Content uploaded by Scott Zona
Author content
All content in this area was uploaded by Scott Zona on Jan 13, 2016
Content may be subject to copyright.
Selbyana
1 1:
6-21
A REVIEW OF ANIMAL-MEDIATED
SEED
DISPERSAL OF PALMS
Scorr ZoNn
Rancho Santa
Ana Botanic Garden. 1500 North College
Avenue,
Claremont. California 9l7l 1
ANonsw HENoeRsoN
' New York Botanical Garden. Bronx. New York 10458
AssrRAcr. Zoochory is a common mode of dispersal
in the Arecaceae
(Palmae),
although little is known
about how-dispersal has influenced the distributions ofmost palms. A survey ofthe literature reveals that
man-v kinds of animals leed on palm fruils and disperse palm seeds.
These animals include birds, bats,
non-flying mammals. reptiles. insects. and hsh. Many morphologicdl features of palm infructescences and
fruits (e.g.,
size. accessibilit_v,
bonl'endocarp) have an influence on the animals which exploit palms, although
the nature ofthis influence is poorl-v understood. Both obligate and opportunistic frugivores are capable
of dispersing seeds.
There is little evidence for obligate plant-animal mutualisms in palm seed dispersal
ecolog.v.
In spite ofa considerable
body ofliterature on
seed dispersal
(Guppy. 1906: Ridle,v.
1930: r.'an
der
Pijl. 1982).
the specifics
of zoochory
(animal-
mediated seed
dispersal)
in regard
to the palm
family have been largel-v ignored (Lrhl & Drans-
flcld. 1987).
Only Beccari
(1877)
addressed
palm
seed dispersal specifrcall-v:
he concluded that lew
animals eat palm fruils although the fruits appear
adapted
to seed dispersal b1'animals.
Dransfield
(198lb) has concluded that palms. in generai.
have a low dispersal abilitl. while Janzen and
Martin (1982) have considered some palms to
bc "anachronisms." moribund spccies
whose
coevolved agents of dispersal
are now extinct.
Long-distance dispersal.
a possible
factor in the
evolution of a large numbcr of island endemics
in this famil.v. is thought b,v some to be unlikel-v
in that manl palms
havc fruits too large for
biotic
long-distance dispersal (Moore & Uhl, 1973:
Dransfield,
l981bl but see Carlquist.1971).
Because seed dispersal
has been so little stud-
ied. we wish to draw altention to known cases
of animal-mcdiated seed
dispersal as well as to
thc need for further freld studies.
The likelihood
of animal-mediated
secd dispersal is high,
gir''en
the
importance of palm fruits as animal food and
thc number of animals
which foragc. hoard. and
consume thcm
(Corner.
1966: Leck. 1969: Snou'.
19811
Roosmalen, 1985). However, man.v
pub-
lished accounts of seed
dispersal inlerred from
dietar-v observations
havc bccn unspecilic or in-
cidentall ver,v felr' hcld stuclies
have addressed
palm seed dispersal specificall)'.
Palm distribu-
iions are weil-known (Moore. 1973a. 1973b:
Dransfield. 198 1b), but the rolc of zoochory in
shaping these distributions is not fulll' under-
stood.
To bring attention to these
palm-animal
interactions.
an overview is
presented
here ofthe
diverse assemblages of animals which feed on
palm fruits along
with a brief examination of the
role fruit and/or infructescence morphology mal,
pla_v
in dispcrsal and subscquent distributions.
i-Icrnoos
Data for fruit consumption and seed dispersal
were taken from personal observations and the
literature. much of it not primarill' concerned
with palm seed dispersal.
The data are
presented
in Tenle 1. The dispersal of non-native palms
is omitted, as is dispersal b,v man and abiotic
means
(e.g..
water dispersal of .\ipa). The s_vs-
tematic arrangement of palm genera
follows
Uhl
and Dransfield
(1987).
and species are arranged
alphabeticalll'within each
genus.
The palm no-
mcnclature
agrees
rvith Moorc (1963. 1973b).
Uhl and Dransfield (1987). and recent mono-
graphs (Read, 1975; Essig. 1978: Dransfield.
l98la: Moore & {Jhl, 1984: Henderson" 1987).
The animal nomenclature is less
standardized.
Where possible.
bird nomenclature
agrees
with
the American Ornithologists' Union (
1983)
checklist: otherwise. the original source
is fol-
lolvcd. The arrangement of animal names for
each palm laxon is not intended to suggest
the
animals' relative importance as dispersers. The
significance of palm fruit in an animal's diet. and
hence the animal's significance as a disperser,
could not be determined at this ler.el of inquiry.
The taronomic class of each animal is indi-
cated
within Tlerr I . In those instances in which
the- specific identitl' of the disperser(s) is not
known. only the class
has
becn
indicated. In cases
designated
u,ith a plus sign (r ). the class
indi-
I 9891 ZONA & HENDERSON: SEED
DISPERSAL OF PALMS
Trslr L Dispersal
agents of palms.
The arrangcmenl of palm genera
follows Uhl and Dransfield
(1987).
Disperser classes are
indicated as follou's: A : Aves
(birds):
C : Mammalia, order Chiroptera
(bats):
I :
lnsecta
(insects);
M : Mammalia
(mammals,
excluding bats):
P : Pisces
(fish):
and R : Reptilia
(reptiles).
A plus
sign
(+) follou'ing
a class designation
indicates
that
other
unspecified
animal dispersers are suspected.
Taxon Drspersal agent
(class) Reference
( orl phoideae:
Corypheae
Thrinax C o lu
rnha leuc
ocep
hala (A)
T. morrisii
H. A. Wendl. Amazona letrcocephala bahamensis Sn-vder et al.. 1982
(A)
Cl,clura
carinata
(R) Iverson. 1979
Coccothrinax
C. a/ta
(Cook)
Becc.
('. jamaicensis Read
Artibetts lituratus
palmarum(C) Greenhall,
1957
Ptilinopus
pulchellus,
I'. superbus,
P. Frith et al.,1.976
iozonus,
Ducula
spilorrhoa
(A)
Rhapidoph.rllum
hystrix L rsus
tttnerrt,tttus./htrilttttus
lMl Maehr
& Brady. 1984
(Pursh)
H. A. Wendl.
& (M) Shuey & Wunderlin,
1977
Drude)
Lit'istona
Pritchardia
Phoenix
P. dactyliferaL.
P. loureirii KunIh
P. paludosa
Roxburgh (C)
P. pusilla Gaertner
P. reclinata Jacq.
C o lu nfu a leucocep h al a (A)
Coltunba leucocephala
(A)
Pteropus
(C)
Ciridops unna
(A)
(c)
(A)
Eidolon,
Rousettus
(C)
Rousettus aegyptiacus
(C)
Lanius
excubitor
(A)
Elephas
naximus (M)
Osmatreron bicincta
(A)
Loxodonta aliicana (M)
Read.1975
Wiley
& Wiley.
1979
Galeano-Garc6s.
1986
Marshall.
1985
Amadon, 1950; but see
Perkins,
l 903
Guppy,1906
Galeano-Garc6s,
1986
Petch. 1924
Docters van Leeuwen, 1935
Docters van Leeuwen, 1936
Smith, l9l0
Wiley
& Wiley,
1979
Brown,1976
Lancaster. 1964
Marshall. 1985
Ridley, 1930
Parrott, 1980; but see Cowan,
I 984
Krishnan,
1972
van
der Ptjl, 1957
Ridley, 1930
Corner.
1966
(A)
lcoelorrhaphe v,rightii C'olurnba leucocephala (A)
(Griseb.
& H. A. Wendl.)
H. A. Wendl.
ex Becc.
Serenoa
repens
(Banr.) .,lphelocoma
coertrlescens
(A) Woolfenden
& Fitzpatrick, 1984
Small Lrrsus amerit'antus
/loridanus
(M) Maehr & Brady. 1984
Washingtonia
ftlifera Canis latrans, Ltrocyon cinereoaryen- Bullock, 1980
(Linden)
Wendl. te,rr
(M), Sialia me.ricana, S. cur-
rucoides, BombyciLla
cedrcrum
(A)
(lorv-pha
umbraculifera
L. Pteropus
edwardsii
(C.)
C. utan Lamark
Sabal Corvus ossifi'agus, Mimus polyglottos, Martin et al., 1951
Turdus migratorius,
Dendroica
co-
ronata, Dry)ocopus
pileatus,
Mela-
nerpes
carolinus
(A), Procyon lotor,
S. etonia Swingle ex Nash Aphelocoma
coerulescens
(A), Ursr.rs Zona, pers.
obs.
a mer ic a nus
fl
o
r
id anu s (M)
S. palmetto (Walt.) Lodd. ex Mimus polyglottos,
Quiscalus
mexi- Cruickshank, 1950
Schultes cantts,
Aphelocoma
coerulescens,
C
yanoc
it ta c r istata, Age laius
p
hoe-
niceus, Cardinalis cardinalis, Larus
delawarensis
(A)
Atnazona
leucocephala bahamensis Snyder et al., 1982
- (A)
Ursus americanus
floridanus
(M) Maehr & Brady, 1984
Sciurus carol i nensis
(M)
()
1:
4116 s 6v
qy 1,n
cas
(A)
S. causiarum
(Cook)
Becc. Columba leucocephala
(A)
(M)
S. yapa Wright ex Becc. Crypturellus boucardi
(A)
Coryphoideae:
Phoeniceae
Hapalemur griseus
occidentalis
(M) Petter et al., 1977
(c) Schonland.1924
8
T.qBr-r 1. Continued
Taxon
SELBYANA
Dispersal agent
(class)
[Volume 1l l'
T
(
Reference
Coryphoideae:
Borasseae
Borassodendron
borneense
Dransf.
Borassus
B. aethiopumMart.
Hyphaene
H. thebaica
(L.) Mart.
H. petersiana
Klotzsch
ex Mart.
Calamoideae:
Calameae
Laccosperma
Eremospatha
E. wendlandianaDammer
ex Becc.
Eugeissona
lrislls Griff.
Kort hal si a laci
nio s a (Griff
.)
Mart.
Salacca
Daemonorops
melanochaetes
Blume
in Shultes
Calamus
C. australis
Mafi.
C. deerratus
Mann &
H. A. Wendl.
C. moti F. Bailey
C. radicalis H. A. Wendl.
& Drude
C. aff. scipionun Loureiro
Plectocomia
elongata
Mart. ex Blume
in
Schultes
P i
gafet
t a fi I ar i s (Griseb.)
Becc.
Raphia
farinifera
(Gaertner)
Hylander
R. hookeri Mann & Wendl.
R. regalis
Becc.
R. taedigera
(Man.) Mart.
R. vinifera Beauv.
Calamoideae:
Lepidocaryeae
Mauritia
flexuosaL. f.
Ceroxyloideae:
Ceroxyleae
C ero
xylon kl op
st ocki a Mart.
Probably:
Pongo
pygmaeus
(M)
Eidolon, Dolosonia,
Pteropus
(C)
Loxodonta africana capensis
(M)
Papio anubis
(M)
Loxodonta africana (M)
(c)
Papio ursinus
(M)
Mandrillus sphinx (M)
Cephalophus
sylvicultor, C. callipygus
(M)
Pan troglodytes
troglodytes
(M)
C e
p
hal
op hus sy lvicult
or (M)
(M)
Anthracoceros convexus
(A)
(M)
Probably:
(R)
Paradoxur us he r map h
rod
it us
j ava ni'
cas
(M)
(M)
Pa
radorurus h er
maph rod i t us
j ava
n i
-
cus
(M)
H
ylobat
e s sy ndac ty lus (M)
Ducula spilorrhoa (A)
Casuarius
casuarius
(A)
Ptilinopus iozonus,
P. magnificus,
P.
superbus
(A)
Argusianus argus
(A)
Casuarius
casuarius
(A)
Pan troglodytes
troglodytes
(M)
Mandrillus sphinx (M)
Casuarius
casuarius
(A)
Casuarius casuarius
(A)
Hylobates lar (M)
P ar adoxurus
he
r
map hroditus
j av ani
-
cas
(M)
(A, M)
G
y
p
o hier ax a ngo le ns i s (A)
(M)
Xerus erythropas
(M), (A, C)
(M)
(A)
Tapirus bairdii (M)
(M)
Daptrius ater (A)
Cebus albifrons
(M)
Tayassu tajacu, T. pecari (M)
Aulacorhynchus sulcatus sulcatus
(A),
(M)
Dransfield
in Moore, 1973a
Marshall,
1985
Burtt,1929
Lieberman et al., 1979
Corner,
1966
van
der
Pijl. 1957
Hamilton
et al.,
1978
Lahm, 1986
Dubost,
1984
Hladik, 1973
Dubost, 1984
Wong, 1959
Rubeli
in Dransfield,
1981a
Ridley, 1930
Beccari, 1877
Bartels, 1964
Ridley,
1930
Bartels,
1964
Chivers,1974
Crome,
1975a
Crome,
1976
Frith et al.,19'/6
Davison.
1981
Stocker
& Irvine, 1983
Hladik,
1973
Lahm, 1986
Stocker & Irvine, 1983
Stocker
& Irvine, 1983
Ellefson, 1974
Bartels,1964
Dransfield,
1976
Austen,
1953
Otedoh,1979
Profizi,1985
Otedoh,
1979
Otedoh,1979
Janzen,
1983b
otedoh,1979
Haverschmidt,
1962
Defler,1979
Kiltie, 1981b
Braun,1976
r 9891
T.r.nlr l. Continued.
ZONA & HENDERSON: SEED
DISPERSAL OF PALMS
Taxon Dispersal agent
(class) Reference
Ceroxyloideae:
Hyophorbeae
Hvophorbe
Chamaedorea
C. lanceolata
(Ruiz &
Pavon) Kunth
C. tepejilote
Liebm. in
Mart.
C. poeppigiana
(Mart.)
Gentry
Arecoideae: Caryoteae
ArengA
A. listeri Becc.
A. obtusifoliaMart.
A. pinnata (Wurmb)
Merrill
Caryota
C. cumingii
Lodd. ex Mart.
C. mitis Loureiro
C. no Becc.
C. rumphianaMart.
C. urens L.
Arecoideae: Iriarteeae
I riartea ventricosa
Mart.
Socratea
S. exorrhiza
(Mart.)
H. A.
Wendl.
Pteropus
(C)
C hamae
p
et es unicolor, Aul ac or hyn
-
chus
prasinus
(A)
Heteromys
(M+)
(A, M)
Agouti
paca (M)
Steatornis caripensis
(A)
Ptilinopus magniJtcus. P.
aurant ii-
frons, Ducula spilorrhoa, D. zoeae
(A)
Pteropus
(C)
Ducula rosacea whartoni (A)
Pa
radox
u rus he r ma
p
h rod i
t
us
j ava
n
i
-
czs
(M)
Hylobates k/ossii
(M)
(c)
P
ar adoxurus her map hr oditus
j arani
-
cr.rs
(M)
Szs
(M)
Paradoxurus
her maphroditus, Viyerra
malaccensis
(M)
Ducula zoeae, Ptilinopus magnificus
(A)
Viverra malaccensrs
(M)
Parado-rurus
hermaphroditus
javani-
cr.rs
(M)
Anthracocerus
coronatus convexus
(A)
(c)
Pa radoxu rut hcrma
ph
rod
it us
j ata n i-
cus
(M)
Canis aureus
(M)
Ateles
belzebuth
(M)
Tayassu
pecari (M)
(A, C, M)
Cebus, Callicebus moloch, Ateles
pa-
niscus, Tayassu
(M)
Steatornis caripens
is (A)
Tayassu
pecari, T. tajacu (M)
Ramphastos
swainso nii (A)
Ramp
has tos b
r
ev icar i natus
(A)
A rt
ibeus
j a maice ns i
s (C)
Ateles
geofroyi (M)
H et er o mys de s marest ianus (M)
Crax (.\)
Cebus
apella
(M)
Alouatta
palliata
(M)
Cebus
capucinus
(M)
(A, M)
Cebus capucinus, Ateles
geffioyi,
Proechimys
semispinosa
(M)
Saguinus
(M)
Marshall, 1985
Wheelwright
et al., 1984
R. Dirzo, pers.
comm.
Foster et al.,
1986
Gallina in Coates-Estrada & Es-
trada, 1986
Snow,1979
Frith
et al.,1976
Marshall, 1985
Powell & Covacevich,
1983
Bartels, 1964
Whiuen,
1980
Docters
van Leeuwen, 1935
Bartels, 1964
Miller, 1964
Dransfield, 1974
Frith et al., 1976
Ridley, 1930
Bartels, 1964
Dransfield, 1974
Docters van Leeuwen, 1935
Bartels, 1964
Ridley, 1930
Klein & Klein, 1975
Kiltie, 1981b
Foster et al., 1986
Terborgh, l986
Snow,1979
Kiltie, 1981b
Howe, 1983
Van Tyne, 1929
Carvahlo,1961
Hladik & Hladik, 1969
Fleming, 1974
Gottsberger, 1978
lzawa, 1979
Milton, 1980
Oppenheimer,
1982
Foster
et al., 1986
Hogan,1986
R. Ulloa, pers.
comm.
Wettinia maynensis Soruce
10
Tl,sl-e 1. Continued
Taxon
SELBYANA
Dispersal agenl (class)
[Volume I I
Relerence
Arecoideae:
Areceae
Orania
aruensis
Becc
Reinhardtia gracilis
(wendl.) Drude ex
Dammer
Dvpsis
Euterpe
E. edulis
Mart.
E. langloisii Burret
E. precatoria
Marr.
Prestoea
P. montana
(Graham)
Nicholson
Oenocarpus
aff. bacaba
Mart.
O. maporaKarst.
Jessenia
J. batauaBurrer
Hyospathe
elegans
Mart.
H. w^eberbaueri Dammer
ex Burret
Royst
o
nea borinqtteana
Cook
R. oleracea
(Jacq.)
Cook
R. regia
(Kunth)
Cook
Archontophoenix
A. alexandrae
(Mueller)
H. A. Wendl.
& Drude
A. cun ni n
g
ha
m i a na (W
endl.)
H. A. Wendl.
& Drude
C
hambeyronia
macrocarpa
(Brongn.) Vieill. ex Becc.
Actinokenl
ia d ivaricat a
(Brongn.)
Dammer
Calyptrocalyx
Linospadix
L. microcarva
(Domin) Burret
L. monoslachra
(Mart.)
Wendl.
Veitchia
C'asuarius
(A)
Heteromvs
(M+)
Hapalemur siriils
(M)
Cotinga
ridgtrat'i
(A)
Per
iss
oc ep ha I us t
ric
o lor (A)
St eatornis c'aripensis
(A)
.lteles bel:ebuth
(M)
('olossorna
bidens,
Electrophorus elec-
tricus
(P)
Cotinga cotinga.
Phoenicircus carni-
fex, Rupicola rupicola (A)
Ramphastos tucanus,
R. ariel
(A+)
Steatorni s cari
pens
i s (
A)
Celtus upella (M)
Steatornis
cdripensis
(A\
Amazona
rittata
(A)
)I a r
gar
o
p
s
.f
us c at u s (
A)
C olttntba
squatnosa, G eol
r1;gon mon-
tana
(A)
Brvcon
(P)
G
1,
m nod e
rus
.foet
id u
s (
A)
C'ebus
apella (M)
Sciurus
granatensi
s (M)
C-ebus
capucinus
(M)
T'ayassu tajacu, T. pecari (M)
St eat ornis caripensis
(A)
Pithecia monachus
(M)
Cebus
albtfrons
(M)
Cebus apella,
Ateles belzebuth
(M).
Ara macao,
Ramphastos tucanus,
Pipile
cumanensis
(A)
(A.
M)
(A,
M)
C- o I u
rnb
a I eut' oc e
p
ha I a (
A)
.lrtibeus lituratus
palmarum
(C)
St eatornis caripensis
(A)
Thraupis
palmarunt (A)
,\I
yiozetete
s slnilrs (A)
Ptilinoptts superbus,
P. magnificus
(A)
Ducula spilorrhoa
(A)
Pt il i nop us s uperbus
(A)
Casuarius
casuarius
(A)
Lop ho lai mus ant arct icus
(A)
Ducula
goliath (A)
(A)
Casttaritts bennett
i pictocollis
(A)
Casuarius casuarius
(A)
Casuarius casuarius
(A)
Ailuroedus crassirostris,
Ptilonorhyn-
chus violacetts
(A)
(A)
Beccari. I 877
R. Dirzo.
pers.
comm.
Petter et al..
I 971
Skutch,
1969
Snow.
1972
Snow.
1979
Klein & Klein.
1975
Goulding.
1980
Snow,1982
Edwards in Ridley, 1930
Snow.
1962
Izawa, 1979
B. Tannenbaum,
pers.
comm
Little & Wadsworth, 1964
Recher & Recher.
1970
Janzen.
1972
Goulding,
1980
Novaes.1980
Izawa, 1979
Heaney & Thorington. 1978
Oppenheimer.
1982
Kiltie. 1981b
Snow.
1979
R. Ulloa,
pers.
comm.
Defler. 1979
Izawa,
1979
Foster et al..
I
986
Foster
et
al.. 1986
Wiley
& wile1.
1979
Greenhall.
1957
Snow,
1962
Snow & Snow.
l97l
Skutch,
I 960
Frith et al..1976
Crome,
1975a
Crome,
1975b
Crome, 1976
Fnth, 1957
MacKee et al..
1985
Pancher in Linden, l88l
Pratt,1983
Crome,
1976
Stocker & Irvine. 1983
Donaghey.1981
Guppy,1906
19891 ZONA & HENDERSON:
SEED DISPERSAL
OF PALMS
TrsI-e
1. Continued.
ll
Taxon Dispersal
agent
(class) Reference
Pinanga
coronata
(Blume Paradorurus
hermaphroditus
javani- Bartels,
1964
Ptychosperma
aff. macar- (A)
thurii (Wendl.
ex
Veitch)
H. A. Wendl.
ex Hook. f.
Nenga
gajah Dransfield (M)
ex Mart.) Blume
4reca
(c)
(A)
Arecoideae:
Cocoeae
Butia leiospatha
(Barb.- (I)
Rod.) Becc.
(M+),
(C.
r, R)
Svagrus loefgrenii Glassm. (A, M, R. C. l)
S. orinocoensis
(Spruce) Cebus albifrons
(M)
Burret
Allagoptera arenaria (I)
(Gomes)
Kuntze
lttalea
Docters
van Leeuwen, 1935
Dransfield.
1975
crs (M)
Casuarius
bennetti
pictocollis (1t) Pratt, 1983
Ono & Sugawara, 1981
MacKee
et al., 1985
House.1984
Docters van Leeuwen, 1935
Docters van Leeuwen, 1936
Silberbauer-Gottsberger,
I 973
Rhea
americana
(A), Cerdocyon Gottsberger
& Silberbauer-Gotts-
thous, Chrvsocyon
brachvttrus berger. 1983
Iguanura
wallichiana Argusianus
argus
(A) Davison.
1981
(Mart.)
Benth.
& Hook.
ex Becc.
Brongniartikentia wginata Cyanorantphus
novae:elandiae
(,\\ L6tocart
in MacKee
et al., 1985
(Brongn.)
Becc.
Clinostigma savoryanum (A)
(Rehder
& Wilson)
Moore & Fosberg
Burretiokentia vieillardii Ducula goliath (A)
(Brongn.
& Griseb.)
Pichi-Ser.
Oncosperma horridum (A, M)
(Griff.) Scheffer
O. tigillarium
(Jack)
Ridle"v Gracula
jatanica, Turtltr
tigrinus
(A) Ridley. 1930
1. regia
(Mart.)
Boer Echimys annatus, Philander
opos- Charles-Dominique et al., 1981
sum, Didelphis rnarsLtpialis
(M)
Cebus,
Saimiri sciureus, Sciurus
(M) Terborgh,
1986
(c)
Cebus albifrons
(M)
Deroptyus
accip itrinus (A)
Gottsberger & Silberbauer-Gotts-
berger,1983
Defler.1979
Morawetz, 1983
van
der
Pijl. 1957
Defler.1979
Mcloughlin & Burton. 1976
Foster et al., 1986
lzawa. 1979
Kaufmann,
1962
Hladik
& Hladik, 1969
Heaney
& Thorington.
1978
Hogan,1986
Scheelea (A, M)
S. attaleoides Karst. Cebus apella (M)
S. zonensis
Barley
S. rostrata
(Oerst.)
Burr. Sciurus
variegatoides,
Agouti
paca, Bradford
& Smith, 1977
Dasyprocta
p
unct
at a, P roec
him-vs
semispinosa
(M)
Nasua narica (M)
Cebus capucinus
(M)
lgouti paca,
Dasyprocta
punctata, Bradford
& Smith, 1977
P r
oe c hi my s s e mi s
p
i nos a (M)
S ci ur us
gr
an at e ns i s (M)
Sciurus
gerrardi (M)
Orbignt,a martiana Barb.- Agouti
paca,
Das
v-procta
punctata
(M) Anderson, 1 983
Rod.
Elaeis
guineensls
Jacq. Gypohierax angolensis
(A) Thomson
& Moreau.
1957
Tockus
.fasciatus,
Ceratog-v-mna elata, Brooke
& Jeffery,
1972
C. atrata, Cortus albtts, Merops al-
bicollis, Falco ardosiaceus
(A)
Tockus alboterminatus, T.
flavirostris, Dean, 1973
Bycanistes sharpii (A)
t2 SELBYANA [Volume I I
Te,sls 1. Continued.
Taxon Dispersal agent
(class) Reference
Acrocomta
A. aculeata
(Jacq.)
Lodd.
ex Mart.
A. vini.fera
Oerst.
Aiphanes
Bactris
B. cuesa
Crueger
ex
Griseb. & H. A. Wendi.
B. gasipaes
Kunth
Desmoncus
Astrocaryum
A. chambira Burrer
A. jauari Mart.
A. mexicanum
Liebman
in
Mart.
A. polystachyum
H. A.
Wendl.
ex Hemsl.
A. standleyanum
Bailey
Ptilostomus
afer (,\)
GaLago
alleni (M)
Papio anubis
(M)
C ricet omys ga
mbianus,
M ast omys
natalensis
(M)
Pan troglodvtes
(M)
Eidolon (C)
(c)
Dasyprocta, Cerdocyon
thous
(M+),
Rhea
ameicana, Tinamus solita-
rlas
(A), (R)
Didelphis albiventris,
Nectomys
squamipes,
Cebus apella,
Agouti
paca, Euphractus
sexcinctus,
DasY'
procta (M), Turdus (A)
Sigmodon
hispidus
(M)
Probably:
Liomys salfini (M)
St eatornis caripe
nsis
(A)
Artibeus
jamaicensis
triniatus,
A. li-
turatus
palmarum
(C)
Steatornis
caripensis
(A)
Cebus albiJrons
(M)
Colossoma
bidens,
Piranha preta (P)
Steatornis caripensis
(A)
Ramphocelus
passerinii (A)
M elanerpes c
hrysauc hen (A)
(c)
H eteromys
des marestianus,
H op lomys
gy
mnurus,
Dasyprocta
punctata
(M+)
Daptrius aler (A)
Steatornis
caripensis
(A)
Cebus
capucinus
(M)
Ramphastos tucanus
(A)
(c)
Rupicola rupicola (A)
Cebus,
Tayassu, Sciurus,
Agouti
(M+),,ara
(A)
Cebus
apella (M)
(P)
Colossoma
macropomum,
C. bidens,
Brycon, Phractocephalus
hemeliote-
rus, Megaladoras irwini, Piranha
preta (P)
Brycon cf . melonopterus,
Myleus, Me-
tynnis. Serrasal
mus.
Leptori nus.
Paulicea lutkeni, Rhamidia schom'
burgkii, Lit hodoras dorsalis,
Mega-
ladoras irwini, Oxydoras
niger, Se-
maprochilodus
(P)
Sciurus aureogaster,
S. deppei
(M)
Ramp hast o s brer ic
ar i natus
(A)
Nasua narica (M)
Cebus
capucinus,
Ateles
geofroyi (M)
D asy
p
roct a p
unct ata (M)
S c iurus
g
ranate
ns i s (M)
Tapirus bairdii (M)
Goodwin, 1976
Molez, 1976
Lieberman et al., 1979
Iwuala et al., 1980
Wrangham
& Waterman,
1983
Marshall,
1985
Leck,
1969
Gottsberger
& Silberbauer-Gotts-
berger,
1983
Scariot,
1987
Baker,
1983
Janzen,1983a
Snow,1962
Greenhall,
1957
Snow,1962
Defler, 1979
Goulding,
1980
Snow,1962
Skutch,1954
Skutch,1969
van
der
Pijl, 1957
Vandermeer, 1983
Haverschmidt,
1962
Snow,1962
Hladik & Hladik, 1969
Bourne,1975
Bonaccorso,
1979
Snow,1982
Terborgh, l986
lzawa, 1979
Gottsberger,
1978
Goulding,
1980
Piedade,1985
Coates-Estrada
& Estrada, 1986
Van Tyne, 1929
Kaufmann,
1962
Hiadik & Hladik, 1969
Smythe,
1970
Heaney & Thorington, 1978
Terwilliger, 1978
l e89l
Tasln 1 Continued.
Taxon
ZONA & HENDERSON: SEED
DISPERSAL OF PALMS
Dispersal agent
(class)
l3
Reference
A. tucumaMart.
A. vulgare Mart.
Arecoideae: Geonomeae
'l4telfia
georgii
H. A. Wendl
Geonoma
G. vaga
Griseb.
& H. A.
Wendl.
Phytelephantoideae
Phytelephas
Ammandra
Deroptyus accipitrinus
(A)
Deroptyus accipitrinus
(A)
Philander opossum, Caluromys
phi-
lander (M)
Potos
flarus. Sciurus. Cebus capuci-
nus,
Dasyprocta
punctata, Hetero-
mys desmarest ianus,
H oplomys
gy
m
nu
rus, P
roec
h i mys se
mi
s
pi
nosa
(M), Probably: Agouti paca, Tayas-
su
pecari (M), Amazona, Ramphas-
/os (A)
P
hai nopti la melanoxant ha, C at harus
gracilorostris,
Myadestes melanops,
C hamaepetes unico
lor (A)
Steatornis caripensis
(A)
Steator
ni s cari
pe
nsis
(A)
Agouti
(M)
Agouti
(M)
Mcloughlin & Burton, 1976
Mcloughlin & Burton, 1976
Charles-Dominique
et al., 198 I
Vandermeer et al., 1979
DeVito, 1983
B. Tannenbaum,
pers.
comm
Snow, 1962
H. Balslev,
pers.
comm.
H. Balslev,
pers.
comm.
cation not only identifies the class ofthe known
dispersers
but also indicates that other unspec-
ified dispersers
ofthat class are suspected.
Rrsulrs
For most palms many different animals are
involved in dispersal
(Tenr-r 1). Not onl-v are
many palms visited by animals from man-v
dif-
ferent classes but also a frugivorous animal ma.v
forage on more than one species of palm. Fur-
thermore. many of the same animals which feed
on palm fruit also feed
on the fruit ofother plants
(e.g.. Lauraceae, Moraceae. Burseraceae) (cf.
Snow,
l98l; Marshall.
1985). As a consequence
of this diversity in feeding ecology, specific an-
imal-palm obligate
mutualisms are
not likel-v
to
have evolved in dispersal (Wheelwright & Ori-
ans. 1982) as they have in pollination (Hender-
son, 1986).
The results
ofthis survey reveal an interesting
diversity of dispersers throughout the tropical
and subtropical areas of Africa. Asia and Ma-
lesia,
Australia and Oceania, and the Americas.
There are comparativell, few palms native to
Africa
(Moore, 1973a, 1973b),
but there are many
unanswered
questions about palm seed dispersal
there.
Burtt (1929) and Corner (1966) have re-
ported that the African elephant is an important
dispersal agent for the widespread Phoenix re-
clinata, Hy,phaene sp., and Borasstts
aethiopum,
although this large animal has a destructive po-
tential which may well lessen its overall effec-
tiveness
(Krishnan. I 972).
In addition, the palm
nut vulture. Gvpohierar angolensis, is well known
for feeding on the fruits of Raphia farinifera.
Shrikes
(Lanitrs excubitor) are thought by Parrott
(
1
980) and others to feed
on the fruits ofthe date
palm, Phoenix dactyli.fera; the partiall-v eaten
fruirs impaled on the leaf spines of the palms
bear
witness
to the shrike's feeding
habits.
Cow-
an (1984).
however, has attributed the partially
eaten impaled dates to infructescences blowing
in the wind against the spines and to the pecking
activities ofthe Spanish sparrow (Passer
hispan-
iolensis).
The activities of the shrike might result
in limited seed dispersal,
but the activities of the
sparrow would not. As with other aspects of the
palms of Africa, much is yet to be learned of
their dispersal biology.
Much of our knowledge of dispersal in Asia
and Malesia comes from observations made by
Ridley (1930) and Bartels
(1964). Of particular
interest are the man-v mammals reported by Rid-
le1- and Bartels which feed on the fruits of Arenga
and Carvota. The pericarps ofthese palm fruits
contain needle-like
crystals of calcium oxalate
which are highly irritating to the mucous mem-
branes of humans, yet wild dogs and palm civets
consume the fruits with no apparent ill effects.
The palm clet Parado,rurtrs hermaphroditus ja-
t'anicus
is especially
important in seed
dispersal.
It is a skilled arborealist. quite capable of climb-
ins even slender
lianas
and is not limited to feed-
t4 SELBYANA [Volume I I
ing on fallen fruits. Bartels
(1964) has observed
the seeds of Pinanga coronata, Daetnonorops
tnclanochaetcs.
and . lrcnga
pirtnuta germinating
from the dung of the paim civet, which usually
defecates
in clearings.
Bartcls has speculated
that
the seeds
of A. pinnata experience
enhanced
ger-
mination after passage
through the gut of the
palm civet;
however,
this h.vpothesis awaits
crit-
ical testing.
Docters van Leeuwen
(1936)
has reported on
the presence
of Corr-pha utan and Oncospenna
tigillarium on the Krakatau islets.
The vegetatlon
of these
areas
was
totally destro-ved
by the 1883
volcanic eruption of Krakatau. but subsequent
explorations
in 1920 and 1
929
revealed the
pres-
ence of O. tigillaritrm and C'. a/an, respectively.
Docters
van Leeuwen
(
1936)
has attributed
their
introduction to the activities of birds. probabl.v
fruit pigeons. Similarly. Atherton and Greeves
(1985)
have attributed the presence
of Calarnus
on Green Island, Queensland, to dispersal from
the Australian mainland b-v the fruit pigeon
Ducula spilorrhoa.
Indecd.
thc palm floras ofis-
lands, especially
volcanic
islands
never connect-
ed
to the
mainland.
provide
the clearest
er
idencc
for long-distancc
dispersal.
In northeastern Queensland. the cassowar-v
(Castnrius casuarius) disperses several rainfor-
cst palms (Stocker
& Irvine. 1983).
The casso-
wary feeds on a great variet.v of fruit including
Calamus and Linospadi,r nticrocart'tt, and seeds
collected from dung germinate satisfactoril-v.
In
Papua New Guinea, the dwarf cassowary.
Casrl-
arius bennetti pictocollis, feeds heavill' on the
fruit of (alyptrocalttx and is also known to feed
on the fruit of an undetermined species
of Arac'a
(Pratt. 1983). Unlike other "bird fruit." palm
fruit taken b.v
cassowaries
must be dropped when
ripe or borne on low-growing palms. The fruits
of the low-growing Linospadir are easil,v
within
reach of the flightless
cassowar-v.
but it must rel)'
on the fallen fruits of CalantLrs and ,lrccct. Al-
though birds are not thought to take yellow fruit
(Gautier-Hion et al.. 1985).
.t-ellow
Calarn
us
fruits
are taken by cassowaries.
In the wa-v it interacts
with palms. the cassowary
behaves
more like a
terrestrial mammal than a bird.
An example
in which animals' foraging
habits
have the potential
to result
in long-distance
dis-
persal is found in the fruit pigeons Ptilinoptt:
superhtts
and P. magnilicus. Birds collected bv
Frith et a|. (1976) near Port Moresby in New
Guinea were found to have in their crops seeds
of ,lrchontophoenix, a palm endemic to Austra-
lia. As Ptilinopu.s
are known to feed in Australia
and regurgitate seeds in a viable condition
(Goodwin. 1983). .lrchontophoeni"t seeds
taken
from the New Guinea birds may well have been
viable.
One might conclude
that the Jtchonto'
phoenix seeds
are taken in Australia and are then
deposited
in New Guinea, in which case
the pr-
geons
ma-v eventually
succeed
in introducing.{r-
chontophoenir to New Guinea. Alternatively,
the) ma) alreadl have introduced.lrchonto-
phoeni,r into New Guinea. and the presence
ol
this palm has escaped
notice: Frith et al.
(1976)
rccorded that the birds thel' observed were no-
tabl.v sedentary. appeared to forage locally' and
gave
no indication
ofnomadic foraging behavior.
In southwestern
Nofih America. Bullock (
1980)
and Cornetl (1985) have demonstratcd
the via-
bility of W'ashinglonia
./tli.fera
secds
lakcn lrom
co-vote
dung. The nomadic foraging of co-votes
strongl.v
suggests
thal the-v
are important in dis-
persing seeds
of W'. filifera, espcciall-v
in trans-
porting seeds
across
unfavorable habitats.
e.g..
between
washes in the desert.
Birds also
leed
on
the fruit. but dispersal
b-v
birds is scasonal
and
thought to be lcss effective.
The co)otes
depenc
on autogenousl-v
dropped fruit and fruit droppec
b1, birds. Limited data suggest
thal autogenous
fruit drop varies among individual trees.
Con-
sequently.
dispersal
b.v
mammals is more likely
for some seeds.
and dispersal
by birds is more
likell' for others.
Palms that readill drop fruit
ma)' or ma-v
not experience
cnhanccd
reproduc-
tive succcss
(b-v n.reans of enhanced dispersal)
over those
with bird dispersed
seeds'
ln Fforida. the bear {'r.strs
attttrit'aritts
dis-
perses
Sabal spp., Serenoa repens, and Rhapi-
dophvllutn hvsn'ix
(Maehr & Brad.v, 1984: Zona,
unpubl.). Saltal etonia, Serenoa repens, and R.
h.v-stri,r
bear their fruit at or near ground level;
the fruit of Sahal palntetto is readil-v
found be-
neath the tree.
Both S. etonia
and R. hJ'strir
are
restricted
to specific
habitats. and in central Flor-
ida bears
are probabl,v
responsible for dispersing
seeds
of these
palms not onll' u'ithin those
hab-
itats but also
across
patches
of unfavorable
hab-
ltat.
ln northern South An.rerica.
thc distribution
of cotingas
(Cotingidac).
birds
which feed on Eu-
tcrpe frujt. is nearl-v
coincidenl with that of Ert-
terpe
(cf
. Lleras
et al..
198'1).
Work b1
Snor'r
(
1982)
sugge
sts that these
frugivores
are
impor.tant
agents
of local dispersal
for palms and othcr fruit trees.
While birds undoubtedl-v
are
important dispers-
ers
of Euterpc', the fruits are also sought b1 hsh
and the eiectric eel. Elctlr, rpllL't
lts el('Ltt
i.tts
(Goulding, 1980).
The local peoplcs
of Amazo-
nia informed Goulding that the eels
congregate
beneath
fruiting Etlerpc trecs
gro*ing in inun-
dated
areas
and "shock" the trees
to induce
fruit
drop.
As it is unlike
l-v
that
electrical
current
could
induce fruit drop. Goulding has noted that this
testimon-v
b;-'
locals seems
to be more folkloric
than factual.
The hsh. including the eel,
feed on
fruit which drop when ripe: moreover. fishermcn
(
l e89l
are able to attract fish b-v
imitating the sound of
iruit falling into water.
The frugivorous fish (Characidae.
Pimelodi-
clae.
Anostomidae. Prochilodontidae. and Do-
radidae) of Amazonia are uniquc components
of
the guild ofdispersers ofthe riverine and inun-
dated forest of that region (Gottsberger.
1978:
Goulding. 1980: Piedade.
1985). They can dc-
stro.v seeds.
as well as disperse
then-r bv cracking
thc cndocarp and digesting
the endosperm:
hou'-
cr-er. thick endocarps
protect some seeds from
lotal predation. Likeu'ise. the bectles
(Coleop-
tcra) described b-v Silberbauer-Gottsbergcr
(
I
973), Gottsberger
and Silberbauer-Gottsbergcr
(
1983), and Morawetz
(1983)
as seed dispersers
of Butia leiospatha, St'agru.s
loe.fgrenii, and -'ll-
ldgoptera arenaria are likell' to be moslll prcd-
atorl' and probabl-v
pla,v
onll' a minor role in
dispersal.
The beetles
oviposit on the fruits and
bur-v them up to 10 cm belou'the soil surfacc.
It
is not known what proportion of thc bcetle-dis-
persed
seeds
survive to germinate.
A summary of TaeI-r I is presented
in Tesle
l. For each tribe, animal dispersers are tabulated
b1' class. Unspecilied
dispersers arc not included
in Ta.er-r 2; hence.
insects.
the
identities of which
are unspecihed
in Te.sI-E l. arc not counted in
Tesr-e 2.
Drscussrox
An effective disperser must remove the peri-
carp from the seed and deposit the seed in a
r-iable
condition at a site suitable
for
germination
and seedling establishment.
Some animals. such
as Indian elephants (Elephas) and civets (Para-
doxtu'us).
swallow the fruit and later defecate the
seed.
Man.v frugivorous birds
(e.g.,
Dttcula, Stea-
rornis caripensis)
simply swallow the whole fruit
and later regurgitate the seed,
which in the casc
oi palms ma.v be too large to pass through the
intestine.
The more dexterous
primates
and bats
arc able to strip the pcricarp from the seed.
which
is thcn discarded.
The effectiveness
of a disperser in depositing
seeds
in a "safe
site" (Harper. 1911).
one
suitable
lor germination
and seedling
establishment. can-
not be evaluated
from dietar.v data.
It is possiblc
that some of the animals which feed on palm
fruits are ineffective dispersal agents: however.
L-\'en
a low level of cffective dispersal ma-v be
!-nough
to maintain a stable
distribution of the
plants
involved
(Janzen.
1970:
Hubbcll. 1979).
The distinction between seed disperser and seed
prcdator ma-v oflen be fine and tenuous. if not
:omewhat artifrcial.
There exists a continuum
from high quality dispersal (effective dispersal)
to low (predation).
According to Janzen
(1970).
a sccd predator eats the seed
(or seed
plus peri-
ZONA & HENDERSON: SEED
DISPERSAL OF PALMS l5
TAsre 2. Number of dispersers
of palms b.v class.
Palm taxa are
follo*'ed bl the number of genera
found
in Trsle l. LJnspccilied
disperscrs
are
not
included:
hence. class
Insecla is
not
counted.
Within
each
palm
tribe. each animal
is counted
onl-v once
er.en
though
it ma1 disperse
man)'
palms.
Class
abbreviations
are the same
as
in TrsI-r 1.
Class
Taxa
Coryphoideae:
Corypheae
(10)
Phoeniceae
(
1
)
Borasseae
(3)
Calamoideae:
Calameae
(10)
Lepidocaryeae
(1)
Ceroxl'loideac:
Ceroxyleae
(
I
)
Hyophorbeae
(2)
Arecoideae:
Caryoteae
(2)
Iriarteeae
(3)
Areceae
(24)
Cocoeae
(
I 2)
Geonomeae
(2)
Ph1'tclcphantoideae
(2)
carp) and destro-vs thc embryo. Alternativcly, a
seed
predalor
ma1'leave
the seed undamaged but
habituatl-v
deposit the seed in an unsuitable site.
Hovu'ever.
seed
predators ma)- at times act as seed
dispersers
("d1,szoochory"
ofvan der Pijl. 1
982).
A predator ma,v occasionall;- eat fruit and discard
seeds
(cf. lzawa. 1919). incompletel.v masticate
sccds
(cf.
Goulding. 1980).
or fail to rccover
scat-
ter-hoarded seeds
(cf. Heaney'
& Thorington.
I
978).
A seed
predator.
iffrightened or distract-
ed. ma1' abandon a potential meal. thereby ef-
fccting dispersal. On the other hand. evcn the
most elhcient dispersers. such as oilbirds. occa-
sionalll' deposit seeds in unsuitable habitats.
Agoutis.
considered to be both predator
and dis-
pcrscr (Bradford & Smith. 1977:
Vandermeer
et
al., 1979), can in fact play an inconsequential
role in either seed
predation
or dispersal
(Larson
& Howe
. 1987).
Onl-v
morc thorough licld study
can determine relative importance ofthe animals
listed in Taslr I in the dispersal
of palms.
For thesc
rcasons.
the distinction between seed
predator and seed disperser
is not emphasized
in this review. We prefer not to maintain the
incongruous
distinction between animals which
are usuall-v dispersers but are occasionall-v
pred-
ators and those which are usuall_v
predators bul
occasionally disperse seeds. The resultant dis-
pcrsal
of seeds differs in the frequenc-v
or quantity
of seed dispersed. but because
quantitative data
for palm seed dispersal are not available, we can-
243501
13300
03300
80900
10300
10000
31200
61500
4 | 1,1 0 0
3211230
23341. 11 0
70900
00100
l6
not discuss the consequences
for palm popula-
tion dynamics of low frequency versus high fre-
quency seed dispersal.
For many plants, aspects of morphology such
as
color, size, and accessibility have been shown
to influence food choice by dispersal agents and
thereby presumably affect ultimate distribution
(Stiles, 1982; van der Pijl, 1982; Willson &
Thompson, 1982;
Janson, 19831 Moermond &
Denslow, 1983:
Gautier-Hion et al., 1985;
Wheelwright, 1985). Doubtless these attributes
of palms also influence their dispersal; however.
the questions of how specific
morphological fea-
tures of palms influence feeding activities and
subsequent dispersal have only rarely been ad-
dressed.
Beccari
(1877) thought that several as-
pects of palm morphology (viz.. the scales on
fruits of the Calameae) actually hindered seed
dispersal activities of animals. but the evidence
presented
here (Teor-E
I ) clearly does
not support
such
a conclusion. Corner (1966) has suggested
that small animals disperse small palm fruit and
that large
animals disperse
large fruit (see
Wheel-
wright, 1985), and Snow
(1971)
has
stated that
fruits of Bactris are too firmly attached to the
infructescence to be taken by the frugivorous
bearded bellbird (Procnias awrano) but are ac-
cessible to the larger oilbird (Steatornis caripen-
JrJr.
The trends in dispersal apparent from Tenrp
2 likely reflect a relationship between palm mor-
phology and disperser behavior or morphology.
For example. the fruits of the Borasseae
and Phy-
telephantoideae tend to be large, fibrous. and
heavy. so, not unexpectedl-v,
no birds are iden-
tified in Tnsre I as dispersers of these palms.
The scaly fruit and spiny infructescences
of the
Calamoideae may preclude bats as dispersers.
The large number of fish
which disperse Cocoeae
is probably accounted for by the large number
of Cocoeae
in the Amazon basin where
this un-
usual form of seed dispersal is most highly de-
veloped. Curiously, reptiles are identified only
once in Tasr-r 1. whereas
birds and mammals
are abundantly represented.
Several
important factors influencing dispersal
and distribution are not evident from Tasr-E 1.
viz., seasonal
variation in seed dispersal and
postdispersal predation.
The quality of dispersal may vary seasonally,
independentl_v
of fruit production. For example.
Steatornis caripensis feeds on the oil-rich fruits
of several different palms. In Venezuela, the oil-
bird roosts in caves, and during the breeding
season, thousands of seeds are deposited in the
caves in what are obviously unsuitable sites for
germination. However. during the rest of the year,
most seeds are regurgitated as the birds forage
throughout the forests
thereby effecting dispersal
[Volume I I
(B.
Tannenbaum,
pers.
comm.). The oilbird sim-
ilarly disperses
palms in Trinidad (Snow & Snow,
I 978) and Ecuador
(Snow, I
979).
Janzen (1971) has shown that Scheelea seeds
deposited by rodents beneath the parent tree ex-
perience up to 80 percent mortality because of
predation by bruchid beetle larvae; moreover,
Wilson and Janzen
(1972) have shown that the
postdispersal distance between a seed and the
parent tree does not influence the probability of
bruchid predation. Bruchid predation is a "filter"
between dispersal and ultimate distribution (see
also Brown. 1976).
Kiltie (198
la) has
shown that
palm seeds scatter-hoarded by squirrels may be
recovered
by peccaries
which eat the endosperm.
Additionally. there may be differences in post-
dispersal
predation depending on dispersal agents:
Bullock (1980) has noted that seeds of I4laslz-
ingtonia.filqfera dispersed by mammals may ex-
perience
less
predation by rodents than seeds dis-
persed
by birds. These limited examples suggest
that postdispersal predation ultimately influ-
ences the distribution of palms: however, biol-
:i:t;: n"". only begun to examine this phenom-
Cotingas, cassowaries, and fruit pigeons are
specialized frugivores in that they feed almost
exclusively on large. highly nutritious fruits.
McKe.v
(1975)
and Snow(1971,
198 l) assert that
a certain degree of "mutual evolution" may have
occurred in both palm and frugivore as the result
of frequent inclusion of palm fruits in the diets
of these animals. The diversitl' of animals that
feed on palm fruit, however, suggests
that highly
coevolved plant-disperser obligate mutualisms
are not in operation (Vandermeer et al., 1979;
Wheelwright
& Orians. 1982). The high nutrient
content of the fruit and the large seed size of
man-v
palms suggest dispersal by specialized
fru-
givores
(McKey. 1975;
Snow. l98l). but the bony
or fibrous endocarp suggests selection pressures
from additional kinds of animal dispersal agents
and/or postdispersal predation. Although spe-
cialized frugivores may provide a higher flre-
quency of quality dispersal
(McKe-v.
1975), op-
portunistic
frugivores.
those
animals
which take
fruit only occasionally
or seasonally. are also
im-
portant in the dispersal of seeds
and the ultimate
distribution of palms. The signilicance of op-
portunistic frugivores has been demonstrated in
several
instances
(Hladik & Hladik. 1969; Bul-
lock, 1980; Cornett,
1985).
CoNcrusroxs
Palm fruits are important sources of food for
many animals, and zoochory is common in the
Palmae. The remarkable diversitf in fruit mor-
phology allows many different classes of seed dis-
SELBYANA
F
"i:
-
fcr
\:\.
aj'
3-
\1
\ l'.
:-.
il
m-
:dS
-'\
-
Of
hr-
L-
ilt
of
:r"
ral
be
n].
\l-
tc
'\l
15.
al
\!-
rll
'ls
JI
l\
,t
9'
nt
ili
.1-
t\
a\
l5
!--
(--
r e89l ZONA & HENDERSON: SEED
DISPERSAL
oF PALMS I7
AusrEN,
W. M. 1953. Palm-nul
vulrures
Gypohierax
angolensis
in Raphia palms at Mtunzini. Zulu_
land.
Osrrich
24: 98-102.
Blxrn, R. H. 1983. Sigmodon
hispidus (rata
algo-
donera
hispida,
hispid
cotron
rar).
pp. 490492 in
D. H. JaNzrN, ed., Costa
Rican natural
historv.
Univ.
of Chicago Press.
Chicago.
BnnrEr-s,
E. 1964. On Paradoxurus
hernaphroditus
jatanicus
(Horsfleld.
1824).
Beaulortia
t0: 193-
201
.
Brccenr,
O. 1877. Della
disseminazione
delle
palme.
Bull.
Soc.
Tosc.
Onic. 2: l6j-173.
BoNacconso.
F. J. 1979. Foraging
and reproductive
ecolog_v
in a Panamanian
bat communitv. Bul ,
Flonda
Stare
Mus.,
Biol.
Sci. 24:
359-40d.
BounNr,
G. R. 1975. The red-billed
toucan
in Guv_
ana.
Liring
Bird
l3:
qq-126.
Bneopono,
D. F. eNo C. C. Slrrrn. 1977. Seed
ore_
dation
and seed
number
in Schcelea
palm
fruits.
Ecology
58:667-673
BRAUN,
A. 1976. Variousobservations
on Ceroxvlon
klopsrockia.
Principes
20: I
58-l 66.
Bnooxr, R. K. lNo R. D. JerrEnv. 19j2. Observa_
trons
on the biology
of Gypohierax
ansolensis
in
western
Angola.
Bull.
Brit. Ornithol.
Club 92: l5-
BnowN,
K. E. 1976. Ecological
studies
ofthe cabbaqe
palmetto,
Saba I
palmetto.II.
Dispersal,
predarion.
and
escape
ofseeds.
Principes
20:49-56.
Bur-locr, S. H. 1980. Dispersal
of a desert
palm by
opportunisric
frugivores.
principes
24:
29-32.
Bunrr. B. D. 1929. A record
of fruirs
and seeds
dis_
persed
b"v mammals
and birds lrom the Sinsidz.
Districr
of Tanganl
ika
Terrirorv.
J.
Ecol.
l7: 3l l-
355.
Cenr-qursr,
S. 1974. Island
biology.
Columbia
Univ.
Press,
New York. 660 pp.
Cenvenlo. C. T. or. 196
l. Sobre
os h6bitos
alimen-
tares
de Phillostomides
(Mammalia,
Chiroptera).
Revista
Biol.
Trop.
9: 53-60.
Cuenles-DovrNrqun. p., M. ArnelrrNrowrcz. M.
CH,cnrEs-Dor\arNrqur.
H. GEn_lno,
A. Hleorr. C.
M. Hrnorx.
eNo M. F. pnfvosr. l9gl. Les
mammilEres
frugivores
arboricoles
nocturnes
d'une
for0t guyanaise:
inter-relalions
plantes-animaux.
Terre
& Vie 35:
341-435.
Curvrns,
D. J. 1974.
The
siamang
in Malaya.
Contr.
Primatol.
4: l-335.
Coarrs-Esrneoe,
R. lxo A. Esruon. 19g6. Manual
de idcntificaci6n
de campo
de los mamiferos
de
la
estaci6n
de
biologia
"Los Tuxtlas.,'Universidad
Nacional
Aut6noma
de M6xico,
Mexico.
l5l pp.
ConNrn.
E.
J.
H. 1966.
The
narural
history
ofpalmi.
Univ. Calilornia
Press,
Berkeley.
393
pp.
ConNerr,
J.
W. 1985. Germination
of Washinetonia
filifera
seeds
eaten
by
co1.ote.
principes
2g: I
q.
CoweN,
P. J. 1984. Impaled
dates:
b1,.
shrikes
or bv
chance?
Bull.
Ornirhol.
Soc.
Middle
E. l3: 7.
g.
Cnovr, F. H. J. 1975a.
Breeding,
feeding.
and
status
of the Torres
Strait pigeon
at Low isles.
north_
eastern
Queensland.
Emu
75: lg9_19g.
- I 975b. The ecology
of lruit pigeons
in trop-
rcal
northern
Queensland.
Austral.
Wildlife Res.
2:155-185.
-. 1976. Some
observations
on the bioloev
of
iL,
t-
lc-
t-
n
1-
pcrsers
to exploit palms in different ways. With_
out more detailed
data analysis,
ir is difficult to
identify
dispersal
"syndromes." but some
classes
ofdispersal agents
show a tendency to avoid cer_
rarn groups of palms. i.e., bats and the cala_
moideae.
birds and the Borasseae.
However. as
more field studies are completed, trends identi_
hed in Tnnre 2 may be obliterated or reversed.
Palms attract a wide variety of frugivores, both
specialized
and opportunistic, many of which
disperse
seeds,
so claims that palms are poorly
drspersed
seem unjustified. Of the 200 known
gcnera
of palms, only 75 are listed in Tesl-r l;
however, as more fieldwork is completed, we ex_
pect to discover that the colorful, fleshy fruits so
characteristic of most palms are indicative of
thcir dispersal
by animals.
There
is
no doubt that
animals
have
influenced
the
distribution of palms.
At this time. however.
onlr,a few examples
of range
extensions
or long_
Jistance
dispersal
(vit.. Calamus. Corypha.
Oi_
.' ).t
p
e r tna, and perhaps
A rc
h o n
t o
p
ho e
ni x) can
be
directly attributed to the activities of dispersal
agents.
Of course.
some
species
and even
genera
have very
restricted
distributions.
but untilmore
is known of their ecology.
we can only speculate
i)r reasons
for limited distributions. Because
scr
nan)' palms with fleshy fruits are adapted to
zoochory. small distributional ranges may be
caused
more by limited habitat,
climate,
or sub_
strate
than by a lack ofdispersal agents.
AcxNowtrocMENTS
Man.v individuals provided invaluable sue_
gt'stions and assistance
in preparing this papei.
\\'e gratefully acknowledge the following:- B. E.
Busenberg. Rodolfo Dirzo, Clifford g. frith.
\limmo Iannelli. Kent D. perkins. David W.
Snow.
Bernice
Tannenbaum. and David R. Wells.
Spc-cial
appreciation
goes
to Sherwin Carlquist
.rnd Kay Tomlinson. borh of whom reviewed
.lrafts
of this manuscript.
Lrrrrurunn Clrro
\rr.cDoN,
D. 1950. The Hawaiian honeycreepers
(Aves,
Drepaniidae).
Bull.
Amer.
Mus.
Nar.
Aist.
95:151-262.
\\{ERrcAN
OnNtrrroLocrsrs'
UNroN. 19g3. Check_
lisr
of North
American
birds,
6th
ed.
g77
pp.
l\DERsoN,
A. B. 1983. The biology of Orbignya
tnartiana
(Palmae),
a
tropical
dry forest
dominint
in Brazil. Ph.D. dissertation.
Univ. of Florida.
Gainesville.
2l I pp.
\rHERToN,
R. G. l.No D. GnErvrs. 19g5. Torresian
imperial-pigeon
Ducula spilorrhoa
on Green
Is_
land.
north-eastem
eueensland.
Emu
g5: 261_263.
18
the cassowary
in nonhern Queensland.
Emu 76:
8-14.
CnutcrsnaNr. A. D. 1950. Ring-billed
gulls
and cab-
bage
palmettos.
Auk 67: 273.
DevrsoN, G. W. H. 198 1
. Diet and dispersion
of the
great
argus Argusianus
argrs. Ibis 123:.185-49.1.
DeeN, W. R. J. 1973. Noles on a lanner with mal-
lormed bill. and on hornbills leeding on oil palm
fruits. Bull. Brit. Ornithol. Club 93: 55.
Drrun. T. R. 1979. On the
ecologl' and behavior
of
Cebus albi.frons in eastern Colombia: I. Ecolog1,.
Primates 20:. 475-190.
l)rVtro. E. D. 1983. Comparison of lruit removal
rates in understory palms. Geononta spp., b.v aVian
dispersers rn tw'o lower cloud forest habitats of
different sizc. Pp. 235-237 rn R. G. ZeHanv eNo
W. A. Hesrn. eds.. Tropical biolog-v: an ecological
approach. Organization for Tropical Studies.
Dur-
ham. Norlh Carolina.
Docrpns vlN LeEUweN. W. M. 1935. The dispersal
of plants b-v
fruit-eating
bats.
Gard. Bull. Straits
Settlem.9: 58-63.
-. 1936. Krakarau.
I 883-1933. A. Botan,v.
Ann.
Jard. Bot. Buitenzorg
1647: l-506.
DoNncnrv, R. H. 1981. The ecolog-v and evolution
of bowerbird mating systems. Ph.D. dissertation.
Monash Univ.. Melbourne. Victoria. Australia.
DnqNsRsro, J. 1974. Notes on Carl:ota no Becc.
and
other Malesian
Can'ota species. Principcs
18:
87-
93. 1975. A remarkable new rVc/?g'd from Su-
matra. Principes 19:
27-35.
1976. A note on rhe habit of Pigafetta /ilaris
in North Celebes. Principes 20: 48.
198 la. A s_vnopsis
of the gcnus Korthal.sia
(Palmae:
Lepidocaryoideae).
Kew Bull. 36: 163-
19 4.l98lb. Palms and Wallace's
line. Pp. .{3-56
in T. C. Wnrruonp. ed..
Wallace's
line and platc
tectonics. Clarendon Press.
Oxford.
Dunosr. G. 1984. Comparison of the diets of lrugir'-
orous forest ruminants of Gabon. J. Mammalog,v
65:298-316.
ErrrrsoN, J. O. 1974. A natural history of white-
handed gibbons in the Mala_van
Peninsula. Pp. 1-
136 in D. M. RuN,rsnucH.
ed.. Gibbon and sia-
mang, Vol. 3. Natural history. social behavior,
reproduction. vocalizations. prehensions. S. Kar-
ger. Basel.
EssIc, F. B. 1978. A revision ofthc genus
Ptt'cho-
s
p
e
r tn
a Lablll. (Arecaceae).
Allertonia 1 : 4 | 5
-17 8.
FrrrnlNc, T.H. 1974. The population ecologl of two
species
of Costa Rican helerom_vid
rodenls. Ecol-
ogy 55:493-510.
Fosrsn. R. B.. B. J. AncE.
eNo T. S.
Wecnrrn. 1986.
Dispersal and thc sequential plant communities
in Amazonian Peru floodplain.
Pp. 357-370 in A.
Esrneoe aNo T. H. Fr-rurNc. eds.. Frugivores and
seed dispersal. Dr. W. Junk. Dordrecht.
Fnrrx, H. L I95l . Food habits ofthe topknot pigeon.
Emu 57:
341-345.
F. H. J. Cnovr. AND T. O. Wolrr. 1916.
Food of fruit-pigeons in New Guinea. Emu 76:
49-58.
Ger-saNo-Ge.nces,
G. 1986. Primer registro de dos
[Volume
I I
generos
de palmas para la flora colombiana. Mu-
tisia 66: l-4.
Geurren-HroN. A.. J.-M. DupeNrrln. R. Qur.rs. F.
Fern. C. Souno. J.-P. Decoux. G. Dusosr. L.
Er,rvoNs. C. Eneno. P. HECKETSWETT-rn.
A.
N{ouNcezr. C. Roussrlnox. LNo J.-M. Tnlolrey.
1985. Fruit characters as a basis ol lruit choice
and seed dispersal in a tropical loresl vertebrate
communill . Occologia
65: 32,1-337.
GooowrN. D. 1976. Crows of the u,orld. Cornell Univ.
Prcss. Ithaca.
354 pp.
-. 1983. Pigeons
and dovcs of the world. 3rd
ed. Cornell Univ. Press.
Ithaca. 363 pp.
Gorrsnrncrn. G. 1978. Sccd dispersal
b-v
fish in in-
undatcd rcgions
of Humait6. Amazonia. Biotro-
pica
l0:170-183.
- AND I. SrI-senseurn-Gorrsspncen. 1983.
Dispersal and distribution in the cerrado vegeta-
tion of Brazil. Pp. 315-352 irr K. Kunrrzzu, ed..
Dispersal and distribution: an international sym-
posium. Sonderbd. Naturwiss. Ver. Hamburg,
No. 7.
GoulrrNc. M. 1980. The fishes and the forest. ex-
ploralions in Amazonian natural hislory. Univ.
Calilornia Press. Berkelcl.
GnarNuALL. A. M. 1957. Food prefercnces
of Trin-
idad fruit bats.
J. Mammalogl 38: 409-410.
Guppv. H. B. 1906. Observations
ola naluralist in
the Pacific betwccn 1896
and 1899.
Vol. II. Plant-
dispersal.
MacMillan & Co.. London. 627 pp.
Herr.rrlroN. W. J.. III. R. E. Buszunr. eNo W. H.
Busxrnx. 1978. Omnir.ory and utilization of lood
rcsources
bv chacma baboons. Papio ursinus. Amer.
Naturalist | 12: 9
| 1-924.
Hrnprn, J. L. 1977. Population biologl' ol plants.
Academic Press. London. 829 pp.
Hevrnsorvror. F. 1962. Notes on the feeding
habits
and food of some hawks of Surinam. Condor 64:
154-158.
HEeNry. L. R. eNo R. W. TnonlNcroN. 1978. Ecol-
og_v of neotropical red-tailed squirrels, Scittrus
granatensis, in the Panama Canal Zone. J. Mam-
malogl'
59: 846-851.
HrNoErsoN, A. 1986. A revicu, of pollination studies
in the Palmac. Bot. Rev. (Lancaster) 52: 221-259.
1987. Sl,stematic studies in the Iriarteinae
(Paimae: Arecoideae: Iriarteeae). Ph.D. disser-
tation. Cit-v Universit-v of New York.
Hr.qox. A. eNo C. M. HLADTK. 1969. Rapports
tro-
phiques entre \.egctation et primates dans la fordt
de Barro Colorado (Panama). Terre & Vie 116:
f < | 1l
Hllorr. C. M. 1973. Alimentation er activire d'un
groupe de chimpanzes rcintroduits en foret Gam-
bonaise. Terre & Vie 27: 3,{3-.113.
HocAN, K. P. 1986. Plant architecture.
pholosl'n-
thetic responses.
and population biolog-v in the
palms Socralea durissima and Scheelea :onensis
on Barro Colorado Island. Panama. Ph.D. disser-
tation. Univ. of Illinois. Urbana-Champaign. 180
pp.
Housr, A. P. N. 1984. The ecology of Oncosperma
horridum on Siberut Island. Indonesia. Principes
28:85-89.
Howr, H. K. 1983. Rhamphastos su,ainsonii (Dios
SELBYANA
I ll
\1
u-
.rJ.
iata-
rd..
\,m-
'. a\-
:I\.
l:rrt-
.l rn
:-H.
: rod
\
':
l al'.
.r:115.
-:lrllS
- h+:
i ,
irl-
'r-1lll-
--Jrc-s
-:nac
' lro-
: rret
. 16:
.t
un
r.i
nl -
\\ ll-
- it.'e
( '1\lJ
\\Cf-
ts0
I t
t]1u
:.lpes
I)ros
r
9891
Tede. Toucan de Swainson. Chcstnut-mandibled
Toucan). Pp.
603, 604 in D. H. JANzEN.
ed.. Costa
Rican natural history. Univ. Chicago Press.
Chi-
cago.
Hunrrrr, S. P. 1979. Tree dispersion. abundance.
and diversity in a tropical drl forest. Science
203:
l 29 9- l 309.
IvrnsoN,
J.
B. 1979. Behaviorand ecology
ofthe rock
rguana Cvclura carinala. Bull. Florida State
Mus..
Biol. Sci. 2.1: 175-358.
lwul.t-n, M. O. E., E. I. Bnqroe. nNo N. Meouxe.
1980. Observations on the lood habits of some
African rodents. Revista Biol. Trop. 28:227-236.
Izewe, K. 1979. Foods
and feeding
behavior of wild
black-capped capuchin (Cehus apalla). Pnmales
2O:
57-76.
JrNsoN, C. H. 1983. Adaptation of fruit morpholog,v
to dispersal agents
in a neotropical forest. Science
219:187-189.
.l{NzEN,
D. H. 1970. Herbivores
and the number of
tree species
in tropical forests. Amer. Naturalist
104: 501-528.
1971. The fate of Scheelea rostratd fr:uits
bcneath the parcnt tree:
predispersal
atlack b-v bru-
chids.
Principes l5: 89-101.
1972. Association of rainloresl palms and
secd-eating beetles in Puerto Rico. Ecolog-v
53:
258-261.
-. 1983a. Acrocomia vinilbra
(Coyol).
Pp. 184.
185 in D. H. JANZEN.
ed.. Costa Rican natural
history. Univ. Chicago Press.
Chicago.
-. 1983b. Tapirus bairdll (Danto, Danta. Baird's
Tapir). Pp. 496, 497 in D. H. J.qNzeN.
ed.. Costa
Rican natural histor-v.
Univ. Chicago Press,
Chi-
cago.
- AND P. S.
Menrru. 1982. Neotropical
anach-
ronisms: the fruits the gomphotheres ate. Science
215: 19-27.
K rurvarw, J
. H. 1962. Ecolog.v and social behar.ior
ofthe coati. Nasua narica, on Barro Colorado Is-
land Panama.
Univ. Calif.
Publ. Zool. 60:95-222.
Krrrre. R. A. 1981a. Distribution of palm fruits on
a rain forest floor: why peccaries
forage near ob-
jects.
Biotropica
l3: l.1l-145.
-. 1981b. Stomach contents
of rain lorcst pec-
caries (Ialassr taja('u and 7-.
pecari). Biotropica
t3: 231-236.
KrprN. L. L. lNo D. J. KlerN. 1975. Social and eco-
logical contrasts between four taxa of neotropical
primates. Pp. 59-86 ln R. H. Tur[e. ed.. So-
cioecology and ps-vcholog-v
of primates. Mouton
Publishers.
The Hague.
KnrsuN,r.N,
M. 1972. An ecological surve.v of the larg-
r'r mammals of peninsular India. J. Bomba_v
Nat.
Hist.
Soc. 69:297-351
I ruv. S. A. 1986. Diet and habital preference
of
.llandrillus sphinx in Gabon: implications of for-
aging
stralegy.
American J. Primatol. Il:9-26.
I \\cASrER, D. A. 1964. Life history of the Boucard
tinamou in British Honduras. Part I: distribution
and general
behavior. Condor 66: 165-181.
I \RSoN, D. lNo H. F. Howe. 1987. Dispersal
and
destruction of Virola surinancnsis seeds
by agou-
1is: appearance and reality. J. Mammalog,v 68: 859,
860.
ZONA & HENDERSON: SEED
DISPERSAL
OF PALMS l9
Lecx. C. F. 1969. Palmae: hic et ubique. Principes
I
3: 80.
Lrpspnr\4eN,
D., J. B. Hlll, M. D. SwArNE. AND M.
LrrsrnN4A.N. 1979. Seed dispersal by baboons in
the Shai Hills. Ghana. Ecolog_v
60: 65-75.
LtNoEN, J. 1881. Kentiopsis dit'aricata, Brongn. Ill.
Hort.. Cand 28: 10. I I
Lrrtr, E. L.. Jn. eNo F. H. WeoswonrH. 1964.
Common lrees of Puerto Rico and the Virgin Is-
lands. Agric. Handb. No. 2219.
U.S. Dept. Agric.,
Washington.
D.C. 548 pp.
Llrnns. E.. D. C. Grecorr.rrrrr,.cNo L. ConeorN. 1984.
Areas criticas de distribuci6n de palmas en las
Americas para colecta,
evaluaci6n y conservaci6n.
Pp. 67-101 ril ANoNyruous.
ed.. Palmeras
poco
utilizadas de Am6rica tropical. FAO & CATIE,
San
Jose.
MecKre. H. S..
PH. Moner. eNo
J.
M. Vrrlr-oN. 1985.
Palms in Neu'Caledonia. Principes
29: 166-169.
Mernn. D. S. nNo J. R. Bnenv. 1984. Food habits
of Florida black bears. J. Wildlife Managem. 48:
230-23
5.
MansHelI-. A. G. 1985. Old World ph-vtophagous
bats
(Megachiroptera) and their food plants: a sur-
ve1,.
Zool. J. Linnean Soc. 83: 35
1-369.
Me.nrrN. A. C.. H. S.
Zrv. eNo A. L. NemoN. 1951.
American wildlile and plants. Dover Publications,
New York. 500 pp.
McKrv. D. 1975. The ecolog-v of coevolved seed
dispersal s-vstems. Pp. 159-191 in L. A. Grr-srnr
rNo P. H. RevEN. eds..
Coevolution of animals
and plants.
Univ. Texas Press. Austin.
Mcl-oucnr-rN. E. eNn P. J. K. BunroN. 1976. Notes
on the hawk-headcd parot Daroptt'us
accipitrinus.
Bull. Brit. Ornithol. Club 96: 68-72.
Mtllrn, R. H. 1964. The versatile
sugar
palm. Prin-
cipes 8: I l5-1,17.
Mu-roN. K. 1980. The foraging strateg_v of howler
monkc-vs.
Columbia Univ. Press. Ncw York. 165
pp.
MoEnuoNo. T. C. eNo J. S. DeNsr-ow. 1983. Fruit
cholce in neotropical birds: effects offruit t_vpe and
accessibilit-v
on selcctivitl'.
J.
Anim. Ecol.
52:407-
420.
Mor-Ez. N. 1976. Adaptation alimentaire du Galago
d'Allen aux milieux forestiers secondaires.
Terre
& Vie 30:
210-228.
Moonr, H. E., Jn. 1963. An annotated checklist
of
cullivated palms. Principes 7: I l9-182.
-. 1973a. Palms in the tropical forest ecosl.s-
tems of Africa and South America. Pp. 63-88 in
B. J. Mrccpns. E. S. AyENsu. AND W. D.
DucxwonrH. eds.. Tropical lorest ecosvstems in
.\flrca and South America: a comparative review.
Smithsonian Institution Press.
Washington.
D.C.
1973b. The major groups
of palms and their
distribution. Gentes Herb. | |
: 27
-I 40.
- AND N. H. Urtl. 1973. Thc monocor-vledons:
their evolution and comparatir.e biolog-v. VI. Palms
and the origin and evolution ofmonocotyledons.
Quan. Rev. Biol.48: 114-436.
- AND 1984. The indigenous
palms of
New Caledonia. Allertonia 3: 313-402.
Monawerz, W. 1983. Dispersal
and succession in an
extremc troDical habitat: coastal sands and xeric
20
woodland in Bahia (Brazil).
Pp. 359-380 in K.
KusrrzKr,
ed.,
Dispersal
and distribution:
an in-
temational symposium. Sonderbd. Naturwiss. Ver.
Hamburg, No. 7.
NoveEs, F. C. 1980. Observaq6es
sobre a avifauna
do alto curso do Rio Paru de Leste. Estado
do
Par6. Bol. Mus. Paraense.
No. Ser.: Zool. 100:
l-
58.
ONo,
M. ero T. SucewARA. 1981. An analysis of
the
flowering
plant
flora
ofthe Ogasawara
(Bonin)
Islands with regard to their mode of dispersal.
Ogasawara
Res.
5: 25-40.
OerrNurrrrarn,
J. R. 1982. Cebus capucinus: home
range,
population
dynamics. and interspecific
re-
lationships. Pp. 253-289
ln E. G. LrrcH.
Jn..
A.
S. ReNo, eNo D. M. WrNsoR,
eds., The ecology
of a tropical forest:
seasonal
rhythms and long
term changes. Smithsonian Institution Press.
Washington,
D.C.
Orroon, M. O. 1979. Methods
of seed dispersal in
Raphia
palms.
Nigerian Field
44: 127-130.
P.nnnorr,
J. 1980. Frugivory
b-v
great grey
shrikes
Lanius excubitor. Ibis 122:
532-533.
PrnzuNs. R. C.
L. 1903. Vertebrata.
Fauna Hawaiien-
sis l: 365-460.
Pnrca, T. 1924. Gregaious flowering.
Ce-vlon
J.
Sci..
Sect. A, Bot. 9: 101-1 17.
Pnrren,
J.-J.,
R. AlsrcNec. e.No Y. Ruvplrn. 1977.
Faune
de Madagascar 44: mammileres
lemuriens
(Primates
Prosimiens).
ORSTOM/CNRS. Paris.
Prroeor, M. T. F. 1985. Ecologia
e biologia repro-
dutiva
de,4s/rocanun
jauar Mart.
(Palmae)
como
exemplo de populacao
adaptada
as areas inun-
daveis do Rio Negro
(Igapos).
Conselho Nacional
de Desenvolvimento
Cientifico e Tecnologico.
Manaus.
Amazonas, Brazil. 184
pp.
Powrrr-, D. .q.NoJ.
Covecrvrcn. 1983. Lister's
palm.
Arenga
listeri, on Christmas Island:
a rare
or vul-
nerable
species?
Principes
27 : 89-93.
Pnerr, T. K. 1983. Diet ofthe dwarfcassowary
Ca-
stnrius bennetti
picticollis
at Wau. Papua
New
Guinea. Emu 82(suppl.): 283-285.
PnonzI,
J.-P. 1985. Raphia
haoken:
a survev ofsome
aspects
of the growth of a useful
swamp lepido-
caryoid
palm
in Benin
(West
Africa).
Principes 29:
108-l
14.
Rreo. R. W. 1975. The
genus
Thrinax
(Palmae:
Cor-
yphoideae).
Smithsonian Contr. Bot. 19:
l-98.
Rpcnnr.,
H. F. ,c.No
J. T. Rrcnn. 1970. Appendix:
a contribution
to the knowledge
of the avilauna
oflhe Sierra de Luquillo. Puerto Rico.
Pp. E86-
E93 in H. T. Oouv eNo R. F. PrcroN,
eds.. A
tropical rain forest.
Book 2. U.S.
Atomic Energv
Commission, Washington.
D.C.
RrDLEy,
H. N. 1930. The
dispersal of plants
through-
out the
world.
L. Reeve
& Co.. London.744
pp.
Roosvrr-rr, M. G. M. veN. 1985. Fruits of rhe
Guianan flora. Instilute of Systematic
Botany.
Utrecht Univ., Utrecht.
,183
pp.
Sclnror, A. O. 1987. Biologia reproductiva
de .tc-
rocotnia
aculeata
(Jacquin)
Loddiges
ex Martius
(Palmae)
no Distrito Federal. Master
of ecolog-v
thesis.
Universidade de Brasilia.
ScnoNI-nNo, S. 1924. On the theoryof 'ageandarea.'
Ann. Bot.
(London)
38: 453472.
[Volume I I
Ssupv,
A. G.,c.ND
R. P.
WurpenrrN. 1977. The needle
palm:
Rhapidophyllum ht,strir. Principes 2I: 47-
Srr-sERseurn-GorrsBERGER. I. 1913. Bliiten- und
Fruchtbiologie
von Butia leiospatha
(Arecaceae).
Osterr.
Bot. Z. l2l,: 171-185.
Srurcn, A. F. 1954. Life histories of Central
Amer-
ican
birds.
Pacific Coast Avifauna 3l: l-448.
-. 1960. Life histories
ofCentral
American birds,
II. Pacific
Coast
Avilauna 34: l-593.
-. 1969. Life histories of Central
American
birds.
III. Pacilic
Coast
Avilauna
35:
l-580.
Surrn. A. P. 1910. Miscellaneous bird notes from
the
lower Rio Grande. Condor
l2: 93-103.
Svvrsr, N. 1970. Relationships between fruiting
seasons and seed dispersal
melhods in a neotrop-
ical forest. Amer. Naturalist 104: 25-35.
SNow,
B. K. 1912. A fie1d
study of the callbird
Pe-
rissocephalus
tricolor.
Ibis I 14: 1,39-162.
1979. The
oilbirdsoflosTayos.
Wilson Bull.
9l:159-161.
- AND D. W. SNow. 1971. The leeding ecology
of tanagers and honel,creepers in Trinidad. Auk
88:
29 l-322.
SNow, D. W 1962. The natural history
of
the oilbird,
Stearornis caripensis, in Trinidad. W.I. Part 2.
Population.
breeding
ecology.
and
food. Zoologica
47(1):199-22t.
-. 197 l. Evolutionary
aspects of fruit-eating
by
birds. Ibis I l3: 194-202.
1981. Tropical frugivorous
birds and their
food
plants:
a world
surve-v. Biotropica l3: l-14.
-. 1S82. The
colingas. Cornell Unir.
Press. Irh-
aca.
203
pp.
- AND B. K. SNow.
1978. Palm fruits in
the diet
ofthe oilbird. Steatornis caripensis.
Principes 22:
I 07- I 09.
Snyopn, N. F. R.. W. B. KrNc, eNo C. B. Krprrn.
1982. Biolog-v
and conservalion of the Bahama
parrot.
Living Bird l9: 9l-1 14.
Srus, E.
W. 1982. Fruit flags: two hypotheses. Amer.
Naturalist 120: 500-509.
Srocxrn. G. C. nNo
A. K. InvlNr. 1983. Seed dis-
persal
b.v cassowaies
((-
asuari us cas uari us) in north
Queensland's
rainforests. Biotropica | 5: 170-17 6.
TpnsorcH,
J. 1986. Community aspects of lrugivory
in tropical forests. Pp. 371-3841r A. Esrn-lo.q.nruo
T. H. FlevrNc. eds..
Frugivores
and seed
dis-
persal.
Dr. W. Junk.
Dordrecht.
TrnwrLrcen, V. J. 1978. Natural hisrorl of Baird's
tapir on Barro Colorado
Island, Panama
Canal
Zone.
Biotropica l0: 2l l-220.
TsonsoN, A. L..q.No R. E. MonE.cu. 1957. Feeding
habits of the palm-nut
vulture Grplohierax. Ibis
99: 608-6 1 3.
UHr-. N. W. eNo
J.
DncNsnelo. 1987.
Genera
pal-
marum. Allen Press. Lawrence.
Kansas. 610
pp.
VeNornvprn,
J.
H. 1983. Pejiba.ve
palm (pejibaye).
Pp. 98-100 ir D. H. JeNzrN.
ed.. Costa Rican
natural historl'. Univ. Chicago Press,
Chicago.
J.
Srour.
.cNo
S.
RrscH. 1979.
Seed dispersal
of a common Costa
Rican rain forest
palm (VL
elfia
georgii).
Trop. Ecol. 20:- 17-26.
vAN DER Pur-. L. 1957. The dispersal of plants
by
bats
(chiropterchory).
Acta Bot. Neerl.
6: 29 l-315.
SELBYANA
r
st9l
- I
982 Principlcs ofdispersal rn highcr
plants.
3rd ed. Springer-Verlag.
Berlin.215 pp.
\ \\ TyNE,
J. 1929. Lile history ofthe toucan Rarn-
t)
h ust o s brer
i car i nal us. U ni\ . Michigan Mus. Zool..
\{isc. Publ. 19: l-.13.
\\ HEELwRTcHT.
N. T. 1985. Fruil size.
gape
width.
and rhe diets offruit-eating birds. Ecolog_v 66: 808-
818.
-. W. A. Hesrn, K. G. Munnay, eND C. GurNDoN.
I
984. Tropical fruit-eating birds and their food
plants: a suwey of a Costa Rican lower montane
forest. Biotropica l 6: 17 3-192.
- AND G. H. Onte.Ns. 1982. Seed dispersal b-v
animals: contrasts
\\'ith pollcn dispcrsal. problems
of terminolog.v. and constraints on coevolution.
\mcr. Naturalist 1 l9: ,{02-.113.
\\ srrrnN. A. J. 1980. .|rt'nga frurl as a lood for gib-
bons.
Principes 24: 113-146.
\\ rrEy.
J. W. AND B. N. Wllrv. 1979. The biolog-v
ZONA & HENDERSON: SEED
DISPERSAL OF PALMS 2l
re
rd
:1,
r-
is.
1s.
m
p-
'd.
t.
aa
:)\
!'1f
1.
:h-
:!'t
::
:R.
ir.
:5-
th
-6.
rn
\D
,15-
of the *'hite-crowned pigeon.
Wildlife Monogr. 64:
t-54.
Wrlr-soN. M. F. ,q.No
J. N. THoupsott. 1982. Phe-
nolog-v
and ecolog-v of color in bird-dispersed
lruits,
or wh1' some lruits are red when the-v are "grecn."
Canad.
J. Bot.60: 701-113.
Wrr-soN.
D. E. eNo D. H. Je.NzrN. 1972. Predation
on Scheelea
palm seeds b1' bruchid beetles: seed
densitl'and distance lrom the parcnt tree.
Ecolog-v
53:954-959.
WoNc. Y. K. 1959. Autecolog-v
of the
bertam
palm-
Eugeissona
lrl.ste,
Griff. Mala-van Forester 22: 301-
3 13.
WoorrnNorN. G. E. eNl J. w. FITZpATnrcrc. 1984.
The Florida scrub ja1. Princeton Univ. Press,
Princcton.406 pp.
WmrvcHev. R. W. rNo P. G. Werenvl.N. 1983.
Condensed tannins in fruits ealen b,v chimpanzces.
Biotrooica |
5: 217-222.
g!
rk
J's
ral
:rls
,al-
'p.
.f).
br
15.
:AI
tta
