 1.2 mm long. Fruit ovoid, 8—12(—15)
cm long, 6-9(-10) cm in diameter, somewhat compressed laterally, apicallypointed, blunt.Seedsnumerous,plano-convex,elliptictoovateinoutline, 10-12
x 6-7 x 2-2.5 mm.
E t y m o l o g y . Named in honor of ADOLPHO DUCKE (1876-1959), collector
ofthetypespecimen and student oftheAmazon flora.

Agricuniv. WageningenPapers87-1 (1987)

33

FIG. 2. Ambelania ducket Markgraf: 1. branch, ] x ; 2 . older branch with inflorescences, | x ; 3 ,opened flower , 3 x ; 4. fruit with quarter-section removed showing seeds, | x . (1. Mathias &Taylor 5116;2 - 3 .Ducke451;4.Zarucchi etal. 2533,ale.mat.).

34

Agricuniv. WageningenPapers87-1 (1987)

MAP 8.Ambelania duckei Markgraf.

Distribution. Knownfrom easternPeru(Loreto,Pasco)andBrazil(AmazonasnearManaus,Rondônia);Map8.
Specimens examined:
BRAZIL. Amazonas: Manaus and vicinity, Réserva Florestal Ducke, 19 Aug 1968 (fl, yng fr),
Aluisio 93 (GH, INPA), 17Apr 1962 (fr), W. Rodrigues & Chagas4398 (INPA), 11Jun 1964 (fr),
W. Rodrigues & Loureiro 5903 (GH, INPA), 11 Sep 1979 (fr), Zarucchi et al. 2531 (GH, INPA,
NY), 2533 (GH, INPA, NY); Manaus, margem do Igarapé do Bindü, 11 Feb 1955 (fr), Chagas
INPA 801 (INPA); Estrada Manaus-Caracarai, variante da BR-174a Manaus-Itacoatiara, 22 Dec
1976 (st), D. Coêlho &Mouräo 875 (INPA); Manaus, BR-17, km 21,28 Dec 1955 (fr), L. Coêlho
& Mello INPA 3227 (INPA, MG); Manaus, Rio Tarumâ, 15 Mar 1936 (fl fr), Ducke RB 35153(G, INPA, K, P, S, U, US), 451 (A, F, K, MO, NY, RB, S, US [frag]), 1Oct 1941 (fl fr), Ducke
451 {IIa Col.) (IAN, MG); basin of Rio Negro, without precise locality, 19 Apr 1941 (st), Fróes
12004/5 (A, F, MICH, NY, US);Manaus, Estrada do Aleixo,26Mar 1947(fl),Fróes22046 (IAN,
NY); Estrada Manaus-Itacoatiara, km 150, 14May 1972(fr), Loureiro et al.INPA 35786 (INPA);
Agricuniv. WageningenPapers87-1 (1987)

35

Manaus, terreno do SIDERAMA, 5May 1973(fr), Loureiro et al. INPA 38014 (INPA); Manaus,
Ig. do Mariano, 31Jul 1956(fr), Mello INPA 4015(INPA, MG, MO, U);km 28do Rodôvia Manaus-Itacoatiara, 30 Oct 1963 (fr), E. Oliveira2829 (IAN); Manaus-Itacoatiara Highway, Réserva
Walter Egler, km 65, 14 Nov 1966 (fr), Prance et al. 3173 (INPA, NY); Est. Manaus-Caracarai,
km 23, 6 Nov 1959 (fl fr), W. Rodrigues & Côelho 1361 (INPA); Manaus, Ig. do Bindü, 20 Apr
1961 (fr), W. Rodrigues & Chagas 2403 (INPA); Manaus, Estrada do Forquilha, 5 Jul 1962 (fr),
W. Rodrigues & Chagas 4528 (INPA); Manaus, Ponte da Bolivia, 18Oct 1955 (st), W. Rodrigues
INPA 2159 (INPA); Manaus, Jardim Tarumäzinho, 11Feb 1977(fr), M. Silva et al.2081 (INPA).
Rondônia: 4 km de Vilhena, 12°45' S., 60° 10'W, 30Oct 1979 (fl fr), Vieiraet al. 798(MO, NY,US).
PERU. Loreto. Rio Aguatia, W bank 8-10 km below Aguatia, 2 Jul 1960 (fr), Mathias & Taylor
5116 (F, LA, MO, US). Pasco: Prov. Oxapampa, Cabeza de Mono, Rio Iscozacin, 10km SW of
Iscozacin, Palcazu Valley, 10° 20' S., 75° 18' W., alt 320 m, 8 Jun 1983 (st), Gentry et al. 41656(MO), 8Jun 1983(fr only),Gentry etal.41693(MO), 13Jun 1983(st),Gentry etal.41976(MO).

Local names. BRAZIL. Pepino(W.Rodrigues &Chagas4528),Pepinodo mato
[damata](D. Coêlho&Mourào 875, W.Rodrigues &Coêlho1361, W. Rodrigues& Chagas 2403, 4398), Pepino bravo (W. Rodrigues &Loureiro 5903), Pepinoverde (D. Coêlho &Mouräo 875, Zarucchi et al. 2531, 2533), Pepino verde damata (Chagas INPA 801, L. Coêlho&Mello INPA 3227). PERU. Auca-sanango(Mathias &Taylor 5116).
Ambelania duckeiwasdescribedbyMARKGRAFin 1935,basedonasinglecollection by ADOLPHO DUCKE from the vicinity of Manaus in central Amazonia.
IntheManaus region,Ambelania duckeiissympatricwithA. acida,andisdistinguished easily by its large ovoid fruit. It is an understory tree up to 10m tall;
I found it growing within a short distance of a small blackwater stream, wellshaded, and insandy soilon terra firma.
In the protologue, MARKGRAF (1935) compared this species with Ambelania
tenuiflora (= A. acida), also found in the vicinity of Manaus, observing thatthe septum which divides the fruit into two cells tears during fruit maturationinA. duckei, yetremainsintact intheother species.Myownstudiesof Ambelania, especially A. duckei in the field near Manaus, contradict this observation,
showing that the fruit dissepiment remains intact in all species of Ambelania.
Many seeds are attached to thiscentral partition by fleshy funicles, and project
into the open cells. Another interesting feature of this species is that the leaves
generally turn yellowish topaleorangewhen dried.
According to MARKGRAF (pers. comm.), the holotype of Ambelania duckei,
Ducke RB 22422, at Berlin, was lost during World War II. It is probable that
aduplicateofthat collectionisdeposited intheherbarium oftheJardin Botanico
in Rio de Janeiro (RB), but it was not seen among material provided on loan
from that institution. If it does exist, it should be designated as the lectotype
for thespecies.
1-3. AmbelaniaoccidentalisZarucchi, sp.nov.
Type:Peru:Loreto:Mishuyacu, nearIquitos,alt 100m,forest, Oct-Nov 1929(fl), Klug 613(US 1455612,holotype;isotypes:F 629963,NY).
36

Agricuniv. WageningenPapers87-1 (1987)

Plantae Ambelaniae acidae etA. duckei affinis seddiffert foliis ellipticis,angusteovatis,basirotundatis velobtusis, nervomediano supra valdeimpresso,
subtusbasimversusprominente;inflorescentiisbrevoribuscompactis paucifloris; fructu in ambitu anguste ovato vellanceolate ovato, 7-12 cm longo, 2.5-4
cmindiametro.
Smalltreeup to 10mtallwith leaf-bearing branches 2-3.5mmin diameter.
Leafbladeselliptictonarrowly ovate,(ll-)14-22(-25) x (4.5-)6-9(-10.5)cm,
glabrous, coriaceous to stiff-coriaceous, sometimes becoming yellowish when
dry,with9—13(—16) pairsoflateralveins;basesbroadlyrounded to obtuseanddecurrentonpetiole;apicesacutebecomingremotelyacuminate;marginsentire,
revolute;midribdeeplyimpressed,especiallynearbaseofblade,aboveandvery
prominent below; lateral veins inconspicuous or slightly raised to deeply impressednearbaseofbladeabove,slightlyraisedtoprominentnearbaseofblade
below. Petioles12—18(—23)mm long, canaliculate. Inflorescences axillary, cymose,(l-)2-5-flowered; pedunclesreduced,upto2mmlong;bractsandbracteolesovate,small,ca 1 x 0.6-0.8mm;pedicels 2—4 mmlong.Flowers: maturebud 25 mmlong;sepalsovate,1.8-2 x 1.3-1.6mm, apicallyroundedtoobtuse.
Corolla white with tube 9-10 long; lobes elliptic to narrowly ovate, 13-15 x
4-5 mm. Stamensinserted ca 3mm from base of corolla tube; anthers 3-3.5mm long, subsessile with filament ca 0.4 mm long, basally sagittate, apicallycaudatewith acumen 0.5mmlong. Ovary conical-ovoid, 2-2.5 mmlong;style
2mmlong;clavuncle0.7mmlong;stigmoidlobes0.5mmlong.Fruitnarrowly
ovoidorellipsoidtolanceovoid,7-12cmlong,2.5-4cmindiameter,somewhat
compressed laterally, apically rounded, basally rounded to truncate. Seedsnumerous,plano-convex,elliptictoovateinoutline,9-11 x 5.5-7 x 3-3.5mm.
Etymology. Latinoccidentalis = ofthewest,inreferencetothegeographic
distributionofthespeciesinwesternAmazonia.
Distribution. Known from western Amazonia in thevicinity of thecommonborderofColombia,Brazil,andPeru.Collectedfromforestonterrafirma;
see Map9.
Specimens examined:
BRAZIL.Amazonas: SàoPaulodeOlivença, 21 Jan 1949(fr),Fróes23949(IAN);Mun. Sào Paulode Olivença, near Palmares, 11Sep-26 Oct 1936 (fr), Krukoff 8135 (NY); Tabatinga, 30Nov 1945(fr), Pires &Black 958 (IAN); Rio Mutum, 11 Nov 1975 (fr), Rosa &Coêlho 538 (IAN); vicinityof Leticia (Colombia), Brazil side, near El Marco, alt 200 m, 7 Sep 1963 (fr), Soejarto 582 (COL,
ECON,K, US).
COLOMBIA. Amazonas: Rio Amazonas, vicinity of Leticia, 29 Aug-12 Sep 1966 (fr), Schuttes etal.24074(ECON);LaVictoria ontheAmazon River [as'Peru' onlabel],2Sep 1929(fr),LI. Williams
3017(F).
PERU. Loreto: University arboretum on Rio Nanay, 7 Aug 1972 (fr), Croat 18866 (MO, NY,
P, U, Z); Puerto Almendra SW of Iquitos on Rio Nanay, 9 Aug 1972 (fr), Croat 19042 (MO),12Oct 1976(fr), Revilla 1463(MO),29Nov 1976(fr), Revilla 1965(MO);Prov. Maynas, Mishana,
Rio Nanay, halfway between Iquitos and Santa Maria de Nanay, alt ca 140m, 14Aug 1978 (fr),Diaz et al. 386 (MO), 30 May 1978 (fr), Gentry et al. 22359 (MO), 25 Feb 1981 (fr), Gentry etAgricuniv. WageningenPapers87-1 (1987)

37

MAP9.AmbelaniaoccidentalisZarucchi.
al. 31719(MO,USF), 1979(fr), Ramirez 75 (MO);Mishuyacu, near Iquitos,alt 100m,Oct-Nov
1929(fr), Klug 21(NY, US);RioJavari behind Angamo Garrison, 4Aug 1973(fr), Lieras etal.
P17156(INPA,NY);Maynas,Dtto.AltoNanay,RioNanay,QuebradadeAnguilla,28Mar1978(fr), Rimachi3482 (F,NY);Maynas,Estación Biologica Callicebus-Mishana-RioNanay, 2horas
rioarriba de Iquitos, 3° 55' S., 73° 35'W., ait ca 130m, 25 Oct 1980(fl fr), Vàsquez et al. 616(MO,US).

Local names. PERU.Cucharahuayo{Vàsquezetal. 616), Mashomicuna(Rimachi3482).
Ambelania occidentalisisheredescribedasanewspeciesbasedupontwenty
collectionsfromwesternAmazoniaofBrazil,Colombia,andPeru.Itisseparated readily from the other two speciesin Ambelania by the fruit shape and bytheleaveswhichhavethemidveindeeplyimpressedaboveandveryprominent
below.
38

Agricuniv.WageningenPapers87-1 (1987)

Excluded Speciesof Ambelania
Ambelania edulis(Roxb.) 3.Presl,Wseob. Rostlin. 2: 1065.1846 = Willughbeia
edulisRoxb.(Apocynaceae) according to MERRILL(1950).
Ambelania cuneata Muell.-Arg. in Martius, Flora Brasiliensis 6(1): 17. 1860 =Molongum lucidum(Humboldt, Bonpland &Kunth) Zarucchi.
Ambelania laxa (Bentham) Muell.-Arg. in Martius, Flora Brasiliensis 6(1): 17,
t.4. 1860[based on Tabernaemontana laxa Bentham,J. Bot. (Hooker) 3:244.
1841] = Molongum laxum (Bentham) Pichon, Mem. Mus. Natl. Hist. Nat.
24: 168.1948.
Ambelania macrophylla Muell.-Arg.inMartius,Flora Brasiliensis6(1): 18.1860
as'macroyhylld = Spongiosperma macrophyllum (Muell.-Arg.) Zarucchi.
Ambelania quadi-angularis Muell.-Arg. in Martius, Flora Brasiliensis 6(1): 18.
1860 = Rhigospira quadt-angularis (Muell.-Arg.) Miers, Apocynaceae SouthAmerica 68,/?/. 70,4.1878.
Ambelania grandiflora Huber, Bol. Mus. Paraense Hist. Nat. 3: 444. 1902 =Spongiosperma grandiflorum (Huber) Zarucchi.
Ambelania camporum Glaziou, Bull.Soc.Bot. France, sér.4(mém. 3e), 10:449.
1910 = Secondatia densiflora A. DC. (Apocynaceae), according to PICHON(1948).
Ambelania zschokkeiformis Markgraf, Notizbl. Bot. Gart. Berlin-Dahlem 12:
295-296. 1935 = Molongum zschokkeiforme (Markgraf) Pichon, Mém. Mus.
Natl. Hist.Nat. 24: 169.1948.
Ambelania lucida (Humboldt, Bonpland & Kunth) Markgraf, Notizbl. Bot.
Gart. Berlin-Dahlem 15:620. 1941[based on Tabernaemontana lucida Humboldt, Bonpland &Kunth, Nov.Gen.etSp.7:162(folio), 209(quarto). 1825]= Molongum lucidum(Humboldt, Bonpland &Kunth) Zarucchi.
Ambelania markgrafiana Monachino, Lloydia 8(2): 122. 1945 [name based on
Neocouma duckeiMarkgraf,Notizbl.Bot.Gart. Berlin-Dahlem 11:337.1932]= Mucoa duckei (Markgraf) Zarucchi.
Ambelania ternstroemiacea (Muell.-Arg.) Monachino, Lloydia 8(2): 123. 1945[based on Tabernaemontana ternstroemiacea Muell.-Arg. in Martius, Flora
Brasiliensis6(1):88.1860] = Neocouma ternstroemiacea (Muell.-Arg.)Pierre,
Bull.Mens.Soc.Linn.Paris,sér.2,5:33-34.1898.
Ambelania lopezii Woodson in Schultes, Bot. Mus. Leafl. 15(2): 76. 1951 =Spongiosperma macrophyllum (Muell.-Arg.) Zarucchi.
Ambelania oleifolia Monachino, Mem. New York Bot. Gard. 10(1): 117-119,
fig. 14a-d.1958as 'oleaefolia"= Spongiosperma oleifolium (Monachino) Zarucchi.
Ambelania oleifolia Monachino var. ripariaMonachino, Mem. New York Bot.
Gard. 10(4):59. 1961as 'oleaefolid' = Spongiosperma riparium (Monachino)
Zarucchi. This variety of Ambelania oleifolia, as described by MONACHINO(1961), contains two taxa: Spongiosperma oleifolium and S. riparium. For acompleteexplanation, seethediscussion under Spongiosperma oleifolium.
Ambelania longiloba Markgraf in Prance, Brittonia 23(4): 441. 1971 = Spongiosperma longilobum(Markgraf) Zarucchi.
Agricuniv. WageningenPapers87-1 (1987)

39

Ambelania lanceolata Markgraf, Acta Bot. Venez. 6: 74. 1971 [1972] = Spongiospermaoleifolium (Markgraf) Zarucchi.
Ambelaniapatchenkoana Markgraf, Acta Bot. Venez. 10:247-249,/ïg. 1. 1975
= Mucoapantchenkoana (Markgraf) Zarucchi.
Ambelania parviflora Markgraf, Ernstia 19:4-5. 1983 = Neocouma parviflora(Markgraf) Zarucchi.

2. MucoaZarucchi, gen.nov.(Apocynaceae;Plumerioideae; Ambelanieae).
Ambelania sect. Neocoumae Monachino subsect. Markgrafianae Monachino,
Lloydia 8(2): 113-114.1945,nom. nud.
Ambelania sect. Neocoumopsis Pichon, Mém. Mus. Natl. Hist. Nat. 24: 167, pi.
IV-13,23,34.1948, nom. illeg.
Ramijuniores compressi, vetustiores teretes,in foliorum axillis multiglandulosi. Nervi tertiarii laminae foliorum supra manifesti, sclerenchymis filiformis
destituti.Inflorescentia cymosa,cymisaxillaribussurculisvegetativissimulcrescentibus. Sepala intus basi multiglandulosa. Antherae sessiles.Baccae globosaevelellipsoideae,pericarpioindurato granuloso.Seminapulpa fibrosa circumdata.
Smalltomedium-sized treeup to 15m tall,lactescent,withyoung stemscompressed, becoming teretewith age;interpetiolar lineor ridgeconspicuous, glandularwithin,becomingresinouswithexudate.Leavesopposite,decussate,petiolate,glabrous,coriaceoustothick-coriaceous,venationbrochidodromous, marginsentireand revolute, bladedecurrent oncanaliculatepetiole;midrib slightly
raised to impressed above, slightly raised to very prominent below;lateral veinsslightly raised to impressed above, raised below; tertiary veins visible above;
foliar filiform sclereids absent; small resin-producing glands numerous in leaf
axils.Inflorescences axillary,developingsimultaneouslywithterminal vegetative
shoot, cymose, few- tomany-flowered; peduncles mostly reduced; bracts ovate.
Flowerssubsessiletoshortpedicellate;bracteolessmall,ovate,nearbaseofpedicel; calyx lobes 5, persistent, quincuncially imbricate, basally with many small
glandsinseveral serieswithin; sepalsfree nearly tobase,eciliate,ovate, apicallyrounded; corolla sympetalous, actinomorphic, salverform, white or yellow toorange,thick;tube narrow, straight, either short or somewhat long,with suprastaminal regionpilose,becoming densenear orifice, dilated atpoint of insertionof stamens; corolla lobes 5, convolute in bud, overlapping to left, becoming
reflexed at anthesis, usually approximating tube in length. Stamens 5, inserted
just belowmiddleoftube;anthers lanceolate,sessile,basally auriculate, apicallycaudate. Ovary superior, ovoid to conical-ovoid, glabrous, apically acute to attenuate, bicarpellate, placentation axile with ovules numerous in each locule;style terete; clavuncle cylindrical to globose, with basal ring of tissue, apically
five-lobed, star-shaped in cross-section, glabrous; stigmoid lobes terete, thin,either subequal to or longer than clavuncle. Fruit a many-seeded, two-celled
40

Agricuniv. WageningenPapers87-1 (1987)

berry,globoseorellipsoid,withseedscompletelyembeddedinastringy,aromaticpulpand acropetally imbricateoncentralplacentas;pericarpindurated withagranularcomposition,surfacesmoothtoverrucose.Seedsplano-convex,naviculate,ovatetoellipticinoutline;testashiny,finelyreticulate;hilumsmall,ovate
toelliptic.Embryo almostaslongasseed,withsmall,thin,plane,ovatecotyledons;radiclestraight,terete,about4-7timesaslongascotyledons.
Typespecies.Mucoaduckei(Markgraf) Zarucchi(basedonNeocoumaduckei
Markgraf,Notizbl.Bot.Gart.Berlin-Dahlem 11:337.1932),herechosen.
Etymology. An anagram of Couma of the apocynaceous tribe Carisseae,
agenuswhichsharesmanycharacterswiththeoneheredescribed.
Distribution. CentralandwesternAmazoniaofBrazil(Amazonas,Roraima),Peru (Loreto), Colombia (Amazonas, Guaviare,Vaupés),and Venezuela(Amazonas,Bolivar);Map2.

MAP 2. Mucoa
Agricuniv. WageningenPapers87-1 (1987)

41

The new genus Mucoa, as here circumscribed, consists of two species from
central and western Amazonia. In 1948,PICHONdescribed Ambelania sect. Neocoumopsis[nom.illeg.]with a single species,A. markgrafiana. This species, firstdescribed by MARKGRAF in 1932 as Neocouma duckei and later transferred toAmbelania {A.markgrafiana, not A. duckei) by MONACHINO(1945),is discussed
indetailunder M. duckei, seebelow.
The distinction between Ambelania and Mucoa isclear, based on gross morphological features and on several less conspicuous characteristics such as theabsence of foliar filiform sclereids and the nature of the calyx lobes, which are
multiglandular atthebasewithin.ThetreesofM. duckeiand M. pantchenkoana
are found in terra firma forest, and, based on specimen records and personalobservations, areofrather rare occurrence.

KEY TO THE SPECIES OF

MUCOA

1. Petioles long, (12-)16-36(-42) mm long; inflorescences 5 - to 12(-20)-flowered, corolla yellow to orange with tube 12-15 mm long; fruit globose;leaf blades elliptic to narrowly ovate or obovate, (14-)18-25(-30) x
7—14(—17) cm (Widespread in lowlands of Western and North Central
Amazonia ofBrazil,Colombia&Peru)
2-1. M.duckei
Petioles short, 2-4 mm long; inflorescences 3- to 5-flowered, corolla whitewith tube 8-10 mm long; fruit (immature) broadly ellipsoid; leaf blades
elliptictoobovate, 10-16 x 4.5-8 cm(Higherelevationsalong VenezuelaBrazilborder)
2-2.M. pantchenkoana

2-1. Mucoaduckei(Markgraf) Zarucchi,comb.nov.
Basionym: Neocouma duckei Markgraf, Notizbl. Bot. Gart. Berlin-Dahlem
11:337.1932.
Type:Brazil:Amazonas:Fontebôa, 5Sep 1929(fl),Ducke RB22418(B,holotype,lost; RB,lectotype [herechosen];isotypes:K, P [Fphoto 38761;GH,US],
S,U,US 1574340).Homotypicsymonym:Ambelania markgrafiana Monachino,
Lloydia 8(2): 122-123. 1945; 9(4): 302. 1946; Pichon, Mém. Mus. Natl. Hist.
Nat.24:167,pi.IV-13,23,34.1948; Schultes,Bot.Mus.Leafl. 13(10):308-309,
pi. 37. 1949;Macbride, Fl. Peru, Publ. Field Mus.Nat. Hist., Bot. Ser. 13(5/1):
373-374.1959.
Small to medium-sized tree up to 15 m tall with leaf-bearing branches 4-7
mm in diameter, young stems compressed with two longitudinal grooves, becoming terete with age. Leaf blades elliptic to narrowly ovate or obovate,(14—)18—25(—30) x 7—14(—17)cm,glabrous,coriaceoustothick-coriaceous, with
14-20pairsoflateralveins;basesobtusetorounded,decurrent onpetiole;apicesrounded to obtuse, usually mucronate; margins entire, revolute; midrib plane
42

Agric univ. WageningenPapers87-1 (1987)

FIG. 3.Mucoaduckei(Markgraf) Zarucchi: 1.flowering and fruiting branch,| x ; 2 . young flower
insection,3 x; 3.sectionedfruit,| x. (1-3.Zarucchietal.1852,incl.ale.mat.).
Agricuniv. WageningenPapers87-1 (1987)43

becoming impressed near base of blade above, very prominent below; lateralveins raised above and below. Petioles (12—)16—36(—42) mm long, canaliculate.
Inflorescences axillary, cymose, 5-12(-20)-flowered; peduncles completely reduced orupto8mmlong;bractsbroadly ovate,2-3 x 1.5mm;pedicels 2-5(-8)
mm long;bracteoles ovate, small,ca0.8-1 x 0.7-0.9mm. Flowers:mature bud
20-25mmlong;sepalsbroadlyovate,2-3 x 2-2.8mm,apicallyrounded. Corolla yellow to orange with tube 12-15 mm long; lobes elliptic, 8-11 x 3-4 mm.
Stamens inserted ca 4 mm from base of corolla tube; anthers 3.5—4mm long,sessile,basally auriculate,apicallycaudatewithacumen0.4-0.5mmlong. Ovary
conical-ovoid, 2-3 mm long, style 1.8-3 mm long; clavuncle 0.8-1 mm long;stigmoid lobes 1.3-1.8mmlong,thin,terete,fused for most oftheirlength.Fruit
globose, 5—8(—10)cm in diameter. Seeds numerous, ca 50-70, plano-convex,
ovatetoellipticinoutline,9-12 x 5-6.5 x 2.5-3.5 mm.
E t y m o l o g y . Named in honor of ADOLPHO DUCKE (1876-1959), collector
ofthetypespecimen and student oftheAmazon flora.
D i s t r i b u t i o n . WidespreadinwesternandnorthcentralAmazoniaofBrazil,
Colombia, and Peru; expected to occur in adjacent areas of Venezuela; Map
10.
Specimens examined:
COLOMBIA. Amazonas: Trocha entre El Encanto y La Chorrera, alt ca 180 m, 31 May-2 Jun
1942(fr), Schuhes 3877(ECON); Rio Caraparanâ entrelasbocas yElEncanto, altca 150m, 22-28
May 1942 (fr), Schuttes 3877a (ECON). Vaupés: Margenes del Rio Vaupés en Pucarón, alt 240
m,2Oct 1939(fl fr), Cuairecasas 7129(COL, F,US);Mitüandvicinity, along RioVaupés between
Rio Yi and Rio Kubiyu, 17 Jul 1976 (fl fr), Zarucchi et al. 1852 (COL, ECON, GH, MO, US,
USF).
PERU. Loreto: Prov. Requena, Dist. Sapuena, Jenaro Herrera, ârbol no. 540,alt 120m, 30 May
1974(st),Abadie 79r(IAN);JenaroHerrera,margen derecha RioUcayali,24May 1982(fr), Encarnaciim 26158 (MO, US); Jenaro Herrera, arboretum, 7 Dec 1977 (fr), Gentry et al. 21215 (MO);Jenaro Herrera, tree no. R-114 PP, alt 120 m, 15 Oct 1973 (fl buds, yng fr), Noriega sn (COL,
F,NY).
BRAZIL. Amazonas: Lago do Castanho-Mirim, estrada da Petrobrâs, 27 Jun 1973 (fl buds, fr),
Albuquerque et al. 917 (INPA); along Rio Negro between Manaus and Sào Gabriel, Acânga, 1°
10' S.,62° 30'W., 27Jun 1979(fr), Alencar etal.84(INPA, MO); Rio Tea, afluente do Rio Negro,
12Jun 1976 (fr), L. Coêlho 457 (INPA); Rio Madeira, Borba, 4 Jul 1936 (fl fr), Ducke RB 30103(G, K, P, RB, S, U, US), 320 (MAD, NY); basin of Rio Solimöes, Mun. Sâo Paulo de Olivença,
Rio Tonantins, Tauriman, 9 Aug 1941 (fl), Fróes 280 (F, G, K, MO, NY, S, US, WIS), 282 (F,G, GH, K, MO, NY, P, S, US); basin of Rio Negro, Mun. Sào Gabriel, Macubeta on Rio Marié,
27 Jan 1941(fr),Fróes548(G,NY, S,WIS);Mun. SàoGabriel, LagodoDodona,basinof UenuichyRiver, 24 Mar 1942 (st), Fróes 825 (NY, US); Fonte Bôa, 5 Apr 1945 (fr), Fróes 20651 (IAN, K,
NY,US);Rio Negro,Tapuruquara, 8Apr 1947(fl fr),Fróes22121(COL,IAN,NY, RB,U, VEN);
Rio Negro, Preto, Maboaby, 8 Nov 1947 (yng fr), Fróes 22786 (IAN, NY, RB, U); Rio Negro,
Estiräo do Cordo, Ig. Jahü, Airâo, 14Jun 1948 (yng fr), Fróes 23247 (IAN, RB, SP, VEN); Rio
Capitari, Mun. de Codajâs, 3 Sep 1950 (fr), Fróes 26552 (IAN); Igarapé Dararrâ, Cachoeira do
Aracu- Ponto 13- SA-20-VA, 3Jun 1976(fr),Marinho 405(IAN);RioMarié,Ponto 2- SA-19-XA,
17 Jun 1976(fr), Marinho 572(IAN);basinofRioNegro,N ofRioNegro2kmabove Tapuruquara,
18 Oct 1971 (old fl, fr), Prance et at. 15390 (F, INPA, K, M, NY, P, S, U, US); Manaus-Pôrto
Velho Highway, km 253, 19 Mar 1974 (fl buds), Prance et al. 20734 (F, INPA, NY, Z); Estrada
Manaus-Pôrto Velho, trecho entre os Rios Castanho e Tupana, 17Jul 1972 (fl fr), M.F. da Silva
44

Agricuniv. WageningenPapers87-1 (1987)

M A P XO.'Mucoaducket(Markgraf) Zarucchi.

et al. 804 (INPA); basin of Rio Negro, without specific locality, 16Aug 1941 (fl buds), Stout 700(US); Mun. de Axinim, Rio Abacaxis, along Petrobras road from Igarapé Axinim, Lat. 4° 15' S,
Long. 58° 43' W, 7 Jul 1983 (fr), Zarucchi et al. 2973 (INPA, NY, US); without specific locality,
1Jun 1945(fr),Fróes21010(IAN,NY).

L o c a l n a m e s . COLOMBIA.Du-kô-gay(Huitotolanguage,Schuhes 3877),Juansoco (Schuttes 3877),Palo deleche{Schuhes 3877). PERU. Yahuar caspi {Abadie
19T), Yahuar huayo {Noriega sn). BRAZIL. Amaparana {Fróes20651), Jacquitaque {Fróes280, 282), Parva-papo do Terra firma {Fróes548), Sorva de Cutia{Stout 700),Sorvarana {Fróes21010).
The species upon which Mucoa duckei isbased was described by MARKGRAF
in 1932as the second speciesin the genus Neocouma, which itself was describedby PIERRE (1898). MARKGRAF named the new species in honor of its collector,
Agricuniv. WageningenPapers87-1 (1987)

45

ADOLPHO DUCKE, and included a short discussion of the relationships between

the two species then accepted in the genus. Although he did contrast the two
species using several characters, he did not indicate that his new species had
axillary inflorescences whereas N. ternstroemiacea had terminal inflorescences.
Dried specimensofboth specieshavea similarappearancewheninsterilecondition.
MONACHINO (1945)transferred Neocouma duckeito Ambelania inhisrevisionof Ambelania (sensu lato), but since the name A. duckei was already in use(MARKGRAF, 1935), Monachino provided the name A. markgrafiana. MONACHINOtreated A. markgrafiana and A. ternstroemiacea asclosely related species
in the genus. In contrast, PICHON (1948) stressed the distinction between these
two species, treating them in separate genera in his tribe 'Ambelaniées.' Ambelaniamarkgrafiana was treated as the onlymember of Ambelania sect. Neocoumopsis;however, thissectionwasnot validly published.
TheholotypeofNeocouma duckeiwaslostduringWorld War II (MARKGRAF,
pers.comm.)andtherefore thelectotypeofDucke RB 22418from the herbarium
oftheJardin Botanicoin RiodeJaneiro (RB)hasbeen chosen.
Mucoa duckei is a rather poorly collected species in light of its widespread
distribution; seeMap 10.Individuals reach up to about 15m talland are foundwithin the lower story on terrafirma. Mucoa duckei is separated easily fromthe only other species in the genus, M. pantchenkoana, by its long petioles,globosefruits, and many-flowered inflorescences.

2-2.Mucoapantchenkoana(Markgraf) Zarucchi,comb.nov.
Basionym: Ambelania pantchenkoana Markgraf, Acta Bot. Venez. 10:
247-249, fig.1.1975.
Type: Venezuela: Bolivar: Sierra Pakaraima, Cabeceras del Rio Paragua(Agua-pira), a lo largo de la frontera Venezolana-Brasilera, frontera No. 15,
3°40' N., 63° 00' W., alt 1400m, 4-5 May 1973(fl fr), Steyermark 107227 (Z,holotype;isotypes:MO 2729215,NY,VEN 95284).
Small tree up to 6 m tall with leaf-bearing branches 3-8 mm in diameter,
young stems compressed becoming terete with age. Leaf blades elliptic to obovate, 10-16 x 4.5-8 cm, glabrous, coriaceous to thick-coriaceous, with 12-20
pairs of lateral veins; bases rounded to obtuse, decurrent on petiole; apicesrounded;marginsentire,revolute;midribslightlyraisedaboveandbelow;lateralveinsslightlyimpressedabove,slightlyraisedbelow.Petioles2-4mmlong,canaliculate.Inflorescences axillary, cymose, 3-5-flowered; peduncles completely reduced, up to 5mm long; bracts ovate, small, ca 2 x 1.2 mm; pedicels 3-6 mm
long; bracteoles ovate, small, ca 1.2 x 1mm, near base of pedicel. Flowers up
to 17-20mm long;sepalsovate, 1.5-2 x 1.2-1.5mm,apically rounded. Corolla
white with tube 8-10 mm long; lobes narrowly obovate, 8-11 x 3-4 mm. Stamensinsertedca 3mm from baseofcorolla tube;anthers 3.5-4 mmlong,sessile,
46

Agricuniv. WageningenPapers87-1 (1987)

M A P 11.Mucoapantchenkoana (Markgraf) Zarucchi.

basally auriculate, apically caudate with acumen ca 0.3mm long. Ovary ovoid,ca 1.5 mm long; style 1 mm long; clavuncle 0.7 mm long; stigmoid lobes short,
ca 0.5 mm long.Fruit(immature) broadly ellipsoid, 6cm long,4.5in diameter.
Seeds numerous, plano-convex, oVate to elliptic in outline, 9-11 x 5.5-6 x
2-2.5 mm.
E t y m o l o g y . Named in honor of GEORGES PANTCHENKO, longtime explorerof the Venezuela-Brazil frontier, who assisted in securing the collection of thetype material.
D i s t r i b u t i o n . Known onlyfrom along theVenezuelan Bolivar and Brazilian Roraima border; Map 11.

Agricuniv. WageningenPapers87-1 (1987)

47

Specimens examined:
VENEZUELA. Bolivar: Distrito Hères, Cerro Marutani, a lo largo del Rio Carla, afluente de lascabeceras del Rio Paragua, 3° 50'N., 62° 15'W., downstream from camp, alt 1200m, 12Jan 1981
(fl fr), Steyermark et al. 123913(MO, VEN); SW of camp following Rio Carla, alt 1000-1050 m,
14Jan 1981(fl),Steyermark etal.124064(MO,VEN).

No localnamesarerecorded for thisspecies.
Mucoapantchenkoana isknownonlyfrom threecollections.Oneofthecollection localities, on the Venezuela-Brazil border at 1400m, is the highest known
elevation recorded for a tree classified in the Ambelanieae. Mucoa pantchenkoana isdistinguished easily from M. duckeibythecharacters listedinthekey.
In describing Ambelaniapantchenkoana, MARKGRAF (1975) stated that its affinity was with A. duckei of the region near Manaus. Although there may be
a superficial resemblance to thisspecies,M. pantchenkoana ismostclosely relatedtoM. duckei, primarily onthebasis ofitsreproductive structures.

3. Spongiosperma Zarucchi, gen.nov.(Apocynaceae; Plumerioideae; Ambelanieae).
Basionym: Molongum sect. TrichosiphonPichon, Mem.Mus.Natl.Hist. Nat.
24: 168, pi. IV - 14, 24, 35. 1948. Type species. Molongum macrophyllum(Mueller-Argoviensis) Pichon.
Shrubs or small treesup to 6m tall, lactescent, with stems terete or subterete
and with conspicuous interpetiolar line or ridge. Leaves decussate, petiolate,glabrous, thin- to thick-coriaceous, venation brochidodromous, margins entire
andrevolute;midribgenerallyimpressedaboveandprominent below,especiallyatbase;lateralveinsmany,joiningtoform anintramarginalvein;petiolescanaliculate.Inflorescencesterminal,cymose,congestedorlax,few-to many-flowered;peduncles short to long;bracts and bracteoles small, ovate to deltoid, with twobracteoles borneusually on lowerportion ofpedicels.Flowerslarge,calyx lobes
5,persistent,subequal,quincunciallyimbricate,outerlobessmaller,apicallyobtuse,innerlobeslarger,apicallyrounded,basallywithmanysmallglandswithin.
Corolla sympetalous, actinomorphic, salverform, white, thick; tube narrow,straight or slightly curved, long, with suprastaminal region pilose, becoming
dense near orifice, dilated at point of insertion of stamens; lobes 5, convoluteinbud,overlappingtoleft, becomingreflexed atanthesis,usually approximatingtube in length. Stamens 5, inserted below middle of tube; anthers lanceolate,sessile,basally sagittate, apically caudate. Ovary superior, conical, bicarpellate,placentation axile with ovules numerous in each locule; style terete, short tolong;clavuncle obconical to cylindrical, with ring of slightly undulate tissue basally,5-lobedapicallywithveryshort,densepubescenceonlobes;stigmoid lobes
short,blunt.Fruitamany-seeded, one-celledberry,withseedsacropetally imbricateand notembedded inpulp;seedsattached tofruit wallbylowfleshy funicles
on two opposite regions adjacent to remnants of torn dissepiment; pericarp
48

Agricuniv. WageningenPapers87-1 (1987)

MAP 3. Spongiosperma

thick,leathery.Seedsirregularlyplano-convex,generallyelliptictoovateinoutline,marginserose;testacoveredwithirregularprotuberances;surfacespongelike in texture andfinelypitted; hilum small, surrounded by a toroid, slightly
raised structure. Embryo almost aslong asseed,with small, thin, plane, ovate
orcircularcotyledons;radiclestraight,terete,about4-8timesaslongascotyledons.
Type species. Spongiosperma macrophyllum (Mueller-Argoviensis) Zarucchi(basedonAmbelaniamacrophyllaMueller-ArgoviensisinMartius,FloraBrasiliensis6(1):18.1860,as'A.macroyhylld').
Etymology. Greekspongios = sponge,sperma = seed;referring totheappearanceoftheseeds.
Distribution. Eastern, central, and northwestern Amazonia of Brazil
Agricuniv. WageningenPapers87-1 (1987)

49

(Amapâ,Amazonas,Maranhäo,Para),Colombia (Guainia,Vaupés),andVenezuela (Amazonas); also upper Rio Orinoco region in Venezuela (Amazonas,
Bolivar);Map 3.
Spongiosperma includes six species of small trees of riparian habitats fromwidespread areasofAmazonia. MONACHINO (1945)treated thetwo then-known
species as species of Ambelania; PICHON (1948) treated them in Molongum sect.
Trichosiphon. MONACHINO (1945)pointed out arelationship between the species
pairs Ambelania macrophylla—A. grandiflora [= Spongiosperma spp.] and A.
laxa—A.cuneata[= Molongum spp.],and alsosuggestedthatA. zschokkeiformis[= Molongum] may be related to this latter pair. PICHON (1948),in his descriptionofMolongum, treatedthegenusasconsistingofthreesections: Trichosiphon,
Coilanthera, and Oligoon;section Trichosiphon included M. macrophyllum andA. grandiflora [= Spongiosperma spp.], the latter species listed without making
the actual combination in Molongum, probably since PICHON did not actuallyseematerialofit.Itisobviousthatboth MONACHINOand PICHONwere hampered
intheir efforts by thelack of available fruiting material. On the basis of distinct
fruit types, among other characters, I have elevated Pichon's section Trichosiphon to generic status. The name Trichosiphon has been used previously at thegeneric levelby SCHOTTand ENDLICHER, for a genus now treated as a synonymof Sterculia L. (Sterculiaceae); the new generic name Spongiosperma has therefore been created.
K E Y TO T H E S P E C I E S O FSPONGIOSPERMA

1. Flowers with corolla tube 10-13 mm long, lobes 13-16 x 4-6 mm; fruits
fusiform tonarrowly ellipsoid (Region ofManaus,Amazonas, Brazil) . .
3-3. S.longilobum
Flowers with corolla tube 18-30 mm long, lobes 20-40 x 7-15 mm; fruits
narrowly ovoid, ellipsoid, obovoid, or globose
2
2. Leaves with 30-35 pairs of lateral veins;fruits globose (Northwestern Amazonia)
3-4S. macrophyllum
Leaveswithlessthan 30pairsoflateralveins;fruits notglobose
3
3. Petiole2.5-4.0mmlong;leafbasessubcordate,rarelyrounded,not decurrent
onpetiole (Upper Rio Paragua, Bolivar,Venezuela) 3-1. S.cataractarum
Petiole greater than 4 mm long; leaf bases rounded to obtuse and usually
decurrent onpetiole
4
4. Leaves large, 8-19 x 3-8 cmwith apices acute and ultimately short acuminate;inflorescences 7-to 18-flowered (Eastern Amazonia)
3-2.S.grandiflorum
Leaves small, 3-8.5(-12) x 0.7^4(-4.5) cm with apices rounded, obtuse, or
subacute;inflorescences 3-to9-flowered
5
5. Leaves narrowly elliptic, elliptic, to lanceolate, 3—8(—12) x 0.7-3(-4.5) cm;
anthers 4.5-5 mm long (vicinity of Cerro Yapacana, Amazonas, Venezuela)
3-5. S. oleifolium
50

Agricuniv. WageningenPapers87-1 (1987)

Leaves obovate to oblong-elliptic, 6.5-8.5 x 2.5-4 cm; anthers 6-6.5 mm
long(Rio Guainia alongColombia-Venezuelafrontier) . 3-6. S.riparium

3-1. SpongiospermacataractarumZarucchi, sp.nov.
Type:Venezuela: Bolivar: Isla del Rio Hacha, ca 600m subiendo elrio desdeel Salto Hacha; region de Canaima, ca 6° 15'N. - 62° 47' W., ait 200-500 m,
15 Feb 1964 (fl fr), Agostini 227 (US 2438548, holotype; isotypes: MY 25109,
NY [3sheets],VEN 58416&58417).
Plantae Spongiospermati longiloboet S. macrophyllo affïnis, sed foliis oblongo-ellipticisveloblongo-ovatis,apicesubacutis,obtusisvelrotundatis, basisubcordatis vel raro rotundatis; petiolo 2.5-4 mm longo; inflorescentiis congestis,
6-14-floris,pedunculo 10-16mmlongo,pedicello9-12mmlongo;floribus albis,
40-50 mm longis, tubo 18-21mm longo,lobis25-33 mm longis 8-10 mm latis,
fructibus anguste ovoideisusque ad 5.5cmlongis,2cmindiametro differt.
Small tree up to 4 m tall with leaf-bearing branches 2-4 mm in diameter.
Leaf bladeselliptic to oblong-ovate, 9-14.5 x 4-5.5 cm, glabrous, coriaceous
to somewhat thick-coriaceous, with 15-23 pairs of lateral veins; bases subcordate or rarely rounded; apices obtuse to subacute, ultimately rounded; margins
entire,revolute;midrib plane to impressed above, especially near base of blade,and prominent below; lateral veins plane or slightly raised both above and below.Petioles 2.5-4 mm long,canaliculate.Inflorescences terminal, cymose, congested, 6-14-flowered; peduncles 10-16mm long; bracts ovate to deltoid, 2.5-3
x 1.5 mm; pedicels 9-12 mm long; bracteoles broadly ovate, ca 1 x 1.5 mm,
usuallynear baseofpedicels.Flowers: mature bud 40-50mmlong;sepals ovate,
5-4 x 2.5-3.5 mm, apically rounded. Corolla white with tube 18-21 mm long,
lobeselliptictooblanceolate, 25-33 x 8-10 mm. Stamens inserted 8-9 mm frombase of corolla tube; anthers 7mm long, sessile, basally sagittate, apically caudate with acumen 0.8-1 mm long. Ovary conical, 2-3 mm long; style 3.5-4 mm
long; clavuncle ca 1.3 mm long; stigmoid lobes short, 0.2-0.3mm, blunt. Fruitnarrowly ovoid to 5.5 cm long and 2 cm in diameter, apically pointed. Seeds
numerous,irregularly plano-convex, 8.5-10 x 4-5 x 2.5-3 mm.
E t y m o l o g y . Latincataractarum = ofthecataracts,inreference tothelocalitieswherethisspecieshasbeen collected.
D i s t r i b u t i o n . Known only from two localities on the upper Rio Paragua(tributary oftheRio Orinoco), Bolivar, Venezuela;Map 12.
Specimens examined:
VENEZUELA. Bolivar: Dtto. Piar, alrededores delCampamento Canaima, selvade galena del Rio
Carrao, 4 May 1979 (fl), Benitez de Rojas 2555 (MY, VEN); Salto Ichün, Alto [Rio] Paragua, ait
380 m, 1 Mar 1958 (fl fr), Cardona 2852 (MY, NY, VEN) Rio Carroni, Canaima am Ufer des
RioCarrao,ait400m, 16Mar 1958(fl),Mägdefrau 539(M,NY [frag, assn}), Vareschi& Mägdefrau
Agric univ. WageningenPapers87-1 (1987)

51

MAP 12.Spongiosperma cataractarum Zarucchi.

6901 (VEN); am Rio Moroco, Nebenfluss des Rio Carao, bei Camp Ucaima, bei Canaima, alt
560-580m, 2Apr 1969 (fl), Oberwinkler et al. 15405(M);Sierra Ichûn, cercancias del Salto Maria
Espuma (Salto Ichûn) del Rio Ichûn, base de la Sierra de Ichûn, tributario del Rio Paragua, Lat.
4°46'N., Long. 63° 18'W., alt 500m, 28Dec 1961(fl), Steyermark 90307(MO,NY, VEN); Canaima,Jun 1962(fl), Tejera&Braun 9(NY, US,VEN),sn(NY),9Nov 1963(fl), Trujillo6013(MY).

Local name. VENEZUELA. Naranja brava (Agostini227).
Spongiosperma cataractarum isperhaps most closely related to S. longilobumof the Manaus region, from which it differs in the following characters: the leaf
bases are usually subcordate, not rounded; the blades do not narrow toward
the base; the leaf apices are more blunt; the petioles are generally shorter; theflowers are larger and fewer per inflorescence; and the fruit is narrowly ovoid
rather than fusiform to narrowly elliptic. Some collections of S. cataractarum
52

Agricuniv. WageningenPapers87-1 (1987)

were previously determined as Ambelania lopezii (a synonym of S. macrophyllum)duetothesuperficial resemblancetothisspeciesaswellasto S. longilobum.
Fruitingmaterial ofthesespecies,however, offers clear distinctions.

3-2.Spongiospermagrandiflorum (Huber) Zarucchi,comb.nov.
Basionym: Ambelania grandiflora Huber, Bol. Mus. Paraense Hist. Nat. 3:444. 1902;Ducke,Arch. Jard. Bot. Rio deJaneiro 3:240. 1922;Markgraf, Notizbl. Bot. Gart. Berlin-Dahlem 15: 620. 1941; Monachino, Lloydia 8(2):128-129. 1945;Pichon, Mém. Mus. Natl. Hist. Nat. 24: 168. 1948 [sub Molongum sect. Trichosiphon].
Type: Brazil: Para: Aramâ, 26 Feb 1900(fl), J. Huber MG 1836(MG, lectotype, here chosen [F photo sn; F 602773 with frag]; isotypes: BM, G [F photo
26842; F, GH, MO, NY, US]). Homotypic synonym: Molongum grandiflorum(Huber) Lemée,Fl. Guy. Franc. 3:297.1953[comb,illeg.].
Small tree up to 6 m tall with leaf-bearing branches 3-4 mm in diameter.
Leaf blades elliptic to narrowly ovate, 8-19 x 3-8 cm, glabrous, coriaceous,with 17-23pairsoflateralveins;basesrounded toobtuse;apicesacute,ultimatelyshortlyacuminate;marginsentire,revolute;midribimpressedabove,especially near base of blade, and prominent below; lateral veins plane or slightly impressed above,slightlyraisedbelow.Petioles 6-12mmlong,canaliculate. Inflorescencesterminal,cymose,congested,7-18-flowered;peduncles 10-26mm long;bracts ovate to deltoid, 3 x 2.5 mm; pedicels 9—14(—15)mm long; bracteolesdeltoid, 1.8-2.3 x 1.3 mm, on basal third of pedicel. Flowers:mature bud ca
48-60mmlong;sepalsbroadlyelliptictoovate,4.5—6 x 3-4mm,apically obtuse
to rounded. Corolla white with tube 20-30 mm long, lobes elliptic to narrowly
obovate, 30—40 x 10-15 mm. Stamens inserted 9-11 mm from base of corolla
tube; anthers 8-9 mm long, sessile, basally sagittate, 1.1-1.4mm long, apicallycaudate with acumen ca 1 mm long. Ovary conical, 4-5 mm long;style 6-9 mm
long;clavuncle 1.5-2mm long;stigmoid lobesshort, blunt, ca0.3-0.4mm long.
Fruit ellipsoid, 6-7 cm long and 3-3.5 cm in diameter, apically rounded. Seeds
numerous,irregularly plano-convex, ca 10-12 x 4-5 x 1.5-2.5mm.
E t y m o l o g y . Latin grandiflorum = large-flowered, in reference to the large
sizeofthe corolla.
D i s t r i b u t i o n . Known only from eastern Amazonia, in Territorio Amapâ,Maranhäo, and Para, Brazil;Map 13.
Specimens examined:
BRAZIL. Amapâ: Coastal Region, Lago Cujubim, 1°45' N - 50° 58'W, 10Aug 1962(fl, yngfr), Pires & Cavalcante 52450 (IAN, K, NY, US);Vai-quemquer-Pacui-Mac, 20Jul 1980(flyngfr), Rabelo 585(NY);Est.S.Joaquim-Pacui-Mcp., 22Jul 1980(flyngfr),Rabelo616(NY).Para:
Belém,Jardin BotanicodoMuseuGoeldi,cult,3Sep 1942(fl fr), Archer 7625 (IAN,NY), 12Dec
1942(fl fr), Archer 7958 (IAN); RioAcarâ, between Tomé-Açu &Acarâ, 3Jun 1969(fr),Austin
Agricuniv. WageningenPapers87-1 (1987)

53

Map 13.Spongiosperma grandiflorum (Huber) Zarucchi.

&Cavalcante4112(MO),3Jun 1969(flfr), Cavalcante &Austin 2239/4112 (MG);Mun.de IgarapéAçu,casadecampodoSr.Natan, 11 Dec 1978(flfr),Bastosetal.112(F,MG);Regiâodo Ariramba,
margem direita do Rio Jamaracaru, 5 Jun 1957 (fl), Black et al. 57-19873 (IAN); Belém, Museu
Goeldi, ârv. 428,cuit, 7 Apr 1958 (fl), Cavalcante 359 (MG); Mun. Acarâ, Fazenda Borba Gato,
near Rio Acarâ, 30 km W of Tomé-Açu--Paragominas road, ca 2° 40' S, 48° 35' W, 8 Nov 1980(fl), Daly D887 (NY); Mun. de Bragança, 7 km E of Bragança along road to Augusto Correa,
ca 1°03' S, 46° 40' W, alt 25 m, 7 Apr 1980 (fl fr), Davidse et al. 17977(F, MO, NY, US); Mun.
Belém,Lagoa Agua Prêta(Utinga),29Jun 1935(fl),Drouet 1949(F,GH, MICH, MO,US);Regiâo
do Alto Ariramba, Caramagal na campina rana, 20Dec 1906 (fr), Ducke MG 8002 (MG); Regiâo
Alto Ariramba, campina do [Rio]Jaramacarû, 4 Dec 1910 (fl fr), Ducke MG 11357(BM, G, MG,
RB, US), 8 Jul 1980 (fl yng fr), Martinelli et al. 6922 (INPA); Belém, Lago da Agua Prêta, 29
Oct 1914 (fl fr), Ducke MG 15519 (BM, MG), Feb 1926 (fl fr), Ducke RB 15820 (K, RB, S, U,US); Belém, Catû, 21 May 1943 (fl), Ducke 1254 (IAN, K, MG, MO, NY, UC, US); Trombetas,
Jaramacarû, 6 Jun 1957 (fl fr), Egler 478 (INPA, MG); Belém, terreno do Utinga, 5Jul 1966 (fl),Elias dePaula 207 (MG, NY), 15Dec 1944(fl), Fróes20775 (IAN, NY, US); Lago Cuçari, regiâo
do Planalto de Santarém, 13Apr 1955 (fl fr), Fróes 31727 (IAN, SP); Cabeceiras do Rio Uruarâ,54

Agricuniv. WageningenPapers87-1 (1987)

Mun.dePrainha, May 1955(fifr),Fróes31875(IAN,SP);RioUna,regiäodoPlanaltodeSantarém,
30 Jul 1955 (fl), Fróes 32014 (IAN, UB); Rio Jacaru, Mun. de Porto de Móz, 17 Sep 1955 (fl),Fróes32074 (IAN); Regiäo do Anapü, Rio Pracajai, Portel, 14Sep 1956 (fl fr), Fróes32737 (IAN,
MO); Regiäo do Rio Mojü, Rio Cairari, 25 Mar 1957 (fl fr), Fróes 33138 (IAN); Belém, IPEAN
grounds, 7 Dec 1974 (fl), Gentry & Pinheira 13129 (INPA, MO); Rio Capim, Approizi, 18 Jun
1897 (fl), J. Huber MG 769(G, MG); Bragança, Dec 1899(fl), J. Huber MG 1725(MG); Rodovia
Belém-Brasilia, km 97, 12 Oct 1959 (fl), M. Kuhlmann &Jimbo 346 (IAN, SP, US); Dist. Acarâ,Thomé Assü, up Rio Acarâ 1/2 km, alt 45 m, 27 Jul 1931 (fl), Mexia 5982a (MO, UC); Santa
Izabel (Belém-Bragança), 27 Aug 1908 (fr), MG 9577 (MG, RB), 28 Sep 1908 (fl), MG 9683 (BM,G, MG); Tomé Acu, margens do R. Acarâ, 2 Jan 1978 (fl fr), Nascimento 397 (MG, WAG); km
203 da Rodovia Belém-Brasilia, 24 May 1960 (fl), E. Oliveira813 (IAN); Belém, Estrada Tavares
Bastos, 7Apr 1970(fl), E. Oliveira5186 (IAN); BR-010Belém-Brasilia, Faz. Maravilha, Rio Ipixuna, 8 Feb 1973 (fl), E. Oliveira 6055 (IAN); Belém, Utinga, 1Dec 1969 (fl fr), M. Pinheiro 15(MG); Belém, Agua Prêta, 7 Nov 1945 (fl), Pires & Black 553 (GH, IAN, P, RB, US); Belém,
7 Nov 1945 (fl fr), Pires &Black 567 (IAN); Belém, beira do Utinga, 10 Apr 1947 (fl), Pires &Black 1471 (IAN, NY, P, RB); Belém, Catü, Dec 1952 (fl), Pires 4421 (IAN), 26 Oct 1953 (fr),
Pires 4549 (IAN, NY); Rio Guamâ, acima de Ourém, Jul 1953(fl), Pires &Silva 4612 (IAN, US);
Belém, Igapó do Catü, 3 Oct 1966 (fr), Pires & Silva 10260 (IAN); Belém, IPEAN, 5 Sep 1967(st), Pires & Silva 10846 (IAN); Belém, IPEAN, Igapó do Catü, 10 Apr 1968 (fl), Pires & Silva
11683(IAN),24Apr 1968(fl), Pires &Silva 11694(IAN); Belém, Bôa Vista, Campina do Guajarâ,margem esquerda da Rio Guamâ, 23 May 1968 (fl), Pires &Silva 11745 (IAN); Belém, groundsof IAN, 1Nov 1960 (fl), Pires 48894 (K, NY, P, US); vicinity of Belém, Sep-Oct 1961 (st), Pires
51875 (NY, UB, US); Mun. Benevides, 30 km ENE of Belém, 24 Jan 1980 (fl fr), Plowman et al.
8092(ECON-Prod. Coll.8550,GH, NY, US);Belém,terrenoda EMBRAPA,proximoao Mocambo, 23Apr 1979(fl), M. Ramos 02(GH, MG, NY);Igarapé-assü, 10Feb 1903(fl), R. S. Rodrigues
MG 3367 (MG), 14Feb 1903 (fr), R. S. Rodrigues MG 3396 (G, MG); Ananindeua, Maguari, 15
Apr 1975(fl fr), M. Silva 2709(MG); Sta. Izabel, Caraparu, 25Apr 1977(fr), M. Silva 3390 (MG),
25Apr 1977(fl), M. Silva 3407 (MG, MO, NY); Mun. de Acarâ, beira do Rio Acarâ, 23Oct 1979
(fl fr), M. Silva &Pinheiro 5078 (NY); Tomé-Acü, margen do Rio Acarâ, estrada para a fazenda
Borba Gato, 2 Nov 1979(fr), M. Silva &Pinheiro 5125 (NY); Belém,Estrada do Cafezal do IAN,
5 Jan 1948 (fl fr), N. T. Silva 120 (IAN, NY, RB); Gurupâ, igarapé Jacopi, 8 Feb 1979 (fl), N.
T. Silva &Rosârio 5058 (MG, NY). Maranhäo: km 55 da BR 316, a 10 km de Araguanâ, 8 Dec
1978(fr), Rosa &Vilar2887(F,NY).

Local names. BRAZIL. Açucena d'agua (Ducke RB 15820),Angelica do Igapó(Huber MG 769,Pires 48894), Angelica-igapó (Pires 51875), Molongó (Huber
MG 1836,Rabelo 585, 616), Pepino (Nascimento 397), Pepino do mato (Archer
7625),Tajarana (R. S. Rodrigues MG 3396).
Spongiosperma grandiflorumisthemostthoroughly collected species oiSpongiosperma,withthemajority ofthecollectionscomingfrom theimmediatevicinity of Belém in eastern Amazonia. This species shows a close resemblance to
S. macrophyllum of the northwest Amazon basin, a similarity which was noted
by HUBER (1902)whendescribingAmbelaniagrandiflora. Ambelania grandiflora
Huber was included by PICHON (1948) in hisMolongum sect. Trichosiphon, but
he did not make the combination in Molongum apparently since he had notseen an actual specimen of this species. Several years later, LEMÉE (1953) madean illegitimate combination in Molongum (attributing the combination toPICHON) since he did not provide a clear and complete bibliographic reference
tothe basionym.
Thelarge,white,showy,fragrant flowers indicate that speciesof SpongiosperAgricuniv. WageningenPapers87-1 (1987)

55

mamay beofhorticultural interest. Spongiospermagrandiflorum hasbeencultivated on the grounds of the Museu Goeldi in Belém(Cavalcante, pers.comm.),whereithasflowered and setfruit {Archer 7625,7958,and Cavalcante 359).
Unlike Spongiosperma macrophyllum and other speciesin the genus, S. grandiflorumisnot recordedbycollectorsasgrowinginlargepopulations,but rather
as individuals of somewhat rare occurrence. Some collections made by DUCKE,
from the western extent of the species range in the Rio Trombetas drainage,
are reported to occur in campinarana (low tree forest on sandy soil). Habitatand environmental factors affect the morphological expression of this species,
as well as that of other species in the tribe. For example, the leaves of somespecimens from the Rio Trombetas region, primarily at Ariramba, bear slightly
smaller and thicker leaves than collections made from riparian habitats. This
has also been noted in specimens of S. oleifolium of the middle Rio Orinoco
basin of Venezuela.
Various characters distinguish Spongiosperma grandiflorum from S. macrophyllum. Ithasfewerpairsoflateralveins,theleafapexismoreblunt,thepetioles
are longer, the flowers are fewer per inflorescence, and the fruit is ellipsoid instead ofglobose.

3-3. Spongiospermalongilobum(Markgraf) Zarucchi,comb.nov.
Basionym: Ambelania longiloba Markgraf in Prance, Brittonia 23(4): 441.
1971.
Type: Brazil: Amazonas: Manaus, km 74 da Estrada Manaus-Itacoatiara,
Rio Preto, 25 Jul 1966 (fl), Duarte [& L. Coêlho]9845 (Z, holotype; isotypes:
HBnv,INPA 17580[4sheets]).
Small tree up to 5m tall with leaf-bearing branches 2.5-5 mm in diameter.
Leaf blades oblong-elliptic to lanceolate, 10—14(—19) x 4.5-6(-6.5) cm, glabrous,coriaceous,with 15-22pairsoflateral veins;basesrounded or somewhat
subcordate with blade narrowed toward the base; apices acute or shortly acuminate;marginsentire,slightlyrevolute;midribimpressedabove,especiallynearbase of blade, and prominent below; lateral veins slightly raised both aboveand below. Petioles 3—5.5(—8)mm long, canaliculate. Inflorescences terminal,
cymose,congested, 12-25-flowered; peduncles 5—15(—23)mm long;bracts ovate
to deltoid, ca 1.5 x 1mm; pedicels 3-6 mm long; bracteoles small, deltoid,ca 1 x 0.8 mm, on basal third of pedicel.Flowers:mature bud 22-28 mm long;
sepals ovate, 3 x 2.5-3 mm, apically obtuse to rounded. Corolla white withtube 10-13mm long,lobesnarrowly ovate, 13-16 x 4-6 mm. Stamens inserted
4-5 mm from base ofcorolla tube;anthers 5-5.5 mm long, sessile,basally sagittate,0.6mmlong,apicallycaudatewithacumen0.4-0.5mmlong.Ovaryconical,
3-4 mm long; style 1.5-2 mm long; clavuncle ca 1.2 mm long; stigmoid lobes
short, 0.25 mm, blunt. Fruit fusiform to narrowly elliptic, up to 6-9 cm longand 1.5-2cmindiameter, apically and basallypointed. Seeds numerous, irregu56

Agricuniv. WageningenPapers87-1 (1987)

M A P 14.Spongiosperma longilobum(Markgraf) Zarucchi.

larlyplano-convex, ca7-10 x 3.5-5 x 2-2-2.5 mm.
E t y m o l o g y . Latin longilobum = long-lobed, in reference to the size of thecorolla lobes.
D i s t r i b u t i o n . Known onlyfrom thevicinity ofManaus,Amazonas, Brazil,
west of the Rio Negro and north of the Rio Amazonas, where it is found frequently alongblackwater streams;Map 14.
Specimens examined:
BRAZIL. Amazonas: Estrada Manaus-Caracarai, km 61,Réserva Biólogica de Campina, 3 May
1973(flfr),Albuquerque elal.657(GH,INPA);estrada Manaus-Caracarai, km 60,Estaçào Experimental de Silviculture Tropical, 16Aug 1973 (fl fr), Albuquerque & Coêlho INPA 99868 (INPA);
Rio Urubü, Iracema waterfall and nearby stream, 6 Aug 1979 (fl), Calderon et al. 2941 (INPA,
Agricuniv. WageningenPapers87-1 (1987)

57

MO, US); Manaus, Rio Preto, km 78 estrada Manaus-Itacoatiara, 14Jan 1966 (fl fr), L. Coêlho
INPA 16782(ECON,INPA); Rio Urubu,Cachoeira Lindoya, 26Sep 1941(fl),Ducke 798(F, IAN,
MG, MO, NY, RB, US); Manaus, beira do Rio Tarumà, 8Aug 1949(fl), Frôes24948 (IAN); Rio
Urubu,cachoeira Lindoia,26Sep 1949(fl),Frôes25410(IAN);SEPLAC, km 60on Manaus-Caracarai Road (BR 174), 1Dec 1974 (fr), Gentry 12991 (INPA, MG, MO, NY); Manaus-Itacoatiara
Highway, Rio Preto, km 80, 14 Nov 1966 (fl fr), Prance et al. 3162 (INPA, K, MG, NY, US);banks of Rio Urubu between Cachoeira Iracema and Manaus-Itacoatiara Road, 8 Jun 1968 (fl),Pranceet al.5075 [Paratype collection of Ambelania longilobaMarkgraf] (INPA, K, NY, US); Rio
Cuieras, above mouth of Rio Brancinho, 11Sep 1973(fl fr), Prance et al. 17734(INPA, MG, NY,Z); Estrada Manaus-Itacoatiara, km 90, Rio Preto, 19Jul 1961 (fl fr), W. Rodrigues &Lima 2192
(INPA); Rio Branquinho, afluente do Cuieiras, 28 Nov 1962 (fr), W. Rodrigues & Coêlho 4904
(INPA); Manaus-Caracarai Road (BR-174), Igarapé at km 46, 14Sep 1979 (fl fr), Zarucchi et al.
2541 (GH, INPA, NY); 18 Jun 1983 (fl buds, yng fr), Zarucchi et al. 2839 (INPA, NY, US); 9
Aug 1983(st),Zarucchi etal.3243(INPA, NY, US).

Local names. BRAZIL. Jasmin d'agua (Duarte [& L. Coêlho] 9845), Molongó{W. Rodrigues & Coêlho4904).
The first reference to Spongiosperma longilobum was by ADOLPHO DUCKE(1944)inhis 'Flora do rio Urubu,' wherehecited acollection {Ducke 798)from
the Cachoeira Lindoya under Ambelania macrophylla. This latter species wasthen known only from the type collection, made by RICHARD SPRUCE along
thelower Rio Uaupésin Brazil.
MARKGRAF cited the type collection of Ambelania longiloba as having beencollectedbyA.P.DUARTEin 1966.TheisotypeatINPAinManauslistsa second
collector, Sr. Luis COÊLHO,a longtimemember oftheINPA botanical staff.
Spongiosperma longilobum is distinguished from other species in the genus
by its narrowed, rounded or rarely subcordate leaf bases, its smaller flowers
and floral contents,and itslong fusiform ornarrowly ellipsoid fruits.
3-4.Spongiospermamacrophyllum(Muell.-Arg.)Zarucchi,comb.nov.
Basionym:Ambelania macrophylla Muell.-Arg. inMartius, Flora Brasiliensis
6(1): 18.1860as 'A. macroyhylld; Schumann inEngler &Prantl, Nat. Pflanzenfam.4(2): 124.1895;Markgraf,Notizbl.Bot.Gart. Berlin-Dahlem 15:620.1941;
Monachino, Lloydia 8(2): 127-128. 1945; 9(4):302.1946.
Type: Brazil: Amazonas: 'Prope Panure ad Rio Uaupés,' Sep 1852 (fl [fr]),
Spruce 2483 (BR, lectotype [here chosen]; isotypes: BM [in part], CGE, KBenth., K-Hook., P [Fphoto 38763;GH, US],W).Homotypic synonyms: Hancornia macrophylla Spruce ex Muell.-Arg. in Martius, Flora Brasiliensis 6(1):18. 1860,nom. nud. in synon. Rhigospira venulosaMiers, Apoc. S. Am. 68-69.
1878.Ambelania cucumerinaMiers,Apoc.S.Am. 13-14,pi. IB. 1878as 'Ambellania\ proparte(= K-Hook. sheet,citedas'2413'),nonlectotypus[= Ambelaniaacida].Molongum macrophyllum (Muell.-Arg.)Pichon,Mém.Mus.Natl. Nat.
24:168,pi.I V - 14,24,35.1948.
Heterotypic synonym: Ambelania lopezii Woodson in Schultes, Bot. Mus.
Leafl. 15(2): 76. 1951;Schultes, Bot. Mus. Leafl. 22 (10): 347, pi. 80. 1970; J.
58Agricuniv. WageningenPapers87-1 (1987)

FIG. 4. Spongiosperma macrophyllum (Muell.-Arg.) Zarucchi: 1: flowering branch, | x ;2. opened
flower, 1 x ;3.sectioned fruit, | x ; 4 . seeds, 3 x .(1. Schuhes &Cabrera 19516;2.Zarucchi 2435,
ale.mat.; 3-4. Zarucchi 2434,ale.mat.).
Agricuniv. WageningenPapers87-1 (1987)

59

Ethnopharm. 1:166. 1979,synon. nov.Type:Colombia:Vaupés [as'Caqueta']:
Rio Taraira [border with Amazonas, Brazil], at second rapids, 8-11 Jul 1948(fl), Schuhes &Lopez 10204 (MO 1579973,holotype; isotypes: IAN 60209, P,
US 1997073[NYphoto ns 5576;NY]).
Small tree up to 6 m tall with leaf-bearing branches 3-5 mm in diameter.
Leaf bladeselliptic to narrowly ovate or obovate, 12—18(—22) x 4-7(—8) cm,
glabrous, thin-coriaceous, with ca 30-35 pairs of lateral veins; bases rounded
to rarely subcordate; apices acute, distally acuminate with acumen 10-15 mm
long;margins entire,revolute;midribplaneto slightlyimpressed above, becoming deeply impressed at base of blade, prominent below; lateral veins plane
above, slightly raised below. Petioles 5-8 mm long, canaliculate. Inflorescences
terminal, cymose, very congested, 10-25(-35)-flowered; peduncles(9-)12-20(-32) mm long; bracts deltoid, 2.5-3.5 x 2-2.5 mm; pedicels 3-6(-7)
mm long; bracteoles ovate to deltoid, 1.8-2.5 x 1.5-2 mm, usually on basal
third of pedicel. Flowers:mature bud 55-65 mm long; sepals ovate to oblong
3-4 x 3-4 mm, apically rounded to obtuse. Corolla whitewith tube 22-25 mm
long, lobes narrowly obovate, 35-40 x 12-14 mm. Stamens inserted 8-10 mmfrom baseofcorollatube;anthers7-8mmlong,sessile,basallysagittate,apicallycaudate with acumen ca 1mm long. Ovary conical, 3-4 mm long; style 5-8 mm
long; clavuncle 1.6-1.8 mm long; stigmoid lobes short, ca 0.3 mm long, blunt.
Fruitgloboseorslightlycompressed,apicallyveryslightlypointed.Seeds numerous, upto 80,irregularly plano-convex, 12-15 x 5-7.5 x 2-3.5 mm.
E t y m o l o g y . Greekmacrophyllum = large-leaved.
D i s t r i b u t i o n . Known from seasonally inundated blackwater streams innorthwestern Amazonia ofVenezuela, Colombia, and Brazil;Map 15.
Specimens examined:
COLOMBIA. Amazonas-Vaupés: Rio Apaporis, Raudal Yayacopi (La Playa) and vicinity, alt ca
800 ft, 18Aug 1952 (fl), Schuhes & Cabrera 16968 (BM, GH, NY, US). Vaupés: Rio Piraparanâ,environsofCatholicmissionofSanMiguel,23Oct 1976(flfr),Davis135(COL,GH);orilla derechadel Rio Vaupés, Raudal Macucü, 14 Nov 1952 (fr), Romero-Castaneda 3493 (COL); Piracuara,
27 Nov 1952 (fl), Romero-Castaneda 3752 (COL, NY); entre Wacaricuara y el varador al Rio Yi,
9-12 Dec 1952 (fr), Romero-Castaneda 3917 (COL); Rio Kananari, Cachivera Palito, alt ca 250
m,25Jul 1951(fl),Schuttes & Cabrera13175(BM,C,GH, LE,US);RioPiraparanâ, middlecourse,
1Sep 1952(fl),Schuttes & Cabrera17135 (BM,COL,GH,MO,US); RioPiraparanâ,Cafio Oo-moofla, 3 Sep 1952 (fl), Schuhes &Cabrera 17154(COL, GH, US); Rio Piraparanâ, Cafio Teemeefia,
6 Sep 1952 (fl), Schuhes & Cabrera 17245 (BM, COL, GH, MO, US); Rio Vaupés between Mitüand Javareté, Tipiaca, Igarapé Murutinga, 14-24 May 1953 (fl), Schuttes & Cabrera 19290 (US);RioPaca (tributary ofRioPapuri),Wacaricuara andvicinity,altca650ft, 1-3 Jun 1953 (fl), Schuttes& Cabrera 19516 (GH, MO, US); Mitu and vicinity, along stream near base of Cerro de Mitü,
26Jun 1976 (fl fr), Zarucchi &Balick 1763(COL, GH, MO, US), 29Aug 1976(fl), Zarucchi 1903
(COL, ECON, GH, INPA, VEN), 6Jul 1979(fl fr), Zarucchi 2434 (COL, GH, MO, WAG), 2435
(COL,GH, MO,USF),21Jul 1979(fl fr),Zarucchi 2504(COL, GH, MO, USF).
VENEZUELA. Amazonas: Rio Orinoco, Cafio Cupavén (right bank of Orinoco opposite mouthof Rio Atabapo), alt 125-150 m, 4 Aug 1959 (fl), Wurdack &Adderley 43749 (F, G, GH, NY,
RB, S, US, VEN).
60

Agricuniv. WageningenPapers87-1 (1987)

MAP 15.Spongiospermamacrophyllum (Mueller-Argoviensis) Zarucchi.
BRAZIL. Amazonas: along Rio Curicuriari and Igarapé Cariua to Cachoeira Piraiauara, 0° 20'
S, 66° 55' W, 12Jul 1979 (fl fr), Alencar et al.632 (NY); Rio Papuri near Mello Franco, 31 Aug
1943 (fl), Allen 3090 (COL). 'Minas: Gandarela dans le campo, 24 Feb 1884 (fl), Glaziou 1408V(P)(probably apirated specimen of Spruce 2483).

Local names. COLOMBIA. Hia-WAHO-ka-ki (Kubeo language, Zarucchi
2504), Palo bälsamo {Zarucchi 2434, 2435), Wy-gaw-fiö-mee-kö (Makuna language,Schultes & Cabrera 16968).
This species was first collected by RICHARD SPRUCE on the upper Rio Negroin September 1852 where he considered it to represent an undescribed speciesof Hancornia (H.macrophylld). Subsequently during hisjourney, Spruce madeanother collection which he questionably equated to the earlier gathering.
MUELLER(1860a)placed Spruce'sfieldidentifications under theirrespectivespeAgricuniv. WageningenPapers87-1 (1987)

61

cies in Ambelania, as 'A. macroyhylld' and A. quadrangularis. Because of theapparentconfusion withtheuseofthespecificepithetmacrophylla, MIERS(1878)
proposed the epithet venulosa for the species that MUELLER had described as'A. macroyhylla,' and placed itwithA. quadrangularisinhisgenus Rhigospira.
Several oftheduplicate specimens of Spruce's typecollection of Spongiospermamacrophyllum contain extra or conflicting information. The Bentham sheet
at Kew has a small drawing of the fruit attached to the sheet, but no preserved
fruit has been located. The duplicate specimen at the British Museum (Natural
History) [BM] contains not only dried fragments of S. macrophyllum but also
six packets of analyzed floral and fruit fragments and the pencilled annotation'Ambellania cucumerina, Spruce 2413' attached at the upper left hand corner.
Thehandwriting onthepacketisthat ofJOHN MIERS,and thepacketwas probablyattached totheBM sheetafter hisdeath.Thefragments inthepacket correspond to 'Ambellania cucumerina Miers'- a mixture of two elements. The floral
fragments weremost likely removed from theHooker sheet at Kew (incorrectlycited as '2413'); this duplicate has only detached inflorescences accompanying
the vegetative shoots. The fruit fragments are from a gathering made in thevicinity of Barra [Manaus] by SPRUCE and sent by him to Kew where they currently form part of the Museums collection. In the present work, these spiritpreserved fruits are designated the lectotype for Ambelania cucumerina Miersand placed in synonymy under A. acida.
Ambelania lopeziiWoodson (in SCHULTES [1951])ishere placed in synonymy
under Spongiosperma macrophyllum. The protologue for A. lopezii includes a
discussionofitsrelationshipwithA.grandiflora[= Spongiosperma]from eastern
Amazonia. This latter species was described by HUBER (1902), who contrasted
it with A. macrophylla; the connection between A. lopezii and A. macrophylla
was not made by WOODSON. Several subsequent articles by SCHULTES (1970,1979) discuss the use of S. macrophyllum (under A. lopezii) as both an arrow
poison and afishpoison.

3-5. Spongiospermaoleifolium (Monachino) Zarucchi, comb.nov.
Basionym: Ambelania oleifolia Monachino, Mem. New York Bot. Gard.
10(1): 117-119, fig. 14a-das 'A. oleaefoM.
Type: Venezuela: Amazonas: alto Rio Orinoco, at edge of Savanna No. 3,
NW base of Cerro Yapacanâ, alt 150m, 17Mar 1953(fl), Maguire & Wurdack
34512 (NY, holotype; isotypes: F 1490247 [F photo 51090; F], G, GH, IAN
96061,K, RB 101908,S,UC 275969,US2253278,VEN42160,W).
Heterotypic synonym: Ambelania lanceolata Markgraf, Acta Bot. Venez. 6:
74.1971[1972],synon.nov.Type:Venezuela:Amazonas:CerroYapacanâ,entre
el Campamento Base y la sabana grande, Lat. 3°45' N., Long. 66° 45'W., alt
125m,7May 1970(flfr),Steyermark &Bunting 103265(Znv,holotype;isotypes:
M, US 2622451,VEN 85074&85075).

62

Agricuniv. WageningenPapers87-1 (1987)

Shruborsmalltreeupto5mtallwithleaf-bearing branches 2-4mmindiameter.Leafbladesnarrowly elliptic,elliptic,tolanceolate, 3—8(—12)x 0.7-3(-4.5)cm,glabrous, stiff-coriaceous, with 20-28 pairs of lateral veins;bases rounded
toobtuse and decurrent onpetiole;apicesrounded tosubacute;margins entire,
revolute;midribplaneabove,prominulous toprominent nearbaseofleafbelow;
lateralveinsbarelyvisibleoneithersurface.Petioles4—8(—11)mmlong,canaliculate.Inflorescencesterminal, cymose,lax,3-6(-7)-flowered;peduncles 8-15mm
long;bracts ovate todeltoid, ca 1 x 0.8mm;pedicels 5—9(—15)mmlong;bracteoles small, ovate to deltoid, ca 0.6 x 0.5 mm, usually at base of pedicel.
Flowers: mature bud ca40-50(-60)mmlong;sepals ovate, 2-3 x 1.5-2.5mm,
apically rounded to obtuse. Corolla white with tube ca 20-28 mm long, lobes
elliptic to oblanceolate, ca 20-30(-35) x 8-11mm. Stamensinserted 5-6 mmfrom baseofcorolla tube,anthers4.5-5mmlong, sessile,basally sagittate, apicallycaudatewithacumen ca0.3-0.6mmlong.Ovaryconical, 1.7-2.5mmlong;style 2-2.5 mm long; clavuncle ca 0.8 mm long; stigmoid lobes short, ca 0.2
mm long, blunt. Fruitobovoid, to 5cm long and 2.5cmin diameter, apicallyrounded. Seedsnumerous,generallyplano-convex, 5-8 x 3.5-5 x 1.5-2 mm.
Etymology. The epithet refers to the similarity of the leaves to those in
Olea,thecommon olivetree.
Distribution. Known onlyfrom thevicinity ofCerro Yapacana, Amazonas,Venezuela; Map 16.
Specimens examined:
VENEZUELA.Amazonas:Depto.Atabapo, area between thewestern baseofCerro Yapacana andthe headwaters of Cano Cotua, Savanna III, 3° 38' N, 66° 52' W, alt ca 100 m, 6 May 1979 (flfr), Davidse et al. 17223(MO), 6 May 1979 (fl), Davidse et al. 17258(MO), 3 Jun 1978 (fl fr), O.
Huber 2034 (MO, VEN), 10 Dec 1978 (fl fr), O. Huber & Tillen 2995 (VEN); I sabana entre el
Cafio Cotùa y el Cerro Yapacana, 3° 40' N, 66° 50' W, ait 100 m, 25 May 1978 (fl), O. Huber
1796 (VEN); pequena sabana ubicada entre la punta 0 del Cerro Yapacana y el Cano Yagua al
S., 3° 36' N, 67°47' W, ait ca 100m, 24 Aug 1978 (v yng fr), O.Huber 2538 (VEN), 24 Aug 1978(fl), O.Huber 2541 (VEN); a unos 10 km al N del sector NE del Cerro Yapacana, 3° 44' N, 66°
48'W,aitca 100m,28Feb 1980(fr), O.Huber 4807(VEN);RioOrinoco,CerroYapacana, savanna
at base, alt 175m, 31 Dec 1950(fl fr), Maguire et al. 30474 (NT); Cerro Yapacana, Savanna No.
3, 1 Jan 1951 (fl fr), Maguire et al. 30553 (NY); NW base of Cerro Yapacana, Savanna No. 2,
ait 150 m, 17 Mar 1953 (fl fr), Maguire & Wurdack 34504 (NY, US, VEN); NW base of Cerro
Yapacana, edge of Savanna No. 3, alt 125m, 16Sep 1957(fl), Maguire et al.41494(M, NY, US),
16 Sep 1957 (fr), Maguire et al. 41507 (LE, M, NY, US, VEN); savanna along Rio Cotua W ofcampsite on laguna section, near base of Cerro Yapacana, ait 75 m, 8 May 1970 (fl), Steyermark&Bunting 103268 (MO); Cano Yapacana, 25 Jan 1958 (fl), Vareschi &Mägdefrau 6602a (VEN);
along Cano Yapacana, alt 125m, 17Jun 1959 (fl fr), Wurdack &Adderley 43021 (MO, NY, P,
US, VEN).

Nolocalnamesarereported for thisspecies.
In the original description of Ambelania oleifolia var. riparia, MONACHINO(1961)citedtwocollections.Iinterpret theseasrepresenting twodistinct species,
Spongiosperma riparium and S. oleifolium. With the additional material from
Agricuniv. WageningenPapers87-1 (1987)

63

MAP 16.Spongiosperma oleifolium (Monachino) Zarucchi.

the Rio Guainia on the Colombia-Venezuela border {Schuhes & Lopez 10132),
thedistinction betweenthesetwocloselyrelated speciesismadeapparent. Much
confusion haspersisted inregard tothesespecies,astheSchuhes &Lopez collectionwasidentified originallyasAmbelania lopezii[= S. macrophyllum].
The paratype collection for Ambelania oleifolia var. riparia, Wurdack & Adderley 43021, wascollected along the Carlo Yapacana. This stream flows fromthesavannaareawherethecollectionsofA. oleifoliacitedbyMONACHINO (1958)
weregathered. It appears that thecollections citedin theprotologue by MONACHINO(1958)for A. oleifoliamayrepresentdepauperateindividualswhich reflect
the rigors of the local environmental conditions by their smaller leaves and diminished stature.
Another recentlydescribed speciesby MARKGRAF(1971),Ambelania lanceolata, is also here considered as representing Spongiosperma oleifolium and so isplacedin synonymy.
64

Agricuniv. WageningenPapers87-1 (1987)

Spongiosperma oleifolium isdistinguished from S. riparium and other speciesof Spongiosperma byitsfewer flowers, longer pedicels,and small obovoid fruit.
Further field studies are needed in order to clarify the relationships between
the morphology exhibited by S. oleifolium and related species and the various
environmental, especially edaphic,factors which affect their appearance.

3-6.Spongiospermariparium(Monachino) Zarucchi, comb.& stat.nov.
Basionym: Ambelania oleifolia Monachino var. riparia Monachino, Mem.
New York Bot. Gard. 10(4): 59. 1961 [as 'A. oleaefolia'] pro parte, as to typecollection only.
Type:Venezuela: Amazonas:alongCano Pimichin between Rio Guainia andPimichin, alt 130-140m, 2Jul 1959(flyngfr), Wurdack & Adderley 43279 (NY,holotype;isotypes: F 1589355,GH, IAN 114667,K, LE, MICH, MO 1782971,
NY,P,RB 115008,UC221540,US 2373308,VEN 52814).
Small tree up to 6 m tall with leaf-bearing branches 2-3 mm in diameter.
Leaf blades obovate to oblong-elliptic, 6.5-8.5 x 2.5^4 cm, glabrous, thickcoriaceous, with ca 20-25 pairs of lateral veins;bases rounded to subacute anddecurrent onpetioles;apicesobtuseorrounded;marginsentire,revolute;midrib
impressed above, especially near base of blade, prominent to quite prominent
below; lateral veinsplane and difficult to distinguish above and below. Petioles
6-9 mm, canaliculate, slightly winged. Inflorescences terminal, cymose, 6-9flowered; peduncles 6—9(—11)mm long; bracts ovate to deltoid, ca 1.5 x 1.2mm; pedicels 3-6(-7)mm long; bracteoles small, ovate to deltoid, ca 0.7 x 0.5mm, near base of pedicel. Flowers:mature bud 45-55 mm long; sepals ovate,
3-4 x 2.5-3 mm, apically obtuse to rounded. Corolla white with tube 18-20mm long, lobes elliptic to oblanceolate, 25-30 x 7-10 mm. Stamens inserted
6mmfrom baseofcorollatube;anthers6-6.5 mmlong,sessile,basally sagittate,apically caudate with acumen 0.6-0.8mm long. Ovary conical, 2-3.5 mm long;style 2-3 mm long; clavuncle 1.6 mm long; stigmoid lobes short, ca 0.2 mm
long,blunt.Maturefruits and seeds unknown.
E t y m o l o g y . The epithet riparium refers to the habitat in which the species
wascollected,alongthebanks ofrivers or streams.
D i s t r i b u t i o n . Known only from four collections made in the Rio Guainia
drainage alongtheColombia-Venezuelaborder; Map 17.
S p e c i m e n s examined:
COLOMBIA. Guainia: Rio Guainia below San Antonio, Caiio del Loro, Jun 1948 (fl), Schuhes&Lopez 10132(COL,GH, MO).
VENEZUELA. Amazonas:Depto.Casiquiare, alrededoresdeYavita (RioTerni)ycerca delacarreterra Yavita-Pimichin hasta el km 5 hacia Pimichin, en orilla inundata del Rio Temi, alt 125-140m, 6-19Jul 1969(fl),Bunting et al,4038(MY);Cano Pimichin entre Pimichin yla desembocadura,
altca 128m,6-19Jun 1969(fl),Bunting etal.4066(MY).
Agricuniv. WageningenPapers87-1 (1987)

65

MAP 17.Spongiosperma riparium(Monachino)Zarucchi.

Local name. Molongó{Schuhes&Lopez10132).
SeetheabovediscussionunderSpongiospermaoleifoliuminregardtoitstaxonomieconfusion withS. riparium.
Spongiosperma riparium is distinguished from most of the other species inthegenusbyitssmaller,thick-coriaceousleaves,characterswhichitshareswith
S.oleifolium.ItcanbeseparatedeasilyfromS.oleifolium,however,byitsshorter
pedicels and greater number offlowers. Spongiosperma riparium isnot known
withmature fruit.
4.MolongumPichon,Mem.Mus.Natl.Hist.Nat. 24:167-169,pi.IV- 14-17,
24-27,35-38.1948proparte.
66

Agricuniv. WageningenPapers87-1 (1987)

Ambelania sect. Terminales Markgraf, Notizbl. Bot. Gart. Berlin-Dahlem 15:
620.1941pro parte,nom. nud.
Ambelania sect. Laxae Monachino, Lloydia 8(2): 114. 1945 pro parte, nom.
nud.
Molongum sect. Coilanthera Pichon, Mém. Mus. Natl. Hist. Nat. 24: 168,/?/.
IV-15,25,36.1948.
Molongum sect. Oligoon Pichon, Mém. Mus. Natl. Hist. Nat. 24: 168-169,
pi.
IV-16-17,26-27,37-38.1948.
Spreading shrub or small tree up to 12 m tall, lactescent, with stems tereteor subterete, nodes somewhat swollen; interpetiolar line or ridge conspicuous.
Leaves decussate, petiolate, glabrous, subcoriaceous to thick-coriaceous, venation brochidodromous, margins entire and revolute, blade decurrent on partor most of canaliculate petiole; midrib plane to impressed above, prominent
below. Inflorescences terminal, cymose, congested to lax and spreading, fewtomany-flowered; peduncleselongate;bractsdeltoidtoovate.Flowerssubsessile
to pedicellate, few to many; bracteoles deltoid to ovate, usually near base of
pedicel; calyx lobes 5, persistent, quincuncially imbricate, basally with several
smallglandswithin;sepalsfree nearlyto base,ovate,rounded toobtuse, eciliate
atapex.Corollasympetalous,actinomorphic,salverform, whitetoyellow,thick;tubenarrow, straight, short,glabrouswithin, slightlydilatated at point of insertionofstamens;lobes5,convoluteinbud,overlappingtoleft, becoming reflexedat anthesis, narrowly ovate to obovate, slightly longer than tube. Stamens 5,
insertednearmiddleoftube;antherslanceolate,small,sessile,basallyauriculate,apically caudate. Ovary superior, ovoid or cylindrical, glabrous, apicallyrounded, obtuse,ortruncate,bicarpellate,placentation axilewithovules numerousineach locule;styleterete;clavunclecylindrical or ovoid, with basal flange,
five-lobed apically, star-shaped incross-section, glabrous;stigmoid lobesterete,thin, one-third to subequal to length of clavuncle. Fruit few-seeded, two-celled
berry, narrowly ellipsoid to lanceovoid, with seeds embedded in a fluffy, white
pulp;pericarp thin, fleshy, smooth, glandular. Seeds piano-to concavo-convex,
ovate to elliptic in outline, testa muriculate and very finely reticulate; hilum
small, usually broadly ovate or circular. Embryo almost as long as seed, withsmall, thin, plane, ovate cotyledons; radicle straight, terete, about 4-7 times aslongascotyledons.
Type species. Molongum laxum (Bentham) Pichon, Mém. Mus. Natl. Hist.
Nat. 24:168,pi. IV-15,25,36.1948 (basedon Tabernaemontanalaxa Bentham,
J. Bot. [Hooker] 3:244.1841).
E t y m o l o g y . From thelocal Brazilian name, 'Molongó.'
D i s t r i b u t i o n . Central and northwestern Amazonia of Brazil (Amazonas),
Colombia(Guainia,Vaupés),andVenezuela(Amazonas)andtheadjacent areas
oftheupper Rio Orinoco inVenezuela; Map 4.

Agricuniv. WageningenPapers87-1 (1987)

67

MAP 4. Molongum

Molongum was described by PICHON (1948)to encompass six speciesin three
sections: Trichosiphon, Coilanthera, and Oligoon.In thispresent work, sect. Trichosiphonisremovedfrom Molongum andtreatedasthedistinctyetclosely relatedgenusSpongiosperma. PICHONwashampered inhisstudybythelackofavailable fruiting material, which Ihavefound to provide some of themore obvious
differences useful inseparating thetwogenera. Molongum isdistinguished from
Spongiosperma by such characters as the relatively short corolla tube which isglabrous within, the thin pericarp which encloses few, rather than many seeds,
and the seeds which are attached to a central septum and embedded in a fluffy
white pulp, instead of projecting into a single open cell attached by low fleshyfunicles.
The three species of Molongum are found growing along rivers or streams
in Amazonia and the upper Rio Orinoco tributaries, with M. lucidum known
68

Agricuniv. WageningenPapers87-1 (1987)

alsofrom lowlandsavannaregionsofeasternColombia,Venezuela,andBrazil.
Allthreespecieshaveverylightweightwoodwhichisreportedlyusedbynative
peoplesasasubstitutefor cork.
KEY TO THE SPECIES OF MOLONGUM
1. Corollatube3-A5mmlong,lobesobovate,5 x 3 mm;flowerpedicels2-3.5
mmlong;leavessubcoriaceoustothin-coriaceous,marginsentiretocrenulate,with 9-16 pairs of lateral veins;petioles 9—15(—28) mmlong (Lower
RioMadeiraregion,Amazonas,Brazil)
4-3.M.zschokkeiforme
Corolla tube 5-9 mmlong,lobesoblong-elliptic to narrowly ovate,7-12 x
3.5-4mm;flowerpedicels4-14mmlong;leavescoriaceoustothick-coriaceous,margins entire,with 15-25pairs of lateralveins;petioles (3—)5—12
mmlong2
2. Leaf bladeselliptic,rarely narrowly ovate or obovate, base rounded toobtuse,apexobtusetoacute,sometimesshortlymucronate;corollatube7-9
mm long, lobes oblong-elliptic to narrowly ovate, 10-12 x 3.5-4 mm(NorthwesternAmazonia)
4-1.M.laxum
Leafbladesobovatetooblanceolate,basebroadlyto sharplycuneate,apex
rounded to obtuse;corolla tube 5-7 mm long; lobes oblong-elliptic, 7-8
x 4mm(NorthwesternAmazonia)
4-2.M.lucidum
4-1. Molongumlaxum(Bentham)Pichon,Mem.Mus.Natl.Hist.Nat. 24: 168,
pi.IV-15,25, 36.1948.
Basionym:TabernaemontanalaxaBentham,J.Bot.(Hooker)3:244.1841.
Type:Brazil:Amazonas:OntheRioNegro,nodate[probablycollectednear
Barcelos, May 1839] (fl), Schomburgk 919 (K, lectotype [here designated];
isotypes, BM, CGE, E-GL, F 870259, FI-W, G, G-DC, K, L, NY, OXF, P,
US702978,W).Homotypic synonyms:Hancomia laxa(Bentham)A.DeCandolle, Prodr. 8:326. 1844.Ambelania laxa(Bentham) Muell.-Arg. in Martius,
FloraBrasiliensis6(1):17,pi.4.1860;SchumanninEngler&Prantl,Nat.Pflanzenfam. 4(2): 124.1895;Markgraf,Notizbl.Bot.Gart. Berlin-Dahlem 15: 620.
1941;Monachino,Lloydia8(2):126-127.1945;9(4):302. 1946.
Small tree up to 8m tall with leaf-bearing branches 2-3 mm in diameter.
Leaf blades elliptic, rarely narrowly ovate or obovate, (4-)5-10 x (1.5-)2-4
cm, glabrous, coriaceous, with ca 15-25 pairs of lateral veins; bases rounded
toobtuse,slightlydecurrentonupperportionofpetiole;apicesobtusetoacute,sometimes shortly mucronate; margins entire, revolute; midrib plane to impressedabove,especiallynearbaseofblade,prominentbelow;lateralveinsplane
onbothsurfaces,barelyperceptible.Petioles(3-)5-7(-10)mm,canaliculate.Inflorescences terminal, cymose, usually lax and spreading, (5-)8-24(-35)-flowAgricuniv. WageningenPapers87-1 (1987)

69

M A P 18.Molongwn laxum (Bentham) Pichon.

ered; peduncles (18—)25—60mmlong; bracts ovate to deltoid, 2-3.5 x 1.5-2mm;pedicels4-6(-8)mmlong;bracteolesovate,cal.5x 1mm.Flowers:maturebud 16-20mmlong;sepals ovate, 1.8-2.5 x 1.7-2.5mm,apically rounded or
obtuse. Corolla white or cream with tube 7-9mmlong, completely glabrouswithin; lobes oblong-elliptic to narrowly ovate, 10-12 x 3.5-4mm.Stamens
inserted 3.5-4mmfrom base ofcorolla tube; anthers 3-3.5mmlong,sessile,basallyauriculate,apicallycaudatewithacumen0.3-0.4mmlong.Ovaryovoid,
apicallyobtuse,1.5-2mmlong;style1.2-2mmlong;clavuncle0.7-0.9mmlong;stigmoidlobesthin,terete,0.3-0.5mmlong.Fruitnarrowlyellipsoidtolanceovoid, 2.5-5(-8) cmlong, apically acute orobtuse, basally obtuse or rounded.
Seedsfew,2-5(-8),planp-toconcavo-convex,ca5-6 x 3-4 x 2mm.
Etymology. Theepithet laxumrefers totheopen orloosearrangementof
theflowersintheinflorescence.
Distribution. Known from themiddletoupper RioNegro in Braziland
70

Agricuniv. WageningenPapers87-1 (1987)

Colombia, and theadjacent area of the upper RioOrinocoinVenezuela; Map
18.
Specimens examined:
COLOMBIA. Guainia: Cafio Bocachico, en el Rio Negro, alt 75 m, 28 Oct 1977 (fl), Espina et al.
274 (COL); 0.5 km N of Boca de Casiquiare (where Rios Negro, Guainia & Casiquiare join), 1-2
km W of Rio Guainia along cafio, 1°57' N, 67°08' W, alt 120m, 5Feb 1980(fl), Liesner &Clark
9119 (MO, VEN); Rio Atabapo, just above Cacagual, alt 130 m, 19 Nov 1953 (fl), Maguire etal.36267(COL,MICH, NY,S).
VENEZUELA. Amazonas: Depto. Atabapo, Cano Yagua, medio (afluente derecho del Rio Orinoco), 3° 36' N., 66° 35' W., ait ca 100 m, 27 May 1981 (fl fr), Cerda sn (VEN); Depto. Atabapo,
southeastern bank of the middle part of Cafio Yagua at Cucurital de Yagua, 3° 36' N., 66° 34'
W., alt ca 120m, 8 May 1979 (fl fr), Davidse et al. 17335 (MO); Rio Casiquiare, without precise
locality, Jan-Feb 1969 (fl fr), Farinas et al. 644 (NY, VEN); Rio Casiquiare, dentro de la laguna
Baciba, Jan-Feb 1969 (fl), Farinas et al. 686 (NY, VEN); Depto. Rio Negro, pequena sabana en
lariberaizquierda (Sur)delbajo RioSiapa,pocodistantedesudesembocadura enel RioCasiquiare,
2° 05'N., 66° 25'W., alt ca 125m, 7 Feb 1981 (fl), O.Huber &Medina 5819 (VEN); Rio Paciba,
Lago Paciba, alt 130m, 3Apr 1953(fl fr), Maguire & Wurdack 34798 (NY, US);along Rio Paciba
between Lago Paciba and Rio Casiquiare, alt 130m, 4Apr 1953(fl fr), Maguire & Wurdack 34866(GH,IAN,K, MO,NY, P,RB,UC,W);Rio Guainia, alongriverbetween Rio Negro and Comunidad, alt 100m, 12 Apr 1953(fl),Maguire & Wurdack 35547(F,NY); Rio Yatua near Laja Catipan,
alt 100-140m, 6Feb 1954(fl fr), Maguire & Wurdack 37528(G, LE,NY), 11 Jan 1958(fl), Maguire
et al. 42627 (NY, RB, VEN); Rio Casiquiare, Cano de Agua Negra (Milla 38), 17 Apr 1968 (fl),Medina 250 (VEN); Rio Orinoco, Cafio Cariche, 27 Apr 1968 (fl fr), Medina 419 (VEN); Cano
Cupavén, cerca de San Fernando de Atabapo, ait 100 m, 2 May 1968 (fl), Medina 492 (VEN);
14 Apr 1978 (fr), Morillo et al. 7580 (MY, VEN); Depts. Atabapo & Casiquiare, a lo largo delRio Atacavi, del Cano Caname y del Rio Atabapo entre los 3° 05' y 3° 50' N., y entre 66° 50'
y67°30'W.,altca 100 m, 10-24Nov
l980(f[),Pinate&Mondolfll044(yEN);adauminaCasiqman,
Vasiva et Pacimoni, Jan 1854 (fl fr), Spruce 2445bis (BM, BP, BR, C, CGE, F, K, NY, P, S,W);
Rio Pacimoni, arriba de la desembocadura del Rio Pacimoni con el Rio Casiquiare, 1°50' N, 66°
30' W, ait 100 m, 8 Apr 1970 (fl), Steyermark & Bunting 102453 (COL, M, MO, S, US, VEN,W); Bajo Rio Atabapo, Isla Sapo, ait 124m, 19Jan 1942(fl), Ll. Williams 13856(F, G, US, VEN[in part]); Rio Guainia, Santa Rita, alt 100m, 17Mar 1942 (fl), LI. Williams 14799 (F, G, VEN);
Rio Guainia, Cano San Miguel, ait 127 m, 25 Mar 1942 (fl fr), Ll. Williams 14874(F, MO, MY,
US, VEN).
BRAZIL. Amazonas: along Rio Curicuriari and Igarapé Cariua to Cachoeira Piraiauara, 0° 20'
S., 66° 55'W., 12Jul 1979 (fl), Alencar et al.594 (NY); Mun. Sâo Gabriel da Cachoeira, margem
direita do Rio Negro, Rio Curicuriari, 3Dec 1978(fl), Damiäo [= Mota] 2967 (COL, INPA); Rio
Negro, Rio Curicuriary, 29 Nov 1929 (fl fr), Ducke RB 22420 (G, K, P, RB, S, U, US), 4 Oct
1935 (fl), Ducke 44 (A, F, K, MO, NY, S, US), 24 Oct 1932 (fl), Ducke 163 (A, F, MAD); Rio
Curicuriary, acima do Cach. Cajû, 22 Feb 1936,Ducke RB 30123 (INPA, RB);Acarâ, Ig. Cubaté(trib. of Rio Içana), 4 Jan 1942 (fl fr), Fróes 491h (G, GH, NY, UC, US); Rio Negro, Enuixy,
Lago do Dondona, 18 May 1947 (fl fr), Fróes 22367 (IAN, NY, RB, U, US, VEN); Rio Negro,
Padauiry, Tapera, 1 Nov 1947(fl fr), Fróes 22692 (IAN), 1 Nov 1947 (fl), Fróes22695 (IAN, NY,
RB, U, US, VEN); Rio Negro, Upper Jurubaxi River, 4 Jul 1948 (fl fr), Fróes 23304 (IAN, RB,
VEN); RioNegro,Barcelos, 15 Apr 1952(flfr),Fróes28265(IAN,NY);Itaubal, RioAracâ, sub-afl.
do Rio Negro, 4 Nov 1952 (fl), Fróes &Addison 29258 (IAN, NY, UB, Z); baixo Rio Negro, Rio
Jufari, Prosperando, 26 Sep 1975(fl), Kubitzki 75-84 (HBG, INPA); Rio Negro, Lago do Atàana,below Barcelos, 10Oct 1978 (fl fr), Madison et al. 6131 (F, INPA, NY); Rio Urubaxi, Ponto 16,
SA-20-VC, 5 Jun 1976 (fl fr), Marinho 424 (IAN); Sâo Gabriel, margem esquerda do Rio Negro,
Lago Atàana, 10Oct 1978 (fl fr), Nascimento 590 (MG, NY); Rio Negro, Rio Curicuriari, 26 Oct
1978(fl), Nascimento 774(MG, NY); Rio Xeriuni, Proj. RADAM, Ponto 07A, SA-20-XA, 15Apr
1974 (fl fr), Pires et al. 13923 (IAN, INPA, MG, Z); Rio Curicuriari, at base of Serra Cujubi, 22
Jan 1948(fl), Schuhes &Lopez 9637B (MO); Prope Panure ad Rio Uaupés, Sep 1852(fl fr), Spruce
Agric univ. WageningenPapers87-1 (1987)

71

2445 (AWH,BM,BR,CGE,E,F,G,GH,GOET,K, LD,LE,MPU,NY,OXF,P,W).Without
definitelocality:nodate(fl),A.R.Ferreira243(LISU,P[lackingcollectorandcollectionnumber]).

Local names. COLOMBIA. Voila (Maguire et al. 36267). VENEZUELA. Boya
(Morillo etal.7580),Guacamari (Kuripako language,LI. Williams 14799),Palode Balso (Spruce 2445bis), Palo de boya (Cerda sn, Medina 250, 492, Pinate&Mondolfi 1044,Maguire etal.42627,LI. Williams14874),Palodeboya blanco(LI. Williams 13856,14799). BRAZIL. Molongó (Ducke RB 22420,44,163, Fróes
491h).
Molongum laxum is a riparian tree found in the upper Rio Negro region andadjacent areas of the upper Rio Orinoco. It was first described by BENTHAM(1841) under Tabernaemontana, based on a collection by SCHOMBURGK fromalongtheRioNegronearBarcelos.Other references (SCHOMBURGK, 1848; M O N ACHINO, 1945)list the type locality incorrectly as the Rio Cotinga on the upper
Rio Branco of Brazil. ALPHONSE D E CANDOLLE (1844) placed this species in
Hancornia.
MUELLER (1860a)transferred Tabernaemontana laxa toAmbelania, citing two
additional gatherings by Richard Spruce under this species. These two collections, from Panure [= Ipanoré] in present-day Brazil and from the Rio Casiquiare 'Vasiva et Pacimoni' in Venezuela, which were both cited under Spruce2445by MUELLER, arelisted herein as2445and 2445bis, respectively.
The earliest known collection of this specieswasmade in 1784by A. R. FERREIRA in Brazilian Amazonia (received on loan from the Lisbon herbarium[LISU]);according toFerreira'sitinerarylistedbyPRANCE(1971a)the collectionwasmadeintheupper RioNegro area.
Molongum laxum occursalong river margins inperiodically inundated areas,
and it is frequently found in large gregarious stands. Molongum laxum is morecloselyrelatedtoM. lucidumthantoM. zschokkeiforme; itiseasily distinguishedfrom thesetwospeciesbycharacterssuchasthelargerflowers and theleaf apex,
which isobtuseto acute, often mucronate.

4-2.Molongumlucidum(Humboldt, Bonpland &Kunth) Zarucchi, comb.nov.
Basionym: Tabernaemontana ? lucida Humboldt, Bonpland & Kunth, Nov.
Gen. et Sp. 7: 162 [folio], 209 [quarto]. 1825; Kunth, Synop. PI. 4: 229. 1825
[1826]; G. Don, Gen. Hist. Dicham. PI. 4: 92. 1837; A. De Candolle, Prodr.
8:377.1844.
Type: Venezuela: Amazonas: 'Crecit in aquis fluminis Pimichin (Missionesdel Atabapo),' May [ca 5May 1800](fl fr), Humboldt &Bonpland 990 (P-H&B[IDCMicrofiche 6209.75:II.2.- F photo 38732;GH, US],holotype; probableisotype [assn]: B-WILLD 5189, first sheet, nv [microfiche in MO]). Homotypicsymonyms: Lacmellea lucida(Humboldt, Bonpland & Kunth) Miers, Apocyn.
S. Amer. 14-15. 1878 as lLacmellia\ Ambelania lucida (Humboldt, Bonpland
72

Agricuniv. WageningenPapers87-1 (1987)

&Kunth)Markgraf, Notizbl.Bot.Gart. Berlin-Dahlem 15:620. 1941;Pichon,
Mém.Mus.Natl.Hist.Nat.24:169.1948subMolongumsect. Oligoon.
Heterotypic synonyms:Psychotria lucida Willdenow ex Roemer&Schultes,
Syst. Veg. 5: 189. 1819 (?Dec) [based on unnumbered duplicate specimen of
Tabernaemontana lucida Humboldt, Bonpland &Kunth in B-WILLD], nom.
illeg.,non Humboldt, Bonpland &Kunth, Nov.Gen.et Sp.3:282[folio],361[quarto], pi. 283. 1819 [21 Nov]. Ambelania cuneata Muell.-Arg. in Martius,
Flora Brasiliensis 6(1):17.1860;Markgraf, Notizbl. Bot.Gart. Berlin-Dahlem
15: 620. 1941;Monachino, Lloydia 8(2): 125-126. 1945; Schultes, Bot. Mus.
Leafl. 14(5):135.1950;J.Ethnopharm. 1:166.1979,synon.nov.Type:Venezuela:Amazonas:'Adfluminis GuainiavelRioNegrosupraostiumfluminisCasiquiari,' Jun 1854 (fl fr), Spruce3528 (BR, lectotype [here chosen]; isotypes:
AWH, BM,CGE, E, F 1539900,G [Fphoto 26841;GH, MO,NY, US],GH,
GOET,K, LD,LE,NY,OXF,P,W[assn]). Homotypic synonym: Molongum
cuneatum (Muell.-Arg.) Pichon, Mém. Mus.Natl. Hist. Nat. 24: 169,pi. IV 17,26,38.1948.
Spreadingshruborsmalltreeupto 12mtallwithleaf-bearing branches1.5-3
mm in diameter. Leaf blades obovate to oblanceolate, (3-)4-6.5(-9) x(1.1—)1.4—3(—4) cm,glabrous,coriaceous to thick-coriaceous, with 16-24pairs
oflateralveins;basesbroadly to sharplycuneateand decurrent onmost ofpetiole;apicesroundedtoobtuse,sometimesretuse;marginsentire,revolute;midribplaneorslightlyimpressedabove,prominentbelow;lateralveinsveryslightly
raised aboveand below.Petiole(4-)7-12mmlong,canaliculate. Inflorescences
terminal,cymose,congestedtosomewhatlax,(4-)15-^40(-55)-flowered;peduncles 15-45(-60) mm long; bracts deltoid, 2-3 x 1.2-1.8 mm; pedicels(4-)6-12(-14)mmlong;bracteolesovate,small,ca0.7-0.8mm.Flowers:maturebudca 13-14mmlong;sepalsovate, 1-1.8 x 1.3-2 mm,apicallyrounded.Corollawhiteorcreamwithtube 5-7mmlong,completelyglabrouswithin;lobes
oblong-elliptic, 7-8 x 4mm. Stamens inserted 2.5-3mmfrom baseofcorolla
tube;anthers2.5-3.3mmlong,sessile,basallyauriculate,apicallycaudatewith
acumen0.3-0.5mmlong.Ovaryovoid,apicallyrounded,ca 1.5mmlong;style
1.4-1.7mmlong;clavuncle0.7-1 mmlong;stigmoid lobesthin, terete,0.7-0.9
mmlong.Fruitellipsoid to narrowly ellipsoid, 1.5—3(—5) cmlong, apically andbasallyrounded.Seedsfew,2-4(-8),piano-toconcavo-convex,5.5-6.5 x 3.5-4
x 2-2.5mm.
Etymology. Latin lucidum = shiny, in reference to the appearance of the
adaxialleafsurface.
Distribution. Known from northwestern Amazonia ineastern Colombia,
southernVenezuela,andadjacent regionsofBrazil;Map19.

Agricuniv. WageningenPapers87-1 (1987)

73

MAP 19. Molongum lucidum(Humboldt, Bonpland &Kunth) Zarucchi.

Specimens examined:
COLOMBIA. Amazonas: Rio Caqueta, alrededores deAraracuara, Aeropuerto Araracuara, 10-22
Nov 1982 (fl fr), Idrobo et al. 11467 (COL). Vaupés: Rio Cuduyari, Yapobodâ, 23 Jan 1944 (fl),Allen 3271 (US); Rio Kubiyù, Cerro de Canendâ (Sabanas), ait 380-680 m, 2-4 Nov 1952 (fl),Garcia-Barriga 15068, 15075 (COL); Rio Vaupés, Cafio Pacû, 6 Mar 1944 (fl fr), Schuttes 5815
(COL,MO);RioKubiyù,CerroKanendâ, savannahsca 15miupstream from mouth, altca 800-900
ft, 19Nov 1952(fl), Schultes &Cabrera 18346(A,US);Mitü and vicinity, along lower Rio Kubiyù,31 Mar 1975 (fl fr), Zarucchi et al. 1137 (COL, GH, MO, US), 15 Apr 1975 (fr), Zarucchi 1210
(COL, GH, MO, US),25 Sep 1976(fl), Zarucchi 2139 (COL, ECON, GH, INPA, MO, US, USF,
WAG), 13Jul 1979(fl fr), Zarucchi 2455(COL, GH, MO), 15Jul 1979(st),Zarucchi 2480 (COL,
GH, MO), 16 Jul 1979 (st), Zarucchi 2482 (COL, GH, MO, WAG), 16 Jul 1979 (fl fr), Zarucchi
2483(COL,GH, MO).
VENEZUELA. Amazonas: Rio Guainia, along Cano Pimichin, below Pimichin, alt 125-135m, 14
Apr 1953 (fl), Maguire & Wurdack 35664 (NY); Rio Guainia, in sabanita 1km W of La Ceiba,
Cano San Miguel 2 km above Limoncito, alt 120-140 m, 14 Oct 1957 (fl), Maguire et al. 41898(NY, US, VEN); Rio Guainia, Maroa, alt 127 m, 11 Feb 1942 (fl fr), LI. Williams 14288 (F, G,

74

Agricuniv. WageningenPapers87-1 (1987)

MY, US); Rio Guainia, Playa Bianca, alt 127 m, 24 Feb 1942 (fl fr), LI. Williams 14454 (F, G,
MO, MY,NY,U, US).
BRAZIL. Amazonas: Rio Negro, S. Felipe, km 2 da Perimetral Norte, 21 May 1975 (fl), N. T.
Silva3858 (IAN).

Local names. COLOMBIA. A-BI-ta-ki (Kubeo language, Zarucchi 2455), Mimi-WAHO-ka-ki (Kubeo language,Zarucchi 2480,2482,2483).
ThespeciesacceptedhereasMolongum lucidumwasfirstdescribedby ROEMER
and SCHULTE?(1819)asPsychotria lucida(Rubiaceae),based onan unnumbered
Humboldt and Bonpland collection deposited in the Willdenow Herbarium at
Berlin.Thisnamewasa later homonym ofan actual rubiaceous plant, P.lucida
Humboldt, Bonpland and Kunth, the description of which also appeared inlate 1819 (21 November). According to MCVAUGH (1955) and STAFLEU and
COWAN (1979), the names published by Humboldt, Bonpland and Kunth inVol. 3(fascicle 12)of the Nova Generaet Species Plantarum have priority overthe competing names which appear in ROEMER and SCHULTES (1819). HUMBOLDT, BONPLAND and KUNTH (1825), also in the Nova Generaet Species Plantarum (Vol. 7), described 'Tabernaemontana ? lucida', based presumably on a
duplicate of the same Humboldt and Bonpland collection. The type specimen
isdeposited inthe separate Humboldt and Bonpland Herbarium at Paris. Latertaxonomietreatmentscontinued toplaceM. lucidumquestionably under Tabernaemontana (e.g.,G. DON, 1837,and A. D E CANDOLLE, 1844).
MUELLER (1860a)didnot account for Tabernaemontana lucidainthe Apocynaceaefor theFloraBrasiliensis.Instead hedescribed Ambelania cuneata [= Molongum lucidum]based on Spruce 3528,from the Rio Guainia of the upper Rio
Negrobasin. MIERS(1878),inhiswork onSouthAmericanApocynaceae, placed
T.lucidainLacmelleaÇLacmellid). MARKGRAF(1941),inhisrevisionofLacmelleaand review ofAmbelania, made thenewcombination A. lucida. MONACHINO(1945) did not consider A. lucida; he was apparently unaware of MARKGRAF'S
1941 articleduetohostilitiesduringWorldWar II. PICHON(1948)listedA. lucida
within Molongum sect.OligoonalongwithM. cuneatum and M. zschokkeiforme,
but did notmake thecombination inthe genus.
Molongum lucidumisa rather variable speciesdistributed inthe northwestern
Amazon, and is known from both savanna and riparian habitats. Specimensof M. lucidum are distinguished easily from M. laxum, partly on the basis ofthebroad tonarrow cuneateleaf bases.
Individuals of Molongum lucidum are able to sucker from the base, enablingthem to survive fire and other forms of destruction in normally inhospitableenvironments.Thosecollectionsmadefrom alongstreamsmay actually beindividualswhichhavegrown from seedsdispersed from thesavanna habitats.

4-3.Molongumzschokkeiforme(Markgraf) Pichon,Mém.Mus.Natl.Hist. Nat.
24: 169,pi.IV-16,27, 37.1948.

Agricuniv. WageningenPapers87-1 (1987)

75

iftfr
FIG. 5.Molongum zschokkeiforme (Markgraf) Pichon: 1. flowering branch, \ x ;2. opened flower,
3 x ; 3. fruits, | x ; 4 . habit. (1.Schuhes & Cordeiro 6514; 2. Zarucchi et al. 2937, ale. mat.; 3.
Zarucchi etal.2902,ale.mat.;4.Zarucchi etal.2902,from slide).

76

Agricuniv. WageningenPapers87-1 (1987)

Basionym:AmbelaniazschokkeiformisMarkgraf,Notizbl.Bot.Gart. BerlinDahlem 12:295-296. 1935;15:620. 1941;Monachino, Lloydia 8(2): 125. 1945;
9(4):302.1946;Schultes,Bot.Mus.Leafl. 14(5):135-136,pi.30. 1950.
Type:Brazil:Amazonas:Maués,adripasinundatas RioMoraes,4Oct 1929
(fl fr), DuckeRB 22421 (B, holotype, lost; isotypes: G, K, P, ?RB nv, S, U,
US1743767).
Small tree up to 5mtall with leaf-bearing branches 1.5-3 mm in diameter.
Leaf blades lanceolate or ovate to elliptic, 5.5—10(—12) x 1.3—3.5(—5) cm,glabrous,subcoriaceous to thin-coriaceous,with9-16pairs oflateralveins;basesrounded anddecurrentonupperportion ofpetiole;apicesacute,rarelyobtuse,distallyrounded;marginsentireorcrenulate,revolute;midribimpressedabove,
especiallynearbaseofblade,prominentbelow;lateralveinsslightlyraisedaboveand below. Petioles 9—15(—28) mm long, shallowly canaliculate. Inflorescences
terminal, cymose, congested, 10-28(-45)-flowered; peduncles 9—16(—26)mm
long;bractsdeltoid,2.5-3 x 1.2-1.5mm;pedicels2-3.5mm;bracteolesdeltoid
to ovate, ca 1.5 x 1 mm near base of pedicel. Flowers: mature bud ca 9mm
long; sepals ovate, 1.5-2 x 1.4-1.6 mm, apically rounded. Corolla white or
cream with tube 3-4.5 mm long, completely glabrous within; lobes obovate,
5 x 3 mm.Stamensinserted ca2mmfrom baseofcorolla tube;anthers 2-2.5
mmlong,sessile,basally auriculate,apically caudatewith acumen0.3-0.5mm
long. Ovarycylindrical to ovoid, apically truncate, 1.1-1.3 mm long; style0.3-0.6mmlong;clavuncle 0.8-1 mmlong; stigmoid lobes terete,0.2-0.3mm
long. Fruitnarrowly ellipsoid or lanceovoid, 2-4(-6.5) cm long and 0.6-1 cm
indiameter,apicallyobtuse,basallyrounded.Seedsfew,2-5(-7),piano-toconcavo-convex,ca6 x 3.5 x 2mm.
Etymology. Theepithetzschokkeiformereferstothesimilarityinleafshape
betweenthisspeciesandZschokkea,agenericsynonymofLacmellea(Apocynaceae- Carisseae).
Distribution. Knownfrom tributaries ofthemiddletolowerRioMadeira
andeastwardinAmazonas,Brazil;Map20.
Specimens examined:
BRAZIL. Amazonas: Regiäo do Rio Madeira, Rio Canumä, 25Oct 1957(fl), Frees 33675 (COL,
IAN), 5 Nov 1957 (fl fr), Fróes 33738 (IAN, MO, NY); basin of Rio Madeira, on shore of lakenear River Marmellos, 29 Nov 1933 (fr), Krukoff 5822 (A, K, NY, S, US); Mun. de Barreirnha,
igarapé-acü do Rio Andirâ, 25Mar 1974(fl fr), Mello U/74 (INPA); Rio Maués, Mun. de Maués,no date (fl), W. Rodrigues &L. Coêlho 6779 (GH, INPA); Rio Marmellos, Igarapé Pote, 6 Aug
1945 (fl), Schultes & Cordeiro 6514 (GH, IAN, K, MO, US); Rio Madeira, Rio Marmellos, nearmouth, 12 Aug 1945 (fl), Schultes & Cordeiro 6533 (GH, IAN, MO); Rio Marmellos, Aug 1948(fl), Schultes &Lopez 10310(GH, IAN, K, MO, US);Mun. de Borba, lower Rio Canumä, E bankof river ca 2 km Sof Vila of Canumä, Lat. 4° 04' S, Long. 59°05'W, 27Jun 1983(fl fr), Zarucchi
et al. 2879 (INPA, NY, US); Mun. de Borba, lower Rio Canumä, W bank of river ca 1km S ofVila of Canumä, Lat. 4° 03' S, Long. 59° 06' W, 28 Jun 1983 (fl fr), Zarucchi et al. 2902 (INPA,
NY, US); Mun. de Axinim, basin of Rio Abacaxis, lower Rio Marimari, ca 2 km below mouthof Rio Paca, Lat. 4° 07' S, Long. 58° 58' W, 2 Jul 1983 (fl fr), Zarucchi et al. 2937 (INPA, NY,
Agricuniv. WageningenPapers87-1 (1987)

11

MAP 20.Molongum zschokkeiforme (Markgraf) Pichon.

US); Mun. de Axinim, along Rio Abacaxis ca 4 km below mouth of Igarapé Surubim, Lat. 4°
29' S, Long. 58° 28' W, 4 Jul 1983 (fl), Zarucchi et al. 2952 (INPA, NY, US); Mun. de Maués,basin of Rio Maués, lower Rio Parauari, ca 3 km N of Mucajâ, Lat. 3° 52' S, Long. 57° 30' W,
18Jul 1983(fl), Zarucchi et al.3071 (INPA, NY, US); Mun. de Maués, basin of Rio Apoquitaua,
lower Rio Pacoval, Lat. 3° 48' S, Long. 58° 04' W, 26 Jul 1983 (fl), Zarucchi et al. 3180 (INPA,
NY, US);Mun.deMaués,RioApoquitaua, Lat. 3°59'S,Long.57°54'W,27Jul 1983(fl), Zarucchi
et al. 3208 (INPA, NY, US); Mun. de Maués, Rio Apoquitaua just above confluence with Rio
Pacoval, Lat. 3°47' S,Long. 57°59'W,27Jul 1983(fl), Zarucchi etal.3217(INPA, NY, US).

Local names. BRAZIL. Boia (Schuttes &Cordeiro 6514, 6533), Molongó (W.
Rodrigues & L. Coêlho6779, Zarucchi et al.2879, 2902,2937, 2952, 3217), Molongódo Igapó(Mello 11/74).
Molongum zschokkeiforme was described under Ambelania by MARKGRAF in
78

Agricuniv. WageningenPapers87-1 (1987)

1935 based on a collection by ADOLPHO DUCKE from central Amazonia at
Maués.The speciesisknown only from thelower Rio Madeira, its southeasterntributaries,andthevariousparanasconnectingtheRioMadeiratotheRio Amazonas.
Intheprotologuefor thespecies,MARKGRAFcompared Ambelania zschokkeiformis withtheillustration ofA. acida(AUBLET, 1775),distinguishingA. zschokkeiformis by its shorter corolla tube. MONACHINO (1945), in his treatment ofAmbelania in thewide sense,indicated that a closerelationship existed between
A. zschokkeiformis and A. laxa, based on their terminal inflorescences (versus
axillary inflorescences inA. acida),and their similar habitat ininundated areas.
SCHULTES (1950)provided further data on A. zschokkeiformis, based on his several gatherings made on the Rio Marmellos, a tributary of the Rio Madeira.
He described them as gregarious trees, usually with the trunk thickened at thebase, growing on sand bars along the river. In this article, SCHULTES provided
a habit and habitat photograph of the trees,which are almost completely submergedfor part oftheyear.
PICHON(1948)placedAmbelania zschokkeiformis inhisnewgenus Molongum,
sect. Oligoon, along with M. cuneatum [= M. lucidum] and A. lucida [= M.
lucidum]. Molongum zschokkeiforme isdistinguished from the other two speciesof Molongum byitssmallerflowers, shorterflower pedicels,thinnerleaveswhichsometimes haveacrenulatemargin,and generally longer petioles.
Theholotypeof Ambelania zschokkeiformis wasdestroyed duringWorld War
II (MARKGRAF, pers.comm.),necessitating theselection of alectotype.A duplicateof thetypecollection, Ducke RB 22421, has not been seen from the Jardim
BotanicoinRiodeJaneiro(RB);therefore, alectotypehasnotbeenchosen.
Excluded Speciesof Molongum
Molongum macrophyllum (Muell.-Arg.) Pichon, Mem. Mus. Natl. Hist. Nat.
24: 168,/»/.IV-14,24, 35. 1948 = Spongiosperma macrophyllum (Muell.-Arg.)
Zarucchi.
Molongum grandiflorum (Huber) Lemée, Fl. Guy, Franc. 3:297. 1953[comb,
illeg.] = Spongiosperma grandiflorum (Huber) Zarucchi.

5. Rhigospira Miers, Apocyn. S.Amer. 67-71,pi. 10A. 1978pro parte; Pichon,
Mém.Mus.Natl. Hist.Nat. 24:169,pi.
IV-18,28,39.1948.
Rigospira O.Kuntze inPost &O.Kuntze, Lexicon 1904,orth. mut.
Ambelania sect. Terminales Markgraf, Notizbl. Bot.Gart. Berlin-Dahlem 15:
620.1941pro parte,nom. nud.
Ambelania sect. Quadrangulariae Monachino, Lloydia 8(2): 114. 1945, nom.
nud.
Smallormedium-sizedtreeupto30mtall,lactescent,withyoungstemssharply quadrangular becoming terete with age and secondary growth; interpetiolar
lineorridgeconspicuous,containingmany small,resin-producing glandswithin
Agricuniv. WageningenPapers87-1 (1987)79

leaf axils on long, canaliculate petiole. Leaves decussate, petiolate, glabrous,
stiff-coriaceous, venation eucamptodromous, margins entire and revolute;
apices obtuse, mucronate; bases variable, acute to subcordate, slightly or markedly decurrent on petiole; midrib plane or slightly raised and striate above,
prominent below; lateral veins slightly raised above and below; tertiary veinsslightly raised above. Inflorescences terminal, cymose, lax and well-branched,
many-flowered; peduncles long, secondary inflorescence axes almost aslong as
peduncle; axes terete to subterete; bracts ovate to deltoid; bracteoles deltoid,
usually on lower half of pedicel;pedicels terete,expanding unequally becoming
frustumic withmaturefruit. Flowerslong,calyxlobes5,persistent, quincuncially
imbricate, basally lacking glands within; sepals free nearly to base, eciliate,
ovate,apically rounded to obtuse.Corolla sympetalous, actinomorphic, salverform, white, thick; tube narrow, straight, long, sparingly pilose from insertionof stamens to orifice; lobes 5, convolute in bud, overlapping to left, becoming
reflexed at anthesis, less than one-half length of tube. Stamens 5,inserted nearbase of tube; anthers lanceolate, subsessile with filaments short, basally auriculate, apically caudate. Ovary superior, cylindrical, glabrous, apically truncate,bicarpellate,placentation axilewithovulesnumerous ineachlocule;styleterete,
short; clavuncle cylindrical with basal flange, five-lobed apically, star-shapedin cross-section, glabrous; stigmoid lobes terete, thin, approximating length ofclavuncle.Fruitmany-seeded,one-celledberry,ellipsoidtoobovoid, occasionally stipitate, with seedsembedded in a crisp, red or maroon, acidic, edible pulp;
pericarp indurated witha granular composition, surface smooth, glandular, becoming rough, muriculate when dry. Seeds plano-convex, naviculate, ovate to
oblonginoutlinewithprominent dorsal ridge;testairregularly reticulate;hilum
small, ovate to circular. Embryo almost as long as seed, with small, thin, plane,broadly ovate cotyledons; radicle straight, terete, about 5-7 times as long as
cotyledons.
Type species. Rhigospira quadrangularis (Mueller-Argoviensis) Miers, Apocyn. S. Amer. 68,pi. 10A. 1878 (based on Ambelania quadrangularis MuellerArgoviensisinMartius, Flora Brasiliensis 6(1): 18.1860);thegenuswas typified
byPichon (1948)bytheexclusion of theother sixspeciesofMiers.
E t y m o l o g y . Greek piyócu {rigescoin Latin) = to become rigid, spiral =CTTieïpa (spirain Latin) which according to MIERS (1878) 'refers to the rigidly
pyramidal form oftheborder ofthecorolla in aestivation.'D i s t r i b u t i o n . Central and western Amazonia of Brazil (Acre, Amazonas,
Roraima), Peru (Amazonas, Loreto), Colombia (Amazonas, Guaviare,
Vaupés),and Venezuela (Amazonas),upper RioOrinocodrainagein Venezuela(Amazonas, Bolivar); also the Pacific coast of Colombia (Chocó, Valle); Map
5or21.
RhigospirawasdescribedbyMIERS(1878)toencompasssevenspeciesofAmazonian treeswhich had previously been treated under Ambelania and Tabernaemontana sect. Stenocephalium by MUELLER (1860a)in Martius' FloraBrasilien80

Agricuniv. WageningenPapers87-1 (1987)

sis.ThispresentworkconsidersRhigospira, ascircumscribed by MIERS,torepresentspeciesinSpongiosperma, Neocouma, Macoubea, andtheemended Rhigospira. Four of Miers' species of Rhigospira are referable to Macoubea, which is
now treated as part of the subfamily Tabernaemontanoideae (BOITEAU &SASTRE, 1975; BOITEAU, ALLORGE, & SASTRE 1978); references to these species
areprovided attheend ofthissectionunder 'Excluded Speciesot Rhigospira.'
Rhigospira wasnot accepted byseveralsubsequent reviewersofthe Apocynaceae(BÂILLON, 1891; SCHUMANN, 1895),whoplaceditinsynonymyunder Tabernaemontana. MONACHINO (1945)mentioned Rhigospira in histreatment of Ambelaniabut didnot recognizeitasavalid genus.
PICHON (1948)accepted Rhigospira asamonotypicgenus,which helectotypified by the exclusion of the other sixspecies.The genus isreadily distinguishedevenwithsterilespecimens,asindicatedinthediscussionunderR.quadt-angularis, and it is also the only genus in the Ambelanieae for which collections havebeengathered westoftheAndean cordillera.

5-1.Rhigospiraquadrangularis(Muell.-Arg.)Miers,Apocyn. S.Am.68,pi. 10A.
1878;Pichon, Mem.Mus.Natl. Hist.Nat. 24: 169,pi. I V - 18,28, 39.1948.
Basionym:Ambelania quadrangularisMuell.-Arg.inMartius,Flora Brasiliensis 6(1): 18. 1860;Markgraf, Notizbl. Bot. Gart. Berlin-Dahlem 15:620.1941;
Monachino, Lloydia 8(2): 124. 1945;9(4):302. 1946;Macbride, Fl. Peru, Publ.
Field Mus.Nat. Hist., Bot. Ser. 13(5/1):374.1959.
Type:Venezuela: Amazonas: 'Prope San Carlos ad Rio Negro, ad basio Cocui,' Jul 1853 (fl), Spruce 3026 (BR, lectotype [here chosen]; isotypes: AWH,
BM,CGE,E,F 1539870,G [Fphoto 26843;F,GH, MO,NY,US],GH, GOET,
K, LD, LE, NY, OXF, P,W).Homotypic symonym: 'Hancornia ?macrophylla
Spruce' ex Muell.-Arg. in Martius, Flora Brasiliensis 6(1): 18. 1860,nom. nud.
insynon. under A. quadrangularis.
Smallormedium-sized treeup to 30mtallwithleaf-bearing branches sharply
quadrangular, 3-8 mmindiameter.Leaf bladesoblong-elliptictonarrowly obovate, 8-22 x 4.5-8 cm, glabrous, stiff-coriaceous, with 11-18 pairs of lateralveins;basesnarrowed,obtusetorounded,rarelysubcordateorsubacute,slightly
decurrent on petiole;apicesobtuse,mucronate;marginsentire,revolute; midrib
plane or slightly raised and striateabove,prominent below;lateralveins slightly
raised above and below; tertiary veins conspicuous and slightly raised above.
Petioles 12-30 mm long, canaliculate, with many glands in leaf axil. Inflorescencesterminal, cymose,well-branched, lax,usually 60-90-flowered; peduncles
25-50(-60) mm long; bracts ovate to deltoid, 3-3.5 x 2-3.5 mm; pedicels 4-8
mm long; bracteoles deltoid, ca 1.5 x 1mm, usually on lower half of pedicel.
Flowers:maturebud 23-28mmlong;sepalsovate, 1.2-2 x 1.3-1.8mm, apicallyrounded to obtuse. Corolla white with tube 11-15 mm long, sparingly pilose
from insertion of stamens to orifice; lobes oblong-elliptic, 10-11 x 3.5-4 mm,
Agricuniv. WageningenPapers87-1 (1987)

81

FIG. 6. Rhigospira quadrangularis(Muell.-Arg.) Miers: 1.flowering branch, \ x ;2.opened flower,
3 x; 3. sectioned fruit, | x . (1-2. Zarucchi 2428, ale. mat. & slides; ; 3. Zarucchi 1570, ale. mat.
&slides).

82

Agricuniv. WageningenPapers87-1 (1987)

MAP 21. Rhigospira quadtangularis (Mueller-Argoviensis) Miers.

reflexed. Stamensinserted 2.5-3 mmfrom base ofcorolla tube;anthers 3 mm
long, subsessile,filaments ca 0.3mmlong;bases auriculate,0.3mmlong;apicallycaudatewithacumen0.4-0.5mmlong.Ovarycylindrical,1.2-1.4mmlong;style0.5-0.7mmlong;clavuncle0.5-0.6mmlong;stigmoidlobesthinandterete,
0.5-0.7mmlong.Fruitellipsoidtoobovoid,8-10cmlongand4-7cmindiameter;apicallyroundedtoobtuse,withlargerfruits usuallystipitate.Seedsnumerous,usually30-60,embedded inaredormaroon pulp,plano-convex,9.5-11.5
x 4-5 x 2-3mm;surfacefinelyandirregularlyreticulate.
Etymology. The epithet quadrangularis refers to the sharply four-angled
youngstemsoftheplants.
Distribution. KnownfromwidespreadlocalitiesinwesternColombia,centralandwesternAmazonia ofBrazil,Colombia, andPeru;alsocollectedinthe
region of the upper Rio Orinoco of Amazonas and Bolivar, Venezuela; Map
21.
Agricuniv. WageningenPapers87-1 (1987)

83

Specimens examined:
COLOMBIA. Chocó: Rio Atrato, Samurindó, alt ca 80 m, 1Apr 1958 (fr), Cuatrecasas & Llano
24100 (MO, US); woods E of beach ca 1mile N of Juarin, alt 10-100 m, 24 May 1967 (yng fr),
Duke 11499 (ECON, NY, OS); Mun. de Quibdó, carretera Quibdó-Guayabal, 11 May 1985 (fr),Moreno et al. 002 (MO); Mun. de Bahia Solano, Cgto. El Valle, colinas detrâs del pueblo, alt 35
m, 4 Oct 1983(fr), Sanchez &Rendon 601 (MEDEL). Guaviare (as 'Vaupés'): Path between Rios
Itilla and Macaya, alt ca 900 ft, 1-7 Apr 1943 (fr), Schuhes 5357 (COL, ECON, K, US). Valle:
Bajo Calima, ca 10 km due N of Buenaventura, Carton de Colombia concession, transect 1, 3°
56' N., 77°08'W.,altca 50m, 6Dec 1981(st),Gentry 35386(COL, MO);Bajo Calima, Concesión
Pulpapel/Buenaventura, 3° 55' N, 77° W, ait ca 100 m, 30 Nov 1984 (fr), Monsalve 594 (MO),26 Feb 1985 (fl fr), Monsalve 702 (MO). Vaupés: Mitû and vicinity, along road from town, 15
May 1976(fr), Zarucchi 1570(COL, ECON, GH, MO,US,USF),22Jun 1976(flbuds,fr), Zarucchi&Balick 1711 (COL, GH, MO, US, USF), 5 Jul 1976 (fl), Zarucchi et al. 1812 (COL, GH, MO,
US, USF), 4 Jul 1979 (fl), Zarucchi 2428 (COL, GH), 2431 (COL, GH, USF), 2432 (COL, GH,
WAG), 22 Jul 1979(fl), Zarucchi 2509 (COL, GH, MO); along Rio Vaupés between Rio Paranâpichuna and Rapids ofMandi, 19Jul 1979(fl),Zarucchi 2497(COL, GH, MO,WAG), 2500(COL,
GH, MO, USF, WAG).
VENEZUELA. Amazonas: Réserva Forestal El Sipapo, bosque a la margen derecha Rio Sipapo,
May 1971 (yng fr), Blanco 1192 (NY, US, VEN). Bolivar: Rio Chizco, Uriman, alt 450-500 m,
Sep 1953 (st), Bernardi 920 (NY); region de los Rios Icabaru, Hacha, y cordillera sin nombre, alt
450-850m, 2Jan 1956(yng fr), Bernardi 2738 (NY, VEN); selvas Rio Curutu, alto [Rio] Paragua,
alt 550 m, Apr 1948 (fr), Cardona 2487 (MO, US, VEN); Chimantâ Massif, lower NW slopes ofAbâcapa-tepui, vicinity ofcamp 1along Rio Abâcapa, alt420m, 30-31 Mar 1953(fr), Steyermark
74748(F, MY, NY, US, VEN); Chimantâ Massif, lower SW-facing forested slopes of Chimantâtepui (Torono-tepui), at extreme S end on N side of Rio Tirica (left-hand branch), alt 1000 m,
14May 1953(fr), Steyermark 75361 (F,NY, VEN).
PERU. Amazonas: Rio Santiago, 2 km bajo de La Poza, alt 180 m, 16Aug 1979 (fr), Asuncion
162 (GH, MO); Mobil Oil Co. trail, 3 km from La Poza, E bank of the Santiago River, alt 180
m,6Aug 1979(fr), Berlin3519(GH, MO,USF);Rio Santiago,2kmâtrasdelacomunidad Caterpiza,bandaestedeCaterpiza,alt 180m, 15Oct 1979(fr),Huashikat 942(MO). Loreto:Prov. Requena,
Dtto. Sapuena, Jenaro Herrera, ait 120 m, ârbol no. 97, 20 Jun 1974 (fl), Abadie 25T (IAN, Z);
Jenaro Herrera, Arboretum, 5 Feb 1974 (fr), Diaz 47A (G), ârbol no. 2/47, 15 Jun 1974 (fl fr),
Diaz 58A (G, IAN); Prov. Maynas, Caserio de Huanta, Rio Orosa, alt ca 120 m, 31 Aug 1978(fr), Diaz et al. 477 (F, GH, MO); Prov. Loreto, Rio Tigre, Campamento Petrolero San Jacinto,
2° 15' S, 75° 50' W, alt ca 200 m, 16Sep 1979 (fr), Diaz &Jaramillo 1471(MO); Jenaro Herrera,margen derecha Rio Ucayali,24May 1982(fl buds),Encarnación26154(MO,US);Jenaro Herrera,
tree number R-114 PP, 15 Oct 1973 (fr), Noriega sn (US); Jenaro Herrera, 4° 50' S, 73° 45' W,
alt ca 170m, 10Dec 1980 (fr), Vasquez &Jaramillo 1029(MO, US), 1Jul 1981 (yng fr), Väsquezetal.2056(MO).
BRAZIL. Acre: Cruzeiro do Sul, Estrada Alemanha, 6 May 1971 (fl fr), Maas et al. P12717 (C,COL, F, G, INPA, M, MG, MO, NY, P, S,U, US), 4 Nov 1966(fr), Prance et al.3013 (F, INPA,
K, MG, NY, S, US), 13 Apr 1971 (fr), Prance et al. 11807 (F, INPA, K, MG, NY, S, U, US);proximo do Aeroporto de Cruzeiro do Sul, 8 Feb 1976 (fr), Marinho 33 (IAN), 6 Feb 1976 (fr),Monteiro &Damiäo [Mola] 142 (INPA), 8 Mar 1976 (fr), J. Ramos &Mola 344 (INPA); vicinityof Serra da Moa, 22 Apr 1971 (fl buds, fr), Prance et al. 12242 (F, INPA, K, M, NY, U, US),12267 (F, INPA, K, M, MG, NY, P, S, U, US); Rio Moa, 8 km above Cachoeira Grande, 27
Apr 1971 (fl buds, fr), Prance et al. 12565(F, GH, INPA, K, M, MG, NY, S, U, US). Amazonas:
Rio Solimôes, Sâo Paulo de Olivença, 20 Aug 1929 (fl), Ducke RB 22419 (K, P, RB, S, U, US);Rio Madeira, Borba, 24 Apr 1937 (fl), Ducke RB 35159 (G, K, P, RB, S, U), 471 (A, F, K, MO,
NY, S,US);Manaus, Réserva Florestal Ducke, 26Feb 1957(fl fr), E. Ferreira57-17 (IAN, INPA,
MG, MO),9Apr 1962(fr), W. Rodrigues & Chagas3583(INPA), 18May 1962(fl fr), W. Rodrigues& Coêlho 4453 (INPA), 13 May 1964 (yng fr), W. Rodrigues & Loureiro 5814 (INPA), 11 Mar
1966 (fr), W. Rodrigues & Coêlho 7559(INPA); Rio Solimôes, Vargem Grande, 31 Jul 1941 (st),Fróes 12153/63 (A, F, MICH, NY, US); Rio Negro, Foz do Caiary, 12Oct 1945(fr), Frees 21149(F, IAN, K, NY, TEX, US);Rio Negro, Padauiry, mata do Tucano, 14Oct 1947(fl), Fróes 22595
84

Agricuniv. WageningenPapers87-1 (1987)

(IAN, NY, RB, SP, U, US, VEN); Rio Negro, Preto, Matupiry, 14Nov 1947 (fl fr), Fróes 22853(IAN); Rio Tefé, Muquentâua, 10Jun 1950 (fr), Fróes 26196 (IAN, NY, UC); Rio Javari behind
Palmeiras Army Post, 72° 49' W, 5° 8' S, 31 Jul 1973 (fl fr), Lieras et al. P16964 (INPA, MG,
NY, Z), 1Aug 1973(fl fr), Lieras et al. P16985 (INPA, NY); Manaus-Pôrto Velho Highway (BR319), behind road camp at km 240, 22 Nov 1973 (fl buds, fr), Lieras et al. P19615 (INPA, MG,
NY, US, Z); Rio Javari, 7 hrs above Paumari, 16 Oct 1976 (fr), Prance et al. 23837 (INPA, MG,
NY, WAG); Upper Rio Negro,without preciselocality, 1907-1908 (fr), Weiss & Schmidt 54 (NY).
Roraima: vicinity of Uaicâ airstrip, Rio Uraricoeira, 3° 33'N, 63° 11'W, 1Mar 1971 (fr), Prance
etal.10804(COL,F,INPA, K MG, NY, S,U, US,VEN).

Local names. COLOMBIA. Caimo morado {Schuhes 5357), Guayabilla {Monsalve594), Guayabilla de monte (Monsalve 702),Juansoco de Danta (Zarucchi
2497, 2500), Mi-mi-WAHO-ka-ki (Kubeo language, Zarucchi 2509), Palo delèche {Schuhes 5357). VENEZUELA. Aratani-yek (Arekuna language, Bemardi920), Guayaba de monte {Bemardi 2738). PERU. Yahuar caspi {Abadie 25T),
Yahuar huayo {Diaz47A, Noriega sn),Yahuar huayo Colorado {Diaz58A), Yapûkait(Huambisa language,Berlin3519,Huashikat 942). BRAZIL. Goiaba d'anta{DuckeRB 22419).
Rhigospira quadi•angularisisa widespread speciesfound growing on terrafirma with individual trees reported up to 30 m tall. It is separated easily fromall other described species of Ambelanieae by its sharply four-angled young
branches.Other distinguishing characters include theelongate, many-flowered,and lax inflorescences and the large, ellipsoid to obovoid, often stipitate, fruitwith a granular, indurated pericarp. Rhigospira quadrangularis exhibits additional interesting characters such as the numerous glands in the leaf axils, theglands on the surface of the fruit, the red to maroon, crisp, acidic fruit pulp,
and the eucamptodromous leaf venation. Foliage characteristics vary widely
within the species, with leaves gathered from the same tree varying markedly
insizeand intheshape oftheleaf base.
The first collection of Rhigospira quadrangularis was made by RICHARDSPRUCE near the base of Serra Cocui, on the present-day border of Venezuela
and Brazil along the Rio Negro. In the field, SPRUCE questionably attached thename lHancornia macrophylla to the specimens, a new species which he had
early encountered and provisionally named on the Rio Uaupés. MUELLER(1860a)included Spruce's questioned determination in synonymy under Ambelaniaquadrangularisand placed H. macrophylla under A. macrophylla. Further
discussion may befound under Spongiosperma macrophyllum.
MIERS (1878) placed Ambelania quadrangularis as one of seven species in his
newgenusRhigospira and accompanied itwithalinedrawingbased on Spruce's
collection.SubsequentworkersdidnotacceptMiers'genusand treated Rhigospira quadrangularis under Ambelania. In 1948, however, PICHON accepted Rhigospira with one species; R. quadrangularis therefore became the generic type
bytheexclusion oftheother sixspecies.
Excluded Speciesof Rhigospira
Rhigospira paucifolia (Spruce ex Muell.-Arg.) Miers, Apocynaceae South
Agricuniv. WageningenPapers 87-1 (1987)

85

America 69. 1878 = Macoubea sprucei (Muell.-Arg.) Markgraf var. paucifolia(SpruceexMuell.-Arg.) Monachino.
Rhigospira reticulata (A. DC.) Miers, Apocynaceae South America 69-70.
1878 = Macoubea guianensis Aublet.
RhigospirasinuosaMiers,Apocynaceae SouthAmerica70.1878 = Macoubea
sprucei (Muell.-Arg.) Markgraf var.paucifolia (Spruce ex Muell.-Arg.) Monachino.
Rhigospirasprucei(Muell.-Arg.)Miers,Apocynaceae SouthAmerica 70.1878= Macoubea sprucei(Muell.-Arg.) Markgraf.
Rhigospira ternstroemiacea (Muell.-Arg.) Miers,Apocynaceae South America 71. 1878 = Neocouma ternstroemiacea (Muell.-Arg.) Pierre.
Rhigospira venulosaMiers,ApocynaceaeSouthAmerica68-69.1878 = Spongiospermamacrophyllum (Muell.-Arg.) Zarucchi.

6.NeocoumaPierre,Buil.Soc.Mens.Soc.Linn.Paris,sér.2,5:33.1898; Pichon,
Mém. Mus.Natl.Hist.Nat. 24:169-170,/?/. IV-19,29,40.1948.
Neocuma O.KuntzeinPost &O.Kuntze, Lexicon 386.1904,orth. mut.
Ambelania sect. Neocoumae Monachino subsect. Ternstroemiaceae Monachino,Lloydia 8(2): 113.1945,nom. nud.
Small or medium-sized trees up to 25 m tall, lactescent, with stems terete or
subterete, covered with large petiole scars and with conspicuous interpetiolar
line or ridge. Leaves with small saucer- or flap-shaped gland in leaf axil, decussate,petiolate,glabrous,coriaceous to thick-coriaceous, sclereidseither present
orabsent,venationbrochidodromous,marginsentireandrevolute,blade decurrentonshortorlong,canaliculatepetiole;midribsomewhatprominent or slightly impressed above, prominent below; lateral veins slightly raised or impressed
above, slightlyraised below;tertiary veinsconspicuous on both surfaces. Inflorescences terminal, cymose, congested, many-flowered; peduncles elongate;
bractsand bracteolescalycine,small,deltoid to ovate, or large,ovate to elliptic;
bracteoles located at or above middle of pedicels.Flowerseither small or large;
calyxlobes5,caducousor apparently persistent, quincuncially imbricate, basally with many small glands in several series within; sepals free nearly to base,ovate toelliptic,apically rounded to acute,either entire or finely ciliateat apex.
Corolla sympetalous, actinomorphic, salverform, white, thick; tube narrow,straight, either short or long, pilose within from below insertion of stamens,
where tube is dilated, up to the orifice; lobes 5, convolute in bud, overlapping
to left, becoming reflexed at anthesis. Stamens 5,inserted near middle of tube;
anthers lanceolate, sessile, basally auriculate or sagittate, apically caudate. Ovarysuperior, ovoid,glabrous,apicallyrounded ortruncate,bicarpellate, placentation axilewith ovulesnumerous ineach locule;style terete,short or long;clavunclecylindrical orglobose,with basal flange, five-lobed apically, star-shaped
incross-section, glabrous;stigmoidlobesterete,thin,equaltoorlessthan lengthof clavuncle. Fruit few- to many-seeded, two-celled berry, ellipsoid or globose
86

Agricuniv. WageningenPapers87-1 (1987)

becoming stipitate, with seeds embedded in a somewhat stringy pulp; pericarp
crustaceous, thin, surface verrucose. Seedsplano-convex or unequally compressed, ovate to elliptic in outline;testa covered with low, irregular protuberances
and irregularly pitted; hilum small, ovate to circular. Embryo almost as longas seed, with small, thin, plane, ovate or circular cotyledons; radicle straight,
terete,about 3-5 timesaslongascotyledons.
Type species. Neocouma ternstroemiacea (Mueller-Argoviensis) Pierre, Bull.
Mens. Soc. Linn. Paris, sér.2, 5:33-34. 1898(based on Tabernaemontana ternstroemiacea Mueller-Argoviensis inMartius, Flora Brasiliensis6(1): 88.1860).
Neocouma was described in 1898 by PIERRE to encompass a single species,
N. ternstroemiacea, which had been first described by MUELLER (1860a)as 'Tabernaemontana ?ternstroemiacea''.Thisspeciesdescriptionwasbasedonacollectionmadeby RICHARD SPRUCEfrom theupper RioNegroin 1853. MIERS(1878)
placedMueller'sspeciesinRhigospira, althoughhenoted thatseveral characterswhich he listed 'might lead to a supposition of its being the type of a distinct
genus.' PIERRE (1898) introduced the genus with a discussion of its apparent
similarities to Couma (Carisseae). In his description of N. ternstroemiacea hecommented upon thepresence offoliar sclereids.
A second species, Neocouma duckei, was added to the genus by MARKGRAF
in 1932,based on a collection by ADOLPHO DUCKE from Fontebôa, Amazonas,
Brazil.In thepresent treatment itisplaced inthenewgenus Mucoa.
MONACHINO (1945) treated Neocouma as a subsection of Ambelania, a genus
whichheacceptedinthewidesense,andnearlyequivalent tothetribe Ambelanieae as here interpreted. PICHON (1948) considered Neocouma as a monotypic
genuseasily separable from Ambelania, Molongum, and Rhigospira.
In the present treatment, two species of Neocouma are recognized, N. ternstroemiacea and N. parviflora.
E t y m o l o g y . The generic name is a modification of Couma, a member oftheapocynaceous tribe Carisseae.
D i s t r i b u t i o n . Northwest Amazonia in Venezuela (Amazonas, Bolivar),
Colombia (Amazonas,Vaupés),and Brazil(Amazonas, Para);Map 6.
K E Y TO T H E S P E C I E S O F

NEOCOUMA

1. Petioleshort, 1-2 mmlong;leafbladeselliptictoovatewithretuseto rounded
apex; bracteoles small, 2.5-3 x 3-4 mm; corolla tube short, 3.5-4 mm
long; fruits stipitate (Amazonas & Bolivar, Venezuela and Serra Aracâ,Amazonas, Brazil)
6-1. N. parviflora
Petiole elongate, 6-30 mm long; leaf blades elliptic with rounded to obtuse
apex; bracteoles large, 12-14 x 5.5-7 mm; corolla tube 10-12 mm long;
fruits not stipitate (Northwestern and Western Amazonia, and Western
Para, Brazil)
6-2.N.ternstroemiacea
Agric univ. WageningenPapers87-1 (1987)

87

MAP 6 Neocouma

6-1.Neocoumaparviflora(Markgraf) Zarucchi,comb.nov.
Basionym:AmbelaniaparvifloraMarkgraf,Ernstia 19:4-5. 1983 [30Jul].
Type:Venezuela:Amazonas: Depto. RioNegro,enla vertiente oriental delMacizo Aracamuni, Lat. 01° 32' N., Long. 65°48'W., alt ca 750m, 10Feb
1981 (fl fr), O.Huber &Medina 5928 (Z nv, holotype; isotypes: NY, VEN
165823).
Smalltree5mtallwithleaf-bearingbranches3-7mmindiameter.Leafblades
elliptic to obovate, 9-14.5 x 5-8.5 cm, glabrous, coriaceous, with 9-13 pairs
oflateral veins;basesobtuseorrounded, decurrent ontheshortpetiole;apices
retuse or rounded; margins entire, revolute; midrib slightly impressed above,
prominent below;lateral veinsimpressed above, slightly raised below;tertiaryveins conspicuous above, slightly raised below; light brown spots on abaxial
Agricuniv. WageningenPapers87-1 (1987)

FIG. 7. Neocouma parviflora (Markgraf) Zarucchi: 1. flowering branch, \ x ; 2. opened flower,
3 x ;3.fruits, J x . ( l - 3 . 0 . Huber &Medina 5928).

Agricuniv. WageningenPapers87-1 (1987)89

surface. Petioles1-2 mm long, winged, shallowly canaliculate. Inflorescences
terminal,cymose,ca25-30-flowered;peduncles 15-36mmlong;bractspossibly
caducous[notseen];pedicels8-15mmlong;bracteolesdeltoidtoovate,calycine,
2.5-3 x 3-4mm,apicallyacute,locatednearmiddleofpedicels.Flowers:maturebud4-5mmlong;sepalsovate,2-2.3 x 1.5-2mm,apicallyobtusetosubacute,finely ciliate only at apex. Corolla tube 3.5-4 mmlong, sparingly pilose fromjust below insertion of stamens to orifice within; lobes deltoid, 2.8 x 2mm,
reflexed. Stamensinserted 2mmfrom baseofcorolla tube;antherslanceolate,
2.5—3mmlong,sessile,basally auriculate, spreading, lobes0.4mmlong,apicallycaudatewithacumen0.4-0.5mmlong.Ovaryovoid,ca1 mmlong,apicallytruncate;style0.3-0.4mmlong;clavunclecylindrical with a basalflange, apicallyfive-lobed,ca0.7mm long;stigmoid lobesterete, thin,ca 0.25mmlong.

MAP 22.Neocoumaparviflora (Markgraf) Zarucchi.

90

Agricuniv. WageningenPapers87-1 (1987)

Fruit(immature)globoseto22mmin diameter,largerfruitssomewhatstipitate;
pericarp crustaceous, ca 1mm thick, with 8-10 seeds embedded in a stringy
pulp;surfaceverrucose.Seedsplano-convex,elliptictoovateinoutline,11-12.5
x 6-7 x 3-4mm, surfacefinelypitted,withirregularprotuberances.
Etymology. Latinparviflora =small-flowered, inreferencetotherelatively
smallcorollasize.
Distribution. Known from Venezuela (Amazonas, Bolivar) and Brazil(SerraAracâ,Amazonas);Map22.
Specimens examined:
VENEZUELA. Bolivar: cumbre del Cerro Guaiquinima, a lo largo del afluente del Rio Carapo,
1 km rio arriba del Salto Szezerbanari, 5°44'4"N, 63°41'8"W, parte SE del cerro, alt 730-750m,23-24May 1978(fr),Steyermark etal.117386(VEN).
BRAZIL. Amazonas: Serra Aracâ, alt 1000 m, 11 Feb 1975 (fl fr), Pires 15048 (IAN 145484);
margensdeumriachodaSerraAracâ, 1Feb 1978 (flfr),Rosa & Lira2350(MG,NY).

Nolocalnamesarerecordedforthisspecies.
Neocoumaparvifloraisaspeciesmarkedlydifferent from N. ternstroemiacea.
Characters which differentiate N. parviflora include the short petiole, elliptic
to ovate leaf bladeswith retuse to rounded apices, small bracteoles, the shortcorollatube,andstipitatefruits.
The speciesisknown onlyfrom four gatherings at above 1000m elevation,
includingontheSerraAracâ,anisolatedmountain.Thesubstrate ofthisserra
isrelatedtothatofthe mainGuianahighlandregion,severalhundredkilometers
to thenorth, wheretheother collections havebeenmade. MAGUIRE (1979)included theSerraAracâasoneofseveralisolatedmountainsinnorthern Amazonia worthy of intense floristic study. Additional populations of Neocouma
parvifloraareexpectedfrom neighboringisolatedmountains.
6-2.Neocoumaternstroemiacea(Muell.-Arg.)Pierre,Bull.Mens.Soc.Linn.Paris, sér.2,5:33-34. 1898;Pichon, Mém.Mus.Natl. Hist.Nat. 24: 169-170,pi.
IV-19,29,40.1948.
Basionym: Tabernaemontana ternstroemiacea Muell.-Arg.in Martius, Flora
Brasiliensis6(1): 88. 1860.
Type:Venezuela: Amazonas: 'Prope San Carlos ad Rio Negro,' Aug 1853(fl), Spruce 3035(BR, lectotype [herechosen];isotypes:B,lost [Fphoto4380;
F, GH, MO, US,VEN],BM,C, CGE, E, F 998981,G, GH, GOET, K, LD,
NY,OXF,P). Homotypicsynonyms:Rhigospiraternstroemiacea(Muell.-Arg.)
Miers,Apocyn.S. Am. 71.1878.Ambelaniaternstroemiacea(Muell.-Arg.)Monachino,Lloydia 8(2): 123-124.1945.

Agricuniv. WageningenPapers87-1 (1987)91

Medium-sized treeupto25mtallwithleaf-bearing branches6—12(—15)mm
in diameter.Leafbladeselliptic,15-30 x 8-16cm, glabrous,coriaceoustothickcoriaceous,with(7-)10-13pairsoflateralveins;basesobtusetorounded,decurrenton petiole;apicesroundedto obtuse,sometimesshortlymucronate;margins
entire, revolute; midrib somewhat prominent above, becoming slightly impressednearbaseofblade,prominent below;lateralveinsslightlyraisedaboveand below;tertiaryveinsconspicuous on both surfaces; numerouslight brownspots scattered on abaxial surface. Petioles (6—)11—24(—30)mm long, winged,
canaliculate. Inflorescences terminal, cymose, congested, (6-)10-23-flowered;peduncles 11—30(—42)mmlong;bractsovatetoelliptic,calycine,11-14 x 5.5-7mm;pedicels4-7mmlong;bracteolesovatetoelliptic,calycine, 12-14 x 5.5-7mm, locatedabovemiddleofpedicel.Flowerslarge,maturebud35-40mm long;
sepalsovatetoelliptic,12-14 x 6-7mm, apexroundedtoacute,eciliate.Corolla
whitewithtube 10-12mm,pilosebelowinsertion ofstamenswithin;lobesoblong-elliptic,20-28 x 7-12mm,reflexed. Stamensinsertedca4mmfrom baseofcorollatube;antherslanceolate,4-5mmlong,sessile,basallyauriculatewith
lobes ca 0.5 mm long, apically caudate with acumen ca 0.6 mm long.Ovary
ovoid,ca2mmlong,apicallyrounded;styleupto4mmlong;clavuncle0.5-0.7
mm long; stigmoid lobes terete, thin, ca 0.6-0.7 mm long. Fruitellipsoid, toca3.5cmlongand 2cmindiameter;pericarpcrustaceous,granularintexture,ca 1mmthick,with(8-)15-20seedsembedded inastringypulp;surface verrucose.Seeds unequally compressed laterally,depressed trullate incross-section,
ovate to elliptic in outline, 14.5-16 x 7-8 x 4-5 mm; surface finely pitted,
withirregularprotuberances.
Etymology. Theepithetternstroemiacearefers tothesimilarityinappearance
ofthecalyxtothatfoundinternstroemias(Theaceae).
Distribution. KnownfromnorthwesternAmazoniain Brazil,Colombia,and
Venezuela;alsocollectedinwesternPara,Brazil;Map 23.
Specimens examined:
COLOMBIA. Amazonas: Trapecio amazónico, interior regions of trapecio between Amazon and
Putumayo watersheds, altabove 100m, Sep 1946(st),Schuhes 8111(NY, UC).Amazonas-Vaupés:
Rio Apaporis, Raudal de Jirijirimo, 7 Aug 1951 (fl, yng fr), Schuttes & Cabrera 13462 (COL, F,
GH,P[asjn],WIS).
VENEZUELA. Amazonas: San Carlos de Rio Negro, in forest along airstrip, alt 115 m, 19 Sep
1975(fl), Berry 1475(VEN); MAB - near San Carlos de Rio Negro, Sep 1975 (fl fr), Brünig VEN
109 (VEN); San Carlos de Rio Negro, 4.3 km NNE on Solano road, ca 1° 56' N, 67° 03' W, alt
119m, 21Dec 1978(fl), Clark & Maquirino 6903(MO,NY); 9.2 km NNE of San Carlos on Solanoroad, near conuco ofJuan Moreno, alt 119m, 1Jan 1979(fl), Clark 6921 (NY);Depto. Rio Negro,.5 to 1.5 km SE of Cerro de La Neblina Base Camp on the Rio Mawarinuma, 0° 50' N, 66° 10'
W, alt 140 m, 21 Feb 1984 (st), Liesner 16154(MO); Rio Guainia, Cafio San Miguel 2 km above
Limoncito, sabanita 1km W of La Ceiba, alt 120-140 m, 14 Oct 1957 (fl), Maguire et al. 41893(NY); carretera San Carlos-Solano, 2-6 km SW de Solano, alt 100 m, 25 Apr 1974 (fl yng fr),
Morillo et al. 3944 (VEN); en camino de San Carlos de Rio Negro hacia Solano, alt 120 m, 27
Oct 1970(fl), Rutkis 221 (VEN); Cerro dela Neblina, vecinidades delcampamento base a lo largodel Rio Mawarinuma afluente del Rio Baria, 0° 50' N, 66° 10' W, ait 110-130 m, 8-10 Jan 1984
(fl buds),Steyermark & Luteyn 129741(MO,VEN).
92

Agricuniv. WageningenPapers87-1 (1987)

MAP 23.Neocouma ternstroemiacia(Mueller-Argoviensis) Pierre.
BRAZIL. Amazonas: Rio Negro, Camanâos, 30 Nov 1929 (fl), Ducke RB 22417 (K, P, RB, S,
U, US); Rio Negro, Cucuhy, 22 Sep 1935 (fl), Ducke RB 35156 (RB), 220 (MAD, NY); basin of
Rio Negro, Mun. Sâo Gabriel, Ca-te-Espera, 14Nov 1941 (old fl, fr), Frees 368 (F, G, MO, NY,
S), 14Nov 1941 (fl), Fróes369 (F, G, NY, S); Mun. Sào Gabriel, Yuco, Rio Xié, 5Dec 1941 (fl),Frees 416 (F, G, GH, NY, S, WIS), 6 Dec 1941 (st), Fróes 425 (G, NY), 9 Dec 1941 (fl), Fróes
431 (G, NY, S, US); Mun. Sâo Gabriel, Santa Ana on Rio Içana, 18Dec 1941 (fl), Fróes 443 (G,NY), 18 Dec 1941 (yng fr), Fróes 444 (NY); Mun. Sâo Gabriel, Acara, Ig. Cubaté (trib. of Rio
Içana), 2 Jan 1942 (fl fr), Fróes 471 (G, NY, S, US), 2Jan 1942 (fl, yng fr), Fróes475 (F, G, GH,
NY, S, US); Mun. Sào Gabriel, Vaupés, Corocoró, 12 Jan 1942 (fr), Fróes 499 (G, NY, WIS),
12Jan 1942(yng fr), Fróes 502 (G, NY, SP), 12Jan 1942 (fr), Fróes 503 (G, NY, US); Mun. Sâo
Gabriel, Macubeta on Rio Marié, 26 Jan 1942 (fl buds), Fróes 539 (F, G, NY, S, WIS), 26 Jan
1942(st), Fróes540 (G, NY, US, WIS), 541 (NY); Mun. Sâo Gabriel, BelaVista, Basin of VaupésRiver, 9 May 1942 (fl), Fróes882 (G, NY, US); Rio Caiary, matas adjacentes â Serra de Tunuhy,
13Nov 1945 (fl), Fróes 21387 (F, IAN, K, NY, US); Rio Negro, Preto, Matupiry, 12 Nov 1947(fl), Fróes22834(IAN,NY, RB,SP,U,US);Rio Solimôes,Ndo IgarapéBelém,next[to]Colombia
border,21 Dec 1948(fr),Fróes23793(IAN,MO),21Dec 1948(oldfl),Fróes23794B(lXiï); margem
esquerda do Rio Negro, Tapuruquara-mirim, Rio Marié, 16Oct 1978 (fl), Nascimento 648 (NY);Agricuniv. WageningenPapers87-1 (1987)93

alto RioNegro, Pariuré, 7Jun 1962 (fr),Pires &Silva 7940 (IAN, UB). Para: Missâo Cururù,11 May 1977 (fr), Rosa &Santos 1901(MG, NY).

L o c a l n a m e s . VENEZUELA.Pendare amargo {Clark &Maquirino 6903), Yayucuri (Rutkis 221). BRAZIL.G o g o deG u a r i b a (Fróes 368,369,
416,425,431,471,
475, 499,502,503),Parva-papo (Fróes 539, 540,541),Sorva d acatinga (Ducke
RB 35156,220), Sorva p a p o (Fróes 882), Uarina G o g o (Fróes 443,444).
Neocouma ternstroemiacea was first described byMUELLER(1860a)asa questionable species in Tabernaemontana, based u p o n a collection by RICHARDSPRUCE made in 1853 near SanCarlos o ntheupper R i oNegro. MIERS (1878)
transferred thespecies toRhigospira, including itwith elements presently attributable to Neocouma, Spongiosperma, Rhigospira, and Macoubea. PIERRE(1898)
created the genus Neocouma to accommodate N. ternstroemiacea as itsonly species. The second collection ofthis species was not made until 1929, when A D O L PHO D U C K E collected it at C a m a n a ó s on the u p p e r Rio N e g r o . M A R K G R A F(1932b) referred tothis collection inhis discussion of TV. duckei, which is treatedhere under Mucoa duckei.
The majority of the specimens of Neocouma ternstroemiacea available
for study were m a d e by R I C A R D O DELEMOS FRÓES in the course of his w o r k

in the 1940so n Amazonian chicle-producing tree species.In recent years, collections have been made in awide variety of localities inwestern Amazonia, including theRio Apaporis at Raudal Jirijirimo a n dthe Trapecio amazónico of Colombia. One recent collection, made in western Para o ntheRioCururu, isan
unexpected disjunction which reflects theclose similarity between this regionof theR i oTapajós/Serra d a Cachimbo a n dthe catinga regions of the upper
Rio Negro.
Excluded Species of Neocouma
Neocouma duckei Markgraf, Notizbl. Bot. Gart. Berlin-Dahlem 11:337.1932= Mucoa duckei (Markgraf) Zarucchi.

BIBLIOGRAPHY
AUBLET,J.B.C.F.1775.Histoire des plantes de la Guiane françoise -Paris.
AZAMBUJA, D.de 1947. Contribuiçào aoConhecimento das Apocynaceae Encontradas no Brasil.
Arq. Serv. Florest. 3:9-112.
BÂILLON,H. 1891 [1889]. Apocynacées. HistoiredesPlantes 10:146-220-Hachette, Paris.
BARTLING, F. G. 1830. Ordines naturales plantarum eorumque characteres et afïïnitates adjecta
generum enumeratione- Göttingen.
BENTHAM, G. 1841. Contributions toward a Flora of South America-Enumeration ofPlants collected by Mr. SchomburgkinBritish Guiana (cont.). J. Bot. (Hooker) 3:212-250.
BENTHAM,G. 1876.Apocynaceae. In: Bentham, G. and J. D. Hooker, Genera Plantarum 2:681-728- Reeve &Co., London.
BOITEAU, P.&C. SASTRE 1975. Surl'arille des Macoubea etlaclassification delasous-famille desTabernaemontanoïdées(Apocynacées). Adansonia, sér. 2,15(2):239-250.
94

Agricuniv. Wageningen Papers 87-1 (1987)

BoiTEAU,P., L. ALLORGE &C. SASTRE 1978.Morphologie florale des Apocynaceae: II. Caractères
distinctifs entre Ambelanieae (Plumerioideae) et Macoubeeae (Tabernaemontanoideae). Adansonia,sér. 2,18(2):267-277.
CRONQUIST, A. 1968. The Evolution and Classification of Flowering Plants - Houghton Mifflin,
Boston.
D E CANDOLLE, A. P. 1844. Apocynaceae. In: Prodromus systematis naturalis regni vegetabilis
8:317-489- Fortin, Masson &Soc, Paris.
DESCOURTILZ,M.E. 1829.Florepittoresque etmedicinaledesAntillesouTraitédesplantes usuelles
descoloniesfrançaises, anglaises,espagnoles etportugaises7:123-126,pi. 480.
DIETRICH,D. 1839.SynopsisPlantarum seuEnumeratio Systematica- Vimariae.
DON, G. 1837. A general history of the Dichlamydeous plants. Vol. 4. Corolliflorae - Rivington
etal., London.
DUCKE, A. 1922.Plantesnouvelles oupeuconnuesdelarégionAmazonienne (IlePartie). Apocynaceae.Arch.Jard. Bot. Rio deJaneiro3:239-248,Errata etCorrigenda, est. 17-18.
DUCKE, A. 1938. A flora doCuricuriari,afluente do RioNegro,observada emviagenscoma Comissâo Demarcadora das Fronteiras do setor oeste. Anais Reuniâo Sul-Amer. Bot. l(3):389-398,
pi. 2.
DUCKE, A. 1944.Flora do rio Urubu. Bol.Minist. Agric.(RiodeJaneiro)6:31^13.
ENDLICHER, S.L. 1838.Apocynaceae. In:GeneraPlantarum 577-586- Wien.
GLAZIOU,A. F. M. 1910.Listesdesplantes du Brésilcentral recueilliesen 1865-1895. Apocynacées.
Bull.Soc.Bot.France, sér.4(mém. 3e),10:448-459.
GMELIN,J. F. 1791. Linnaeus, Systema Naturae,ed. 13. Vol.2(1).
HALLE, F. 1978.Architectural variation at the specific levelin tropical trees. In: Tomlinson, P. B.
& M.H. Zimmermann (eds.),TropicalTreesasLiving Systems,209-221- Cambridge Univ.Press.
HALLE,F.,R.A.A.OLDEMAN&P. B. TOMLINSON 1978. TropicalTreesand Forests:An Architectural
Analysis- Springer-Verlag, Berlin.
HICKEY, L. J. 1979.A revisedclassification ofthe architecture ofdicotyledonous leaves, pp. 25-39.
In: Metcalfe, C. R. & L. Chalk, Anatomy of the Dicotyledons. Second edition. Vol. 1- Clarendon
Press, Oxford.
HOLMGREN, P.K., W. KEUKEN &E. K. SCHOFIELD 1981.Index Herbariorum. Part 1.The Herbaria
oftheWorld, ed. 7.Regnum Veg. 106 - Bohn, Scheltema &Holkema, Utrecht/Antwerpen.
HUBER, J. 1902.Materiaes para a Flora Amazonica V.Plantas vasculares colligidas ou observadas
na regiâo dos Furos de Breves em 1900 e 1901. Apocynaceas. Bol. Mus. Paraense Hist. Nat.
3:443^145.
HUMBOLDT, F. H.A. VON,A.J. BONPLAND&C. S.KUNTH 1819.Nova Genera etSpecies Plantarum
3:282[folio],361[quarto],/»/. 283.
HUMBOLDT,F.H. A. VON,A.J. BONPLAND&C.S. KUNTH 1825. Nova GeneraetSpecies Plantarum
7:162[folio],209[quarto].
JUSSŒU,A.-L. DE 1789.Genera Plantarum 143-151,257-Paris.
KUNTH, C. S. 1825[1826].SynopsisPlantarum 4:229-Levrault,Paris.
LAMARCK, J. B. A. P. 1791.Tableau encyclopédique et méthodique-botanique 1:125-126,pi. 169- Paris.
LAWRENCE,G. H. M. 1965. Taxonomy ofVascular Plants- Macmillan Co.,New York.
LECOINTE,P. 1934.AAmazonia Brasileira III.ArvoresePlantas Uteis- Livraria Classica, Belém.
LEMÉE,A. 1953. FloredelaGuyane française. Vol. 3- Lechevalier, Paris.
MACBRIDE, J. F. 1959. Apocynaceae. Flora of Peru. Publ. Field Mus. Nat. Hist., Bot. Ser. 13(5/l):363^155.
MCVAUGH, R. 1955. Theamerican collectionsofHumboldt and Bonpland described inthe Systema
Vegetabilium ofRoemer and Schuhes.Taxon 4(4): 78-86.
MAGUIRE, B. 1979. Guayana, Region of the Roraima Sandstone Formation. In: Larsen, K. & L.
B. Holm-Nielsen (eds.),Tropical Botany,223-238-Academic Press, London.
MARKGRAF, F. 1932a.Apocynaceae.In:Pulle,Flora of Surinam 4(l):l-65.
MARKGRAF, F. 1932b.Neue Apocynaceen aus Südamerika IV.Notizbl. Bot. Gart. Berlin-Dahlem
11:337-340.

Agric univ. WageningenPapers87-1 (1987)

95

MARKGRAF, F. 1935. Neue Apocynaceen aus Südamerika V. Notizbl. Bot. Gart. Berlin-Dahlem
12:295-301.
MARKGRAF, F. 1941. Die Gattung Lacmellea H. Karsten. Notizbl. Bot. Gart. Berlin-Dahlem
15:615-629.
MARKGRAF, F. 1971[1972].NewApocynaceaeandAsclepidaceaefrom Venezuela.Acta Bot.Venez.
6:65-75.
MARKGRAF, F. 1975.NovedadesVenezolanas enlasApocynaceae.Acta Bot.Venez.10:247-250.
MARKGRAF, F. 1983. TresnuevasespeciasdeApocynaceaedelTerritorio Federal Amazonas,Venezuela.Ernstia 19:1-5.
MEISNER, C. F. 1836-1843. Apocynaceae in Plantarum Vascularium Genera 262-265 [pars prior],
170-173[parsaltera].
MERRILL, E. D. 1950. Unlisted technical plant names in the published works of L. Oken (1841)
and J. S.Presl(1846).J.Arnold Arbor. 31:264-287.
MIERS, J. 1878.On the Apocynaceae of South America, with some Preliminary Remarks on the
Whole Family- Williams &Norgate, London.
MONACHINO, J. 1945. A Revision of Ambelania, Inclusive of Neocouma (Apocynaceae). Lloydia
8(2):109-130.
MONACHINO,J. 1946.A RésuméoftheAmerican Carisseae(Apocynaceae). Lloydia 9(4):293-309.
MONACHINO, J. 1958.Apocynaceae. In: Maguire, B.,J. J. Wurdack & Collaborators, The Botany
oftheGuayana Highland. Part III.Mem.NewYork Bot.Gard. 10(1):117-138,/îg.s. 14-16.
MONACHINO, J. 1961. Apocynaceae. In: Maguire, B.,J. J. Wurdack & Collaborators, The Botany
oftheGuayana Highland. Part IV(2).Mem.NewYork Bot.Gard. 10(4):59-65,./fes. 43-44.
MUELLER, J. 1860a.Apocynaceae.In:Martius, Flora Brasiliensis6(l):l-196,/>/. 1-54.
MUELLER, J. 1860b.Speciesnovae nonnullae americanae exOrdine Apocynearum et observationes
quaedam in species generis Echitis Auctorum earumque distributio in genera emendata et nova.
Linnaea30:387^454.
NECKER,J. DE 1790.Elementa Botanica. Neuwied. 2:54.
OKEN, L. 1841. Asclepiaden und Apocynen. Allgemeine Naturgeschichte für alle Stände
3(2):1025-1051.
PICHON, M. 1948. Classification des Apocynacées I. Carissées et Ambelaniées. Mém. Mus. Natl.
Hist.Nat. 24(3):lll-181,p/. I-IV.
PIERRE, L. 1898. Observations sur quelques Landolphiées. Bull. Mens. Soc. Linn. Paris, sér. 2,
1(5):33^K).
PRANCE, G. T. 1971a. An index of plant collectors in Brazilian Amazonia. Acta Amazonica 1(1):25-65.
PRANCE, G. T. 1971b.New species ofphanerogams from Amazonia. Brittonia 23(4):438-445,/ig.s.
l^f.
PRESL,J. S. 1846.Tojesïowite. Apocyneae.Wseobecny Rostlinopis 2:1051-1069.
PULLE, A. 1909.Apocynaceae. In: Neue Beiträge zur Flora Surinams II. Recueil Trav. Bot. Néerl.
6:285-287.
RECORD, S.J. &R. W. HESS 1943. Timbers ofthe NewWorld-Yale University Press,New Haven.
REES,A. 1819.TheCyclopaedia.Vol. 38 - London.
ROEMER,J. J. &J.A. SCHULTES 1819.Systema Vegetabilium 4:545and 5:189.
SCHOMBURGK, [M.] R. 1848. Reisen in Britisch-Guiana in den Jahren 1840-1844. Vol. 3: Versuch
einer Fauna und Flora von Britisch-Guiana- Leipzig.
SCHREBER,D.J.C.D. 1789.Carolia Linné,GeneraPlantarum, ed.8- Frankfurt.
SCHULTES, R. E. 1950. Plantae Austro-Americanae VI. Plantarum principaliter vallis Amazoniae
novarum velminuscognitarum notaediversae.Bot. Mus.Leafl. 14(5): 109-136,/)/. XXVII-XXXVI.
SCHULTES, R. E. 1951.Plantae Austro-Americanae VII. De festo seculari Ricardi Sprucei America
australi adventu commemoratio atque de plantis principaliter vallis amazonicis diversae observationes.Bot.Mus.Leafl. 15(2):29-78,/>A XV-XX.
SCHULTES, R. E. 1966. Lacticiferous plants of the Karaparanâ-Igaraparanâ region of Colombia.
Acta Bot.Néerl. 15:178-189.

96

Agricuniv. WageningenPapers 87-1 (1987)

SCHULTES,R.E. 1970.DePlantisToxicariis e Mundo NovoTropicaleCommentationes VII.Several
ethnotoxicological notesfrom theColombian Amazon. Bot.Mus.Leafl. 22(10):345-352,/>/. LXXXLXXXIII.
SCHULTES, R. E. 1979.De Plantis Toxicariis e Mundo Novo Tropicale Commentationes XIX. Biodynamicapocynaceous plantsofthenorthwest Amazon.J. Ethnopharmacol. \:\(>5-\92,figs. 1-13.
SCHUMANN, K. 1895.Apocynaceae. In:Engler,A.&K. Prantl (eds.),Dienatürlichen Pflanzenfamilien4(2):109-189.Wilhelm Engelmann- Leipzig.
SCOPOLI,J.A. 1777.Introductio ad historiam naturalem - Wien.
STAFLEU,F.A.&R.S.COWAN. 1979. TaxonomieLiterature(ed.2). Vol.2: H-Le[Regnum Vegetable98] - Bohn, Scheltema&Holkema, Utrecht.
STEUDEL,E.G. 1831.Nomenciator Botanicus- Stuttgart.
WAGENITZ, G. 1964.Gentianales. 405-424. In:Melchior, H. Engler's Syllabus der Pflanzenfamilien- Gebruder Borntraeger, Berlin.
WEBBER,I. E. 1945. Thewood oîAmbelania taxa Muell.Arg. Lilloa 11:49-53,pl. I-II.
WILLDENOW,C. L. 1797.Carolia Linnéspeciesplantarum ed.Willdenow, Vol. 1- Berlin.
WOODSON, R. E. 1935. Observations on the inflorescence of Apocynaceae (with special reference
to theAmerican genera ofEchitoideae).Ann. Missouri Bot.Gard. 22(1):1-48,/»/. 1-3.
WuRDACK, J. J. 1970. Erroneous data in Glaziou collections of Melastomataceae. Taxon
19(6):911-913.

ACKNOWLEDGEMENTS
The author also wishes to acknowledge with gratitude the assistance of thefollowing persons and organizations contributing totheresearch and thereviewof this work. Primary thanks go to Dr. RICHARD EVANS SCHULTES of Harvard
University,for hisvaluabledirectionand for hiswillingnesstosharehisremarkableknowledge of theAmazonian flora. Principal funding for my field researchin South America was generously provided by the Atkins, Anderson, and Fernald Funds of Harvard University; Sigma Xi, The Scientific Research Societyof North America; and the National Science Foundation (Projeto Flora Amazonica;G. T. Prance&T. C.Plowman, Co-Principal Investigators). Additional
workonthisrevisionwascarried outasaNATOPostdoctoral Fellowin London
at the British Museum (Natural History) and Royal Botanic Gardens, Kew;
at theNational Herbarium in Washington, D.C. asa Smithsonian Fellow; andat theMissouri Botanical Garden wheretheauthor isan Assistant Curator.
I wish to thank also the directors and curators of themany botanical institutions whichmade facilities and specimens available to me for this study (*indicates those herbaria visited personally by the author): A*, AWH*, BM*, BP,
BR*, C*, CGE*, COL*, DS, E* (incl. E-GL*), ECON*, F*, FI-W*, G* (incl.
G-DC*), GH*, GOET, HBG*, IAN*, INPA*, K*, L*, LA, LD, LE, LINNSmith*,LISU, M*, MAD, MEDEL*, MG*, MICH, MO*,MPU, MY*, NA*,
NY*, OS,OXF*, P* (incl. P-Humboldt & Bonpland* and P-Rousseau*), RB,
S*, SP, TEX, U*, UB, UC, US*, USF, VEN*, W*, WAG*, WIS, and Z*. In
particular, I wish to thank the following persons: Dr. E. FORERO (COL); Mr.
P. S. GREEN and Dr. G. LI. LUCAS (K); Mr. J. F. M. CANNON (BM); Drs. R.
S. COWAN, J. CUATRECASAS, D. H. NICOLSON, and J. J. WURDACK (US); Dr.
F. MARKGRAF (Z);Dr. S. NILSSON (S); and Dr. A.J. M. LEEUWENBERG (WAG).
Agricuniv. WageningenPapers87-1 (1987)

97

Special thanks alsogoto Ms. KAREN KRAGER forherexecution ofthe line
drawings which appear in this work. Thebase mapused to plot genericand
species distributions isan adaptation of theFlora Neotropica base map no.
1 producedattheDepartmentofSystematicBotany,TheUniversityofUtrecht.

98

Agricuniv. WageningenPapers87-1 (1987)

INDEX OF LOCAL NAMES
A-BI-ta-ki(Kubeo language)
Açucena d'agua
Akusiwalapulu (Wayampi language)
Amaparana
Ambalali (Galibi language)
Ambelani(Galibi language)
Angelica do Igapó
Angelica-igapó
Aratani-yek (Arekuna language)
Auca-sanango
Bat batti
Batibati
Boia
Boya
Caimo morado
Cuchara huayo
Dü-ko-gay (Huitoto language)
Gogode Guariba
Goiaba d'anta
Graine biche(Creole language)
Guacamari (Kuripako language)
Guayaba de monte
Guayabilla (de monte)
Hia-WAHO-ka-ki (Kubeo language)
Jacquitaque
Jasmin d'agua
Juansoco
Juansoco de Danta
Machieraro
Makoriro
Makurero (Arawak language)
Makuriro
Makurriro
Mapa
Masho micunaMi-mi-WAHO-ka-ki(Kubeo language)
MolongóMolongódo Igapó
Molongó semespinho
Naranja brava
Palo bâlsamo
Palo deBalso
Palode boya
Palodeboya bianco
Palodeleche
Papaya biche(Creole language)
Paraveris (Galibi language)
Parva-papo
Agricuniv. WageningenPapers87-1 (1987)

Molongum lucidum
Spongiosperma grandiflorum
Ambelania acida
Mucoa duckei
Ambelania acida
Ambelania acida
Spongiosperma grandiflorum
Spongiosperma grandiflorum
Rhigospira quadrangularis
Ambelania duckei
Ambelania acida
Ambelania acida
Molongum zschokkeiforme
Molongum laxum
Rhigospira quadrangularis
Ambelania occidentalis
Mucoa duckei
Neocouma ternstroemiacea
Rhigospira quadrangularis
Ambelania acida
Molongum laxum
Rhigospira quadrangularis
Rhigospira quadrangularis
Spongiosperma macrophyllum
Mucoa duckei
Spongiosperma longilobum
Mucoa duckei
Rhigospira quadrangularis
Ambelania acida
Ambelania acida
Ambelania acida
Ambelania acida
Ambelania acida
Ambelania acida
Ambelania occidentalis
Molongum lucidum,
Rhigospira quadrangularis
Molongum laxum,
M. zschokkeiforme,
Spongiosperma grandiflorum,
S. longilobum, S. riparium
Molongum zschokkeiforme
Ambelania acida
Spongiosperma cataractarum
Spongiosperma macrophyllum
Molongum laxum
Molongum laxum
Molongum laxum
Mucoa duckei,
Rhigospira quadrangularis
Ambelania acida
Ambelania acida
Neocouma ternstroemiacea

99

Parva-papo doTerra firma
Paudecolher
Pendare amargo
PepinoPepino bravo
Pepino domato [Pipino,and other spelling variants]

Pepino doce
Pepino verde
Pepino verdeda mata
Pipim
PipinoQuienbiendent (Creole language)
Sorvada catinga
Sorvade Cutia
Sorvarana
Sorva papo
Tajarana
Taki-taki
Uarina Gogo
Volla
Wakorira
Wild Cucumber
Wy-gaw-nö-mee-kö (Makuna language)
Yahuar caspi
Yahuar huayo
Yahuar huayo Colorado
Yapukait (Huambisa language)
Yayucuri

Mucoa duckei
Ambelania acida
Neocouma ternstroemiacea
Ambelania acida,A. duckei,
Spongiosperma grandiflorum
Ambelania acida,A. duckei
Ambelania acida,A. duckei,
Spongiosperma grandiflorum
Ambelania acida
Ambelania acida,A. duckei
Ambelania duckei
Ambelania acida
Ambelania acida
Ambelania acida
Neocouma ternstroemiacea
Mucoa duckei
Mucoa duckei
Neocouma ternstroemiacea
Spongiosperma grandiflorum
Ambelania acida
Neocouma ternstroemiacea
Molongum laxum
Ambelania acida
Ambelania acida
Spongiosperma macrophyllum
Mucoa duckei,
Rhigospiraquadrangularis
Mucoa duckei
Rhigospira quadrangularis
Rhigospira quadrangularis
Rhigospira quadrangularis
Neocouma ternstroemiacea

NUMERICAL LIST OF SPECIESOF THE AMBELANIEAE (APOCYNACEAE)
Ambelania Aublet
1-1. Ambelania acida Aublet
1-2. Ambelania duckeiMarkgraf
1-3. Ambelania occidentals Zarucchi
Mucoa Zarucchi
2-1. Mucoa duckei (Markgraf) Zarucchi
2-2. Mucoapantchenkoana (Markgraf) Zarucchi

100

Agricuniv. WageningenPapers87-1 (1987)

Spongiosperma Zarucchi
3-1. Spongiosperma cataractarum Zarucchi
3-2. Spongiospermagrandißorum(Hüber) Zarucchi
3-3. Spongiosperma longilobum(Markgraf) Zarucchi
3-4. Spongiosperma macrophyllum (Mueller-Argoviensis) Zarucchi
3-5. Spongiosperma oleifolium (Monachino) Zarucchi
3-6. Spongiosperma riparium(Monachino) Zarucchi
Molongum Pichon
4-1. Molongum laxum (Bentham) Pichon
4-2. Molongum lucidum(Humboldt, Bonpland &Kunth) Zarucchi
4-3. Molongum zschokkeiforme (Markgraf) Pichon
Rhigospira Miers
5-1. Rhigospira quadrangularis(Mueller-Argoviensis) Miers
Neocouma Pierre
6-1. Neocoumaparviflora (Markgraf) Zarucchi
6-2. Neocouma ternstroemiacea (Mueller-Argoviensis) Pierre

LIST OF EXSICCATAE
Collection numbers whichareprinted initalicsindicate typecollections.
Abadie, B., 19T(2-1),25T(5-1).
Agostini.G., 227(3-1).
Albuquerque, B.P.deetal.,345(1-1),657(3-3),917(2-1),INPA 99868(3-3).
Alencar, L.etal.,84(2-1),92(1-1),594(4-1), 632(3^1).
Allen,P.H., 3090(3-4), 3271(4-2).
Aluisio,J., 93(1-2).
AmaraLI.L.etal., 121,659(1-1).
Archer, W.A., 7625(3-2),7766(1-1),7958(3-2),8009,8106,8234(1-1).
Asuncion L.,J., 162(5-1).
Aublet, J. B.C. F.,s.n.(1-1).
Austin, D. F.etal.,4112(3-2).
BAFOG, 25M,5169,6137(1-1).
Bamps,P.,5324(l-l).
Bastos,M.etal., 134(1-1).
Bastos,N. C.etal., 112(3-2).
BenitezdeRojas,C.E.,2555(3-1).
Berlin,B., 3519(5-1).
Bernardi, L.,920,2738(5-1).
Berry,P.E., 1475(6-2).
Black,G. A.,49-8192, 52-14159 (1-1),57-19873(3-2).
Blanco,C , 1192(5-1).
Brünig, E.F.,VEN 109(6-2).
Bunting,G. S.etal.,4038,4066(3-6).
BW(Boschwezen), 1556,1882,4441,4552,4660,4713,4907,5932,6295,6863(1-1).
Calderon, C. E.etal.,2941(3-3).
Campbell,D.G. etal.,P22293,P22573(1-1).
Cardona,F.,2852(3-l).
Cavalcante,P.B.etal., 359(3-2), 1447(1-1),2239/4112(3-2).
Cerda, J., s.n. (4-1).
Agricuniv. WageningenPapers87-1 (1987)

101

Cezârio, etal.,INPA 20612 (1-1).
Chagas, J., INPA 1 (1-1),INPA 801(1-2).
Cid [Ferreira],C.A.eta l , 1056(1-1).
Clark, H. L.etal.,6903,6921 (6-2).
Coêlho,D.etal.,875(l-2).
Coêlho, L.F.etal.475(2-1), INPA 3227(1-2),INPA 5157,INPA 36021 (1-1).
Croat, T. B., 18866,19042(1-3).
Cruz, J. S.dela, 3068,4423(1-1).
Cuatrecasas, J. etal.,7129(2-1),24100 (5-1).
Curran,H.M.,9(l-l).
Daly, D. C.etal.,D887 (3-2).
Damiào [Mota],C. etal.,2967 (4-1).
Davidse, G. etal., 17223,17258(3-5), 17335(4-1), 17977 (3-2).
Davis, E.W., 135(3^1).
Diaz S., C.etal.,47A,58A (5-1), 386(1-3),477(5-1), 1471(5-1).
Drouet, F., 1949(3-2).
Duarte, A. P.etal.,9845(3-3).
Ducke, A., MG 2640,MG 6829,MG 6959,MG 7156(1-1),MG 8002(3-2),MG 10629,MG 10899(1-1), MG 11357, MG 15519 (3-2), MG 15760, RB 11397 (1-1), RB 15820 (3-2), RB 17458(1-1), RB 22417 (6-2), RB 22418 (2-1), RB 22419 (5-1), RB 22420 (4-1), RB 22421 (4-3), RB
22422 (1-2),RB 30103(2-1), RB 30123(4-1), RB 35153(1-2), RB 35156(6-2), RB 35159 (5-1),
44, 163 (4-1), 212 (1-1), 220 (6-2), 320 (2-1), 451,451 (Ha Col.) (1-2), 471 (5-1), 798 (3-3),1254(3-2).
Duke,J.A., 11499(5-1).
Egler,W. A.etal.,478(3-2),46545 (1-1).
EliasdePaula, J., 207(3-2),216(1-1).
Encarnación, F.,26154 (5-1),26158 (2-1).
Espina Z.,J. etal.,274(4-1).
F photo [Field Museum photograph], 4380 (6-2), 26841 (4-2), 26842 (3-2), 26843 (5-1), 26844(1-1),34363 (1-1),38732 (4-2),38761 (2-1),38763 (3-4),51090(3-5),s.n. (3-2).
Farinas, M. etal.,644,686 (4-1).
FDBG (Forest Department of British Guiana), 3265[F529](1-1).
Ferreira, A. R., 243(4-1), s.n. (1-1).
Ferreira,E., 57-17 (5-1).
Fróes, R. de L. et al., 34 (1-1), 280, 282 (2-1), 368, 369, 416, 425, 431,443, 444, 471,475 (6-2),
491h (4-1), 499, 502, 503, 539, 540, 541 (6-2), 548, 825 (2-1), 882 (6-2), 1956 (1-1), 12004/5(1-2), 12153/63 (5-1), 20651 (2-1), 20751 (1-1), 20775 (3-2), 21010 (2-1), 21149 (5-1), 21387(6-2),22046(1-2),22113(1-1),22121 (2-1),22367(4-1),22595 (5-1),22692,22695(4-1), 22786(2-1), 22834 (6-2), 22853 (5-1), 23247 (2-1), 23304 (4-1), 23793, 23794B (6-2), 23949 (1-3),
24171, 24886 (1-1), 24948, 25410 (3-3), 25720 (1-1), 26196 (5-1), 26552 (2-1), 28265, 29258(4-1), 31046 (1-1), 31727, 31875, 32014, 32074, 32737 (3-2), 32901 (1-1), 33138 (3-2), 33675,
33738 (4-3),34386(1-1).
Gabriel, M., s.n. (1-1).
Garcia-Barriga, H., 15068,15075 (4-2).
Gentry, A. H. et al., 12991 (3-3), 13129 (3-2), 21215 (2-1),22359, 31719 (1-3), 35386 (5-1), 41656,
41693,41976(1-2).
Glaziou, A. F. M., 14078(1-1), 14081 (3^t).
Granville, J. J.de,B.4882(1-1).
Grenand,P.,750(l-l)Guedes, T., 246(1-1).
Harrison,J.B., 1147(1-1).
Herb. Maire, s.n. (1-1).
Herb. L.C. Richard, s.n. (1-1).
Hohenkerk, L. S.,454C(CTS 68A)(1-1).
Hostmann, F.W. R., 1317 (1-1).
102

Agricuniv. Wageningen Papers87-1 (1987)

Huashikat,V.,942(5-1).
Huber, J., MG 642,MG 1044a(1-1),MG 1725,MG 1836(3-2).
Huber, O.etal., 1796,2034,2538,2541,2995,4807(3-5),5819(4-1), 5928(6-1).
Humboldt, A.von &A. Bonpland, 990(4-2).
Idrobo.J.M., 11467(4-2).
INPA (Instituto National dePesquisas da Amazonia), 1,4015,4180,7662,8907(1-1).
Irwin, H. S.etal.,48654,54587, 55676 (1-1).
Jangoux, J.etal.,41(1-1).
Jenman,G. S.,4929 (1-1).
Killip,E.P.etal.,30007 (1-1).
Klug,G., 21,6/3(1-3).
Krukoff, B.A., 5822(4-3),8135(1-3).
Kubitzki,K., 75-81 (4-1).
Kuhlmann, J. G., RB21875 (1-1).
Kuhlmann, M. etal.,346(3-2).
Labroy, O.,s.n. (1-1).
Lanjouw, J.,416 (1-1).
Leeuwenberg, A. J. M., 11779(1-1).
Lems,K., 5253(1-1).
Leprieur, F. R., s.n. (1-1).
Liesner, R. L.etal.,9119(4-1), 16154(6-2), 19460(1-1).
Lieras,E.etal.,P16600 (1-1),P16964,P16985(5-1),P17156(1-3),P19615(5-1).
Loureiro, A.et al.,INPA 35786(1-2),INPA 37638,INPA 38014(1-2),INPA48424 (1-1).
Maas, P.J. M. etal.,P12717 (5-1).
Madison, M. T.étal., 6131 (4-1).
Mägdefrau, K., 539(3-1).
Maguire, B.etal., 30474, 30553,34504,34512(3-5), 34798,34866,35547(4-1), 35664 (4-2), 36267,
37528 (4-1),41494,41507(3-5),41893(6-2),41898(4-2),42627 (4-1).
Marinho, L. R., 33(5-1),405(2-1),424(4-1),572(2-1).
Martin, J., s.n. (1-1).
Martinelli, G. etal.,6922(3-2).
Martius, K. F.P.von, s.n., s.n. (1-1).
Mathias, M. E.etal., 5116(1-2).
Medina, E.,250,419,492 (4-1).
Mélinon, M., 14,82, s.n. (1-1).
Mello,F., 11/74(4-3),INPA 4015(1-2).
Mexia,Y.,5982a(3-2).
MG (Museu Goeldi),MG 9577,MG 9683 (3-2).
Monsalve B.,M., 594,702(5-1).
Monteiro, O.P.etal., 141(1-1), 142(5-1), 144,INPA 99007 (1-1).
Moreno, B.etal.,002(5-1).
Moretti.C, 133,481(1-1).
Mori, S.étal., 15105(1-1).
Morillo, G. etal., 3944(6-2),7580 (4-1).
Moses,T., 55(1-1).
Nascimento,O.C , 397(3-2), 590(4-1),648(6-2),774 (4-1).
Noriega M., R., s.n. (2-1),s.n. (5-1).
NY photo (n.s.),5576 Q-A).
Oberwinkler, F. etal., 15405(3-1).
Oldeman, R. A.A., 1351A, B.4025,B.4161,T-205(1-1).
Oliveira, A.,INPA 60537 (1-1).
Oliveira, E., 813(3-2),2367(1-1),2829(1-2),3248(1-1), 5186(3-2), 5501(1-1),6055(3-2).
Oliveira,R.,2736(1-1).
Persaud, A. C , 150,277(1-1).
Pinate,P.etal., 1044(4-1).
Agricuniv. WageningenPapers87-1 (1987)

103

Pinheiro,M.,15(3-2).
Pires,J.M .etal.,553,567(3-2),732(1-1),958(1-3),1269(1-1),1471,4421,4549,4612(3-2),
7408(1-1),7940(6-2),10260,10846,11683,11694,11745(3-2),13923(4-1),15048(6-1),48894(3-2),48899,50572,51643,51874(1-1),51875(3-2),52450(3-2).
Plowman,T.C.etal.,8092(3-2).
Poeppig,E.,2604(1-1).
Poiteau,P.A.,s.n.(1-1).
Prance,G.T.etal.,1622,1624,1717(1-1),3013(5-1),3162(3-3),3173(1-2),4628(1-1),5075(3-3),9532(1-1),10804,11807,12242,12267,12565(5-1),15390(2-1),17734(3-3),20734(2-1),23837(5-1),24941(1-1).
Rabelo,B.,516(1-1),585,616(3-2).
Ramirez,R.,75(1-3).
Ramos,J.etal.,344(5-1).
Ramos,M .L.S.,02(3-2).
Revilla,J.,1463,1965(1-3).
Ribamar,J.etal.,140(1-1).
Ribeiro,B.G.S.,169,997(1-1).
Riedel, L., 1368(1-1).
Rimachi,Y.,M.,3482(1-3).
Rodrigues,R.S.('R.Siqueira'),M G3367(3-2),M G3377(1-1),M G3396(3-2).
Rodrigues,W .A.etal.,1333(1-1),1361(1-2),1425,2059(1-1),2192(3-3),2402(1-1),2403(1-2),3583(5-1),4398(1-2),4402(1-1),4453(5-1),4528(1-2),4684(1-1),4904(3-3),5200,5380(1-1),5814(5-1),5903(1-2),6779(4-3),6893(1-1),7559(5-1),8931,9967(1-1),I N P A2159(1-2).
Romero-Castaneda,R.,3493,3752,3917(3^).
Rosa,N.A.etal.,538(1-3),1218,1757(1-1),1901(6-2),2350(6-1),2710(1-1),2887(3-2).
Rutkis,E.,221(6-2).
Sagot,P.,393(1-1).
Samuels,J.A.,s.n.(1-1).
Sanchez,D.etal.,601(5-1).
Sandwith,N .Y.,465,537(1-1).
Santini,4170(1-1).
Santos,M .R.,76,147(1-1).
Sastre,C.etal.,94,3931,3955,3972,4164,4192,4341,5449(1-1).
Schomburgk,RobertH.,919(4-1).
Schubert,B.G.,2206(1-1).
Schultes,R.E.etal.,3877,3877a(2-1),5357(5-1),5815(4-2),6514,6533(4-3),8111(6-2),9637B(4-1),10132(3-6), 10204 (3^1),10310(4-3),13175(3-4),13462(6-2),16968,17135,17154,
17245(3^1),18346(4-2),19290,19516(3-4),24074(1-3),26136A(1-1).
Silva,M .etal.,540,1735(1-1),2081(1-2),2438(1-1),2709,3390,3407,5078,5125(3-2).
Silva,M .F.daetal.,804(2-1).
Silva,N .T.,70(1-1),3858(4-2),5058(3-2).
Smith,A.C.,2581(1-1).
Soejarto,D.D.,582(1-3).
Spruce,R.,2445,2445bis(4-1), 2483 (3-4;seealsoundertypesof1-1), 3026 (5-1),3035 (6-2),5525(4-2),s.n.(1-1).
Stahel,G.,265,265a(1-1).
Steyermark,J.A.etal.,74748,75361(5-1),90307(3-1),102453(4-1), 103265,103268 (3-5),107227(2-2),117386(6-1),123913,124064(2-2),129741(6-2).
Stout,G.,700(2-1).
Tejera,E.etal.,9,s.n.(3-1).
Tillett,S.S.etal.,44822,45033(1-1).
Trujillo.B.,6013(3-1).
m e ,E.,5177(1-1).
Vareschi,V.etal.,6602a(3-5),6901(3-1).
104

Agricuniv. WageningenPapers87-1 (1987)

Vâsquez, R. etal.,616(1-3), 1029,2056(5-1).
Versteeg,G. M., 626(1-1).
Vieira,M. G. etal.,798(1-2).
Wachenheim, H.,0.17,202,403(1-1).
Weigelt,C , s.n. (1-1).
Weiss,L.etal.,54(5-1).
Williams,Ll.,3017(1-3),13856(4-1),14288,14454(4-2),14799,14874(4-1).
Wurdack,J.J.etal.,43021(3-5), 43279 (3-6),43749(3-4).
Zarucchi,J.L.,etal.1137,1210(4-2),1570,1711(5-1),1763(3^1),1812(5-1),1852(2-1),1903(3^1),2139(4-2),2428,2431,2432(5-1),2434,2435(3^1),2455,2480,2482,2483(4-2),2497,
2500(5-1),2504(3-4),2509(5-1),2531,2533(1-2),2534,2538(1-1),2541,2839(3-3),2879,
2902(4-3),2903(1-1),2937,2952(4-3),2973(2-1),3071,3180,3208,3217(4-3),3243(3-3).

REGISTER
Synonyms are in italics. Page number of principal entries in bold face; those of figures and photographsin italics.
AmbelaniaAublet 1-5,7,8,11,13,15,17(map),21,22,23,24,25(map), 36,42,46,
50,72,75,78-81,85,87- sect. Laxae Monachino 2,67
- sect.Neocoumae Monachino 40,86- sect.Neocoumopsis Pichon 40,42,46- sect. QuadrangidariaeMonachino 79- sect. TerminalesMarkgraf 67,79- subsect.Markgraflanae Monachino 40- subsect. TernstroemiaceaeMonachino 86- acidaAublet 1-3,9,15,16,18,19,24,26,27,29(map),32,33,36,37, 79- camporum Glaziou 2,39
- cucumerinaMiers 1,2,27,33,58- cuneataMueller-Argoviensis 1,39,50,73,75- duckei Markgraf 2,8,15,26,33,34,35(map),36,37,41,46- edulis(Roxburgh) J.Presl 39- grandiflora Huber 2,39,50,53,55,62- lanceolataMarkgraf 3,40,62- laxa (Bentham)Mueller-Argoviensis 1,39,
50,69,79- longilobaMarkgraf 3,39,56- lopeziiWoodson 2,39,53,58,62- lucida(Humboldt, Bonpland & Kunth)Markgraf 2,39,73,75,79- macrophylla Mueller-Argoviensis 1,2,27,
32,33,39,49,50,58,62,85- markgrafiana Monachino 2,39,42,46- occidentalisZarucchi,sp.nov. 26,36,
38(map)- oleifoliaMonachino 2,39,62,65- oleifoliavar. ripariaMonachino 2,39,63- pantchenkoana Markgraf 3,40,46,47(map)- parviflora Markgraf 3,40,88
Agricuniv. WageningenPapers87-1 (1987)- quadrangularisMueller-Argoviensis 1,39,
62,80,81,85- sagotiiMueller-Argoviensis 1,26
- tenuifloraMueller-Argoviensis 1,26,36- tenuifloravar. tenuiramea Mueller-Argoviensis 1,27
- ternstroemiacea(Mueller-Argoviensis) Monachino 39,46,86,91- zschokkeiformis Markgraf 39,50,77,79
Ambelanieae Pichon exBoiteau,Allorge&
Sastre 2,3,5,7,8,11,13,15,16,18,21,22,
25,40,48,87,100
AmbelaniinaePichon 2,22
BenthecaNeckei 24
Carisseae Bartling 4,7,11,13,16,22,41,87
CoumaAubl. 4,7,41,87- guianensisAubl. 4
Hancornia laxa (Bentham) A.DeCandolle 69- macrophylla Spruceex Mueller-Argoviensis 58,61,81,85
Lacmellea Karst. 2,75, 77- lucida(Humboldt, Bonpland & Kunth)
Miers 72
Macoubea Aublet 1-3,13,22,81,94- guianensisAublet 86- sprucei (Mueller-Argoviensis) Markgraf 14,
86- spruceivar.paucifolia (Spruceex MuellerArgoviensis) Monachino 86
Macoubeeae Boiteau &Sastre 3,22
Macoubeinae Pichon 2,22
MolongumPichon 2-5,11-13,15,17(map),
21-23, 50,55,66,68(map), 79,87- sect. CoilantheraPichon 2, 50,67,68- sect. OligoonPichon 2, 50,67,68,73,75,79

105

Molongum Pichonsect. TrichosiphonPichon 48,50,53, 55,68- cuneaium (Mueller-Argoviensis) Pichon 73,
75,79- grandiflorum (Huber) Lemée 53,79- laxum(Bentham)Pichon 2,5,6,8,18,20,
39,67,69,70(map), 72,75- lucidum(Humboldt, Bonpland &Kunth)
Zarucchi,comb.nov. 3-8,10,14,16,18,39,
68,69,72,74(map),75,79- macrophyllum (Mueller-Argoviensis) Pichon 48, 50,58,79- zschokkeiforme (Markgraf) Pichon 2,8,39,
69,72,75, 76,78(map), 79
MucoaZarucchi,gen.nov. 3,4,7,8,11,13,15,17(map), 18,21-23,40,41(map), 42- duckei(Markgraf) Zarucchi,comb.nov.
3-6,9,15,18,19,39,42,43,45(map),46,
94- pantchenkoana (Markgraf) Zarucchi,comb,
nov. 40,42,46
NeocoumaPierre 1-3,5,8,11,13,15,17(map), 18,21-23,45,81,86,87,94- duckei Markgraf 2,7,39,41,42,46,87,94- parviflora(Markgraf) Zarucchi,comb,
nov. 40,87,88,S9,90(map),91- ternstroemiacea (Mueller-Argoviensis)
Pierre 3,7,8,10,12,15,18,20,39,46,87,
91,93(map), 94
Neocuma O.Kuntze 86
Plumerioideae 22,40,48
Psychotria lucida Humboldt, Bonpland &Kunth 75
- lucidaWilldenowexRoemer &Schultes 7,
73,75
Rhigospira Miers 1-5,7,8,11-13,15,16,17(map), 21-23,62,79,80,81,85,87,94- paucifolia (Spruceex Mueller-Argoviensis)
Miers 85
- quadrangularis (Mueller-Argoviensis)
Miers 1-6,8,10,12,14,15,16,18,20,39,
80,81,S2,83(map),85- reticulata (A.DeCandolle) Miers 86- sinuosaMiers 86- sprucei(Mueller-Argoviensis) Miers 86- ternstroemiacea (Mueller-Argoviensis)
Miers 86,91- venulosaMiers 1,58,86
RigospiraO.Kuntze 79
Secondatia densiflora A.DeCandolle 2,39
SpongiospermaZarucchi,gen.nov. 1-4,11-13,
15,16,17(map), 18,21-23,48,49(map), 50,
55,64,68,81,94- cataractarumZarucchi,sp.nov. 50,51,52(map)106- grandiflorum (Huber)Zarucchi,comb,
nov. 16,18,39,50,53,54(map), 55,56,79- longttobum(Markgraf) Zarucchi,comb,
nov. 39,50-53,56, 57(map)- macrophyllum(Mueller-Argoviensis) Zarucchi,comb.nov. 1,2,4-6,14,16,18,19,27
33,39,49-51,53,55,56,58,59,61(map),62,
79,85,86- oleifolium (Monachino) Zarucchi,comb,
nov. 3,39,40,50, 56,62,63,64(map), 66- riparium (Monachino) Zarucchi,comb.&stat.nov.
9,18,39,51,63,65,66(map)
SterculiaL. 50
Sterculiaceae Bartling 50
Tabernaemontana L. 1,72,75,81,94- sect. Stenocephalium Mueller-Argoviensis 1,80
- laxa Bentham 1,39,67,69,72- lucidaHumboldt, Bonpland &Kunth 39,
72,75- ternstroemiacea Mueller-Argoviensis 2,39,
87,91
Tabernaemontanoideae Stapf 3,22,81
Ternstroemia MutisexL.f. 92
TheaceaeD.Don 92
Trichosiphon Schott &Endlicher 50
Willughbeiaacida(Aublet)Oken 26- edulisRoxburgh 39
WillughbejaScopoliexSchreber 23- acida(Aublet) G. F.Gmelin 26
Zschokkea Mueller-Argoviensis 77

Agricuniv. WageningenPapers87-1 (1987)