ResearchGate

See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/248582071

Vertebrate predation by primates: A review of

hunting patterns and prey

Article Journal of Human Evolution - July 1982

DOI:10.lOl6/8004772484(82)800957X

CITATIONS READS
64 177

1 author:

Thomas M. Butynski
Lolldaiga Hills Research Programme, Sustainability Centre Eastern Africa
143 PUBLICATIONS 3,344 CITATIONS

SEE PROFILE

Some of the authors of this publication are also working on these related projects:

Distribution of the Common warthog (Phacochoerus africanus) and the Desert warthog (P. aethiopicus)
Project
in the Horn of Africa and N Kenya. View project

Project Eastern Africa Primate Conservation and Diversity Program View project

All content following this page was uploaded by Thomas M. Butynski on 12 November 2014.

The user has requested enhancement ofthe downloaded file.

 Call 3(1974w) DateReq: 6/11/2008 Yes

Number: Date Rec: 6/12/2008 A No

Location: Periodical Stacks Borrower: DXU E cond't'onal

ILL: 43486571 Maxcost: $501FM

Request Type: Source: ILLiad LenderString: *MMT,NJM,NJS,RID,ZXC

OCLC Number: 1783947 Afliation: PALCI
Staff Email: ill@drexel.edu (0 F3 6

Billing Notes: DXU prefers paying through IFM but will accept invoices:

Title: Journal of human evolution.

Uniform
Title:

Author:
Edition: Imprint: London, New York, Academic Press.

Article: Butynski, T.: Vertebrate predation by primates: a review of hunting patterns and prey.

Vol: 11 No.: Pages: 421-430 Date: 1982

Dissertation:

Veried: <TN:105757><ODYSSEY:129.25.131.206/DXU> OCLC 0047-2484

Borrowing Ariel: 129125131233

Notes:
ShipTo: Drexel University - Hagerty ILL/3141 Chestnut Street/Philadelphia, PA 19104-2875
E-delivery
Addr: (215) 8952542
Ship Via: IDS: 123/Lib Rate/Ariel/Odyssey/E-mail

ShipVia: IDS:123/Lib Ra NeedBy: 7/13/2008

Borrower: DXU

Return To: ILL: 43486571 Lender: mmt

Monmouth University Library Req Date: 6/11/2008 OCLC #: 1783947

Interlibrary Loan Dcpt' Patron: Butynski, Thomas
Cedar Avenue
w. Long Branch, NJ 07764 Author:
Title: Journal of human evolution.

Ship To; Article: Butynski, T.: Vertebrate predation by primates: a

review of hunting patterns and prey.
Drexel UnlverSIty - Hagerty ILL Vol.: 11 No.:
3141 Chestnut Street Date: 1982 Pages: 421-430
Philadelphia, PA 19104-2875 Veried: <TN:105757><0DYSSEY:129.25.131.206/
Maxcost: $501FM Due Date:
Lending Notes:
Bor Notes: Ariel: 129.25.131.233
z and Depression, pp. 237245.
Mother-infant separation in
Malay 12, 211217.
esus monkey infants. Animal
and conspecics other than
w, Eds.) Advance: in the Study
15 monkey infants from their
m mother on 1832 week old
313342.
) troops of hanuman langurs
Languren in Verschiedenen
ag Paul Parey.
1t Habitats. Proceedings of the
Review of Child Development
Thomas M. Butynski*
Kibale Forest Project,
P-O- Ba 409:
F0" Portal,
Uganda,_
East Afn
Received 6 August 1981 and
accepted 10 November 1981
Keywords: Meat-eating,
Predation, Diet, Hunting,
Scavenging, Prosimian, Simian,
Hominid.
Vertebrate Predation by Primates: a
Review of Hunting Patterns and Prey
Thirty-eight species of free-living non-human primates, representing .
nine families, have been observed to hunt and/or eat vertebrates.
Thirty-two per cent of these are prosimians, and 68% are arboreal
forest-dwellers. The predation pattern involving the search for, and
pursuit, capture and consumption of prey is rarely exhibited in non-
human primates other than chimpanzees and baboons. Scavenging
among non-human primates is uncommon. Only baboons and
chimpanzees are known to stalk their prey, to hunt co-operatively and
to share meat, and only chimpanzees have been observed to kill their
prey by ailing and to carry meat. Man is the only primate that
hunts animals larger than himself, and that incorporates tool-use,
complex vocal communication and bipedalism into his hunting
pattern. It is concluded that: (1) hunting of vertebrates is widespread
but infrequent among free-living nonhuman primates, (2) search for,
and pursuit, killing and eating of vertebrates was probably practised
by early primates, and (3) there are several similarities between the
hunting patterns of man and non-human primates but also several
important differences. .

11 variability in ecology and -_________

tes: Studies in Adaptation and

15). Primates 8, 127154. 1. Introduction

It has been hypothesized that man has been a hunter-gatherer for more than 99 % of
human history (Laughlin, 1968), and that meat has contributed a signicant portion to
the diets of at least some hominids during the last 2-5 million years (Fagan, 1974). If this
is true, the selective pressures associated with a hunting (Read, 1925; Dart, 1953) or
hunting-gathering existence (DeVore & Washburn, 1963; Laughlin, 1968; Washburn &
Lancaster, 1968; Pfei'er, 1969; Schaller & Lowther, 1969; Young, 1971; King, 1975,
1976; Ardrey, 1976) may have been a major force shaping human evolution and cultural
development.
Observations of baboons, Pepin spp. (Harding, 1975; Rose, 1978), chimpanzees, Pan
troglodytes (Teleki, 1975), and contemporary human hunter-gathercrs, however, suggest
that the hunting component of early hominid subsistence was not so much an innovation
as an extension of a basic primate pattern and associated subsistence behaviors. That is,
the earliest hominids were not the rst primates to exhibit hunting behavior.
Harding (1975) and Teleki (1975) reasoned that if hunting were not unique to human
evolution, but rather a continuation of a pattern already present in the ancestors of early
man, the likelihood that hunting behavior and hominization were coupled in their develop-
ment would be substantially reduced. In other words, fundamental developments in the
behavior of man may have evolved long before the appearance of Australopit/zecusbefore
man the hunter emerged.
Several workers involved in this debate (e.g. Dart, 1953; Teleki, 1975; King, 1976)
indicate that many non-human primates hunt vertebrates. Yet, surprisingly, there is no
review of the literature concerned with this behavior.

New York Zoological Society, The Rockefeller University, and

Makerere University.

Journal of Human Evolution (1982) 11, 421430

I 00472484/82/050421 + 10 $03.00/0 1982 Academic Press Inc. (London) Limited
422 'r. M. BUTYNSK!

- h u m a n Diets of several species c

s p r e d a t i o n o n ve rt eb ra te s b y fr ee -l iv in g n o n
This paper summarizes a n d di sc us se
en t d a t a Dominique, 1977) and in
en te r th e a b o v e d e b a t e b u t ra th er to pr es
primates. The intent he r e is n o t to
o n ve rt e- animal matter (Niemitz,
u e n c y a n d pa tt er n of p r i m a t e p r e d a t i o n
relevant to the debate d a t a o n f r e q
es en ta ti on percentages of animal rna
r o v i d e a n or ga ni za ti on al gu id el in e fo r pr
brates. The above two po in ts of v i e w p
h y p o - (Hladik 8: Hladik, 1969,
th e d a t a c a n b e in te gr at ed , a n d a so ur ce of
of the data, a backgr o u n d in to w h i c h 1978), and green monkey
theses against whi c h th e d a t a c a n b e ap pl ie d. Using a sample of 58 C'
from all parts of the worl
all latitudes derive at leas
2. D e n i t i o n s a n d M e t h o d s the median, and the mot
t ki ll ed ei th er b y th e c o n s u m e r or b y addition to meat from In
e d as th e ea ti ng of a n a n i m a l no
Scavenging is den ly pr ac ti se d b y n o n - these data, Lee postulatc
s so ci al gr ou p. B e c a u s e sc av en gi ng is ra re
a member of the cons um er from mammals. Lee (19
d th at ve rt eb ra te fe ed in g b y a n o n - h u m a n
, it w a s a s s u m e
human primates (see below) ve rt eb ra te re ma in s in If present-day hunter
ra te hu nt in g. I n se ve ra l ca se s, th er ef or e,
primate entailed verteb ve rt eb ra te s. D a t a o n matter, it appears that,
ec es w e r e us ed as ev id en ce fo r th e h u n t i n g of
stomach contents or fa diet. We have to look tc
f r o m ca pt iv e an im al s w e r e om it te d.
probable diets, or H u n t i n g pa tt er n a n d most resemble that of the
ar e us ed s y n o n y m o u s l y he re .
Hunting and predation l be ha vi or s di re ct ly to man in the extent to \
al so s y n o n y m o u s te rm s a n d e n c o m p a s s al
predation pattern are c o n s u m p t i o n of pr ey .
us iv e of , th e se ar ch fo r, p r o c u r e m e n t a n d
associated with, and incl th e hu nt . co op er at iv e
o m m u n i c a t i o n a n d di vi si on of la bo r du ri ng
These include vocal c a n d pr ey -s iz e se le ct io n.
n d w e a p o n us e, m e a t ca rr yi ng a n d sh ar in g, Vertebrate Matter in the Pr
hunting, bipedalism a
to th e e sh of ve rt eb ra te s.
Meat" refers only r th e c h a c m a b a b o o n Except for man, the 111.1]
fo ll ow in g in fa nt ic id e ha s b e e n re po rt ed fo
Although cannibalism od al l, 19 77 ; Su zu ki , spread, behavior (Table
n , 19 71 ), c h i m p a n z e e (B yg ot t, 19 72 ; Go
(Papio ursinur) (Saaym a mates were found for nir
a n d re dt ai l m o n k e y (C er co pi th ec ui ar ca ni us )
a) (F os se y, 19 81 )
1971), gorilla (Gorilla gorill di st in gu is he d f r o m th e prosimians, seven specie
r m of ki ll in g a n d me at -e at in g sh ou ld b e
(Struhsaker, 1977), this fo o n d u c t e d pr im ar il y to and three species of ape:
r sp ec ie s. T h e ki ll in g of ot he r sp ec ie s is c
killing and eating of othe p r e s u m a b l y u n d e r t a k e n Amphibians are eaten
e a s th e ki ll in g of co ns pe ci c in fa nt s is
obtain nutrients. wher m p r o v e d op po rt un it y to mals by 19, and reptiles l
o w n in cl us iv e t ne ss t h r o u g h i
primarily to increase ones g, th e ex pl oi ta ti on of th e Kortlandt 8: Kooij (196
el im in at io n of a co mp et it or s of fs pr in
reproduce and/or through 19 79 ). C a n n i b a l i s m is , and Mandrillus sp. also _
ce ap pa re nt ly be in g se co nd ar y (H rd y,
infant as a nutrient sour th is pa pe r. provide specic names f
nd of pr ed at io n a n d wi ll no t b e in cl ud ed in
therefore, a sp ec ia l ki feed.
Except for the feeding
are based upon one or 2
3. R e s u l t s a n d D i s c u s s i o n hunting of vertebrates is
here vertebrates appear
Animal Matter in the Primate Diet (Harding, 1975; Teleki,
u m a n p r i m a t e s h a v e m e m b e r s w h i c h h u n t
fa mi li es o f n o n - h hours that eld research
At least 10 of the 11 extant r i m a t e s f r o m 1 0 fa mi li es ,
r e v i e w o f th e di et s of 1 0 4 sp ec ie s of p of vertebrate hunting at
animals (Jones, 1972). In a ec ie s p r e y u p o n an im al s.
f o u n d th at 6 0 ( 5 8 % ) o f th e sp 800/0 of these are attril:
Gaulin & Konner (1977) ec ie s of p r i m a t e s f r o m ei gh t
(. 19 77 ) p r e s e n t f e e d i n g d a t a fo r 4 6 sp Hausfater, 1976).
Clutton-Brock & Harvey a l m a t t e r , a n d 2 2 (4 8 "/ 0)
) o f th es e sp ec ie s ea t at le as t s o m e a n i m That many primates
families. Thirty (65 % t t e r
a l m a t t e r . M e a n p e r c e n t a g e a n i m a l m a
species have diets consisting o f 5 3 8 5 % a n i m
t t e r most primates regularl
v e r y si mi la r m e a n p e r c e n t a g e a n i m a l m a
in th e di et s o f th es e 4 6 sp ec ie s is 1 2 % . A
- Harding (1975), Teleki
i m a t e s li st ed b y H a m i l t o n & B u s s e (1 97 8)
(13%) was calculated for the 21 sp ec ie s o f p r
of many of the differences
di et o f n o n - h u m a n p r i m a t e s co ns is ts
Almost all of the animal c o m p o n e n t o f th e are not of kind but rat]
invertebrate s ( H a m i l t o n & Bu ss e, 19 78 ).
 VERTEBRATE PREDATION BY PRIMATES 4-23

free-living non-human Diets of several species of prosimians consist of more than 70 9/0 animal matter (Charles-
ather to present data Dominique, 1977) and in Horselds tarsier, Tarsz'us bancanus, the diet is apparently 100"o
:6 predation on verte- animal matter (Niemitz, 1979). Excluding man, the simians with the highest reported
deline for presentation percentages of animal matter in their diets are Georoys tamarin, Saguinusgeorzyti (30 O/(,)
and a source of hypo- (Hladik & Hladik, 1969), talapoin monkey, Miapit/zecus talapoin (36%) (Gautier-Hion,
1978), and green monkey, Cercopit/zecw aethiopr (41%) (Galat & Galat-Luong, 1978).
Using a sample of 58 contemporary subsistence-based human hunter-gatherer societies
from all parts of the world, Lee (1968) found that with a single exception all societies at
all latitudes derive at least 20% of their diet from the hunting of mammals . . . the mean,
the median, and the mode for hunting all converge on a gure of 35 per cent. . . f in
by the consumer or by addition to meat from mammals, sh comprise a mean 01' 26% of the diet. Based upon
rely practised by non- these data, Lee postulated that the diets of prehistoric man consisted of 3040% meat
ling by a non-human from mammals. Lee (1968) did not consider the invertebrate portion of the diet.
vertebrate remains in If present-day hunter-gatherers reect early man in their consumption of animal
Evertebrates. Data on matter, it appears that, among simians, man has the highest animal component in his
diet. We have to look to the prosimians, animals whose behavior and ecology probably
iunting pattern and most resemble that of the rst primates (Nonnihan, 1976), before we nd primates similar
all behaviors directly to man in the extent to which they hunt and eat animals.
consumption of prey.
the hunt, cooperative
,nd prey-size selection.
Vertebrate .lIattt-r in the Primate Diet

)r the ehacma baboon Except for man, the hunting of vertebrates by primates is an uncommon, albeit wide-
oodall, 1977; Suzuki, spread, behavior (Tables 1 8L 2). References to vertebrate feeding by non-human pri-
(Cercopithecus arcanius) mates were found for nine families, 26 genera and 38 species. These include 12 species of
distinguished from the prosimians. seven species of New \IVorld monkeys, 16 species of Old World monkeys
:onducted primarily to and three species ofapes.
~esumably undertaken Amphibians are eaten by nine species of primates, birds (excluding eggs) by 18, main-
moved opportunity to mals by 19, and reptiles by 23. Based upon a questionnaire sent to reliable eld personnel,
the exploitation of the Kortlandt 8: Kooij (1963) report that Alouatta sp., Colnbus sp., Hapalcmw sp., .l/Iacaca sp.
979). Cannibalism is, and iVIandrillur sp. also prey upon vertebrates in the wild. Unfortunately, they do not
this paper. provide specic names for these primates nor the classes of vertebrates upon which they
feed.
Except for the feeding on mammals by baboons and chimpanzees, all scores in Table I
are based upon one or a few sightings. Among non-human primates it appears that the
hunting of vertebrates is most common among baboons and chimpanzees, although even
here vertebrates appear to comprise less than 1 13 of the diet of most groups studies to date
2 members which hunt (Harding, 1975; Teleki, 1975; Wrangham, 1977). During the many tens of thousands of
mates from 10 families, hours that eld researchers have observed nonhuman primates, fewer than +50 sightings
5 prey upon animals. of vertebrate hunting and/or feeding have been reported in the literature, and well over
of primates from eight 80%J of these are attributable to baboons (220) and chimpanzees (143) (Teleki, 1975;
matter, and 22 (48%) Hausfater, 1976). ' .
:entage animal matter That many primates occasionally prey upon vertebrates should not be surprising since
:entage animal matter most primates regularly search for, capture and eat invertebrates. As emphasized by
nilton & Busse (1978). Harding (1975), Teleki (1975), Harding & Strum (1976), and especially Rose (1978),
a primates consists of many of the differences between the subsistence patterns of non-human primates and man
are not of kind but rather of degree.
 T. M. BUTYNSKI
424

Table 1
Summary of th e da te on hu nt in g of ve rt eb ra te s by pr im at e.
1' Table 1 continued
M

ecies________ Amphibia
Reptilia ____________ es al
mm.
Ma_ _ a Re
i fe
re
nc
es
- il_y_a
F#a_m, _
ie_s_._--
n_d_s_pe_c.
and sp_
F mily _
:__ __ _
Papio anubix
Tupaidae Lait (1967)
Ptiloccrcus lowii DSouza (1976)
Tupaia minor
Lemuridae Hladik at al. (1980)
XXX

Chairogaleus medias Page: (1980)

Microcebus caquereli Hladik (1979); Papiu rynnccphalus
Micracebus murinus Martin (1972)
Papio papio
Lorisidae Charles-Dominique (1977) Papio ursinur
Galaga allmi Charles-Dominique
v

Galago crassicaudatus 8: Bearder (1979) Thempillzecux gelada

Charles-Dominique (1977) Hylobatidac
XXXX

Galago elegantulur Hladik (1979) Hylabates lar

Lari: tardigmdes Fooden (1971) Pongidac
Nycticebus caucang Charles-Dominique (1977) Pan panixcm
Pcmdicticur pom; Pan troglodytes
Tarsiidae Hladik (1979)
Tarsiur bamanus Nicmitz (1979
Callithricidae Coimbra-Filho &
Leontopithecus rosalia Mittermeier (1973)
Dawson (1976)
Saguinux oediput Neyman (1977)
van Horn (pers. comm.
in Dawson, 1976)

Cebidae Izawa (1978)

Cebu: apclla Oppenheimer (1968)
X

Cebu: capucinu: Durham (pers. comm.

m 31"
Table 2
I

_______._-
Lagathrix lagot/zrica in Teleki, 1975)
LeNestour (pers. comm.
Pithecia melanoczphala in Moynihan, 1976) Data on hunting
___________.___
Mittermeier (pers. comm.
Saimiri rciureus in Moynihan, 1976) % animal matter in the div.=
Number of species known t
vertebrates
.. Cercopithecidae Waser, (1977) "A, vertebrate matter (meat
/\
v

Cercocebus albigena Quris, (1975) ; Homewood Number of species known t

Cerwcebus galeritus (1976) mammals
Galat 8: Galat-Luong, /2, meat from mammals in
Cercopithecus aethiops (1978) Components of the hunting p
Gartlan 8L Brain (1968) Kill and eat vertebrates
Kavanagh, 1978 Search for and pursue vertl
McGrew el al., 1978 Practice scavenging; males
Struhsaker, 1967 than females
Gautier-Hion, 1978 Cooperative hunting; stalk
Cercopithecus cephus Butynski, in press share meat
Cercapithecu: mitir Mizuno at al. (1976) Kill prey by ailing; carry
Rudran, 1978 Kill large prey; complex v
Kingdon, 1971 communication; weapon
XXX

Ccrcapithecus neglectus Gautier-Hion (1973) bipedalism

Cercopithecus talupoin Hall, 1965 __________.- -
Erythrocebus palm Koster (pers. comm.)
Foodcn (1971)
XXX

Macaw assimmsis Sugiyama (1971)

Macaw radium Dittus (1974)
Macaw :t'nica
samw fez-3r
 VERIEBRATE PREDATION BY PRIMATES 425
:ebx-ates by primates
Table 1 continued Summary of the dam on hunting of vertebrates by primates
__________

"1.
References
Family and species Rept ilia Amp hibia Aves Mammalia References
M

- a):
Papia anubis X X , X DeVore & Washburn
: (1967)
(1963)
ouza (1976)
v-k.. _
Harding (1975)
Harding &. Strum (1976)
:lik at al. (1980) Kingdon (1971)
:s (1980) Rose (1977)
lik (1979); Rowell (1966)
r_

[artin (1972) Papio cynocephalus X X X X Altmann & Altmann (1970)

Hausfater (1976)
_

Papio papia X McGrew et al. (1978)

rles-Dominique (1977)

Papio ursinus x x x Dart (1963)

HesDominique
Hall (1966)
_ 7

Bearder (1979) Thempithecus gelada

ee-Dominique (1977) X Kummer (1968)
..

Hylobatidae
lik (1979) Hylabam lar
,_._.-4~'.._~ Yu_. A*~.f,_r:;_._\lv_

len ( 197 1) X Carpenter (1940)

Pongidae
dos-Dominique ( 1977) Pan panimzr X X Badrian & Badzian (1977)
Pan troglodytes X X Busse (1977)
m; (1979) Goodall (1968)
m (1979 Hladik (1977)
McGrcw et al. (1978, 1979)
Ibra-Filho & Nishida et al. (1979)
ittermeiet (1973) Plooij (1978)
son (1976) Suzuki (1971, 1973)
nan (1977) Wrangham (1977)
---_--.______
___
Horn (pers. comm.
Dawson, 1976) "' X indicates that this species is known to hunt this class of verte-
brate.
TThe data for Tam'u: banmnus were collected under seminatural
a (1978) conditions.
:nheimer (1968)
lam (pers. comm. Table 2 Summary of data on hunting by primates"
Teleki, 1975) --_________
atour (pers. comm.
Other
Moynihan, 1976) Data on hunting Prosimian simiansl' Baboon Chimpanzee Mani
:rmeier (pers. comm. M...
Moynihan, 1976) % animal matter in the diet 0-100 041 03 14 >61
Number of species known to hunt/eat
r, (1977) vertebrates 12 19 5 2
:, (1975); Homewood % vertebrate matter (meat) in the diet 0-5 0-5 1 l 61
76) Number of species known to hunt/eat
8: Galat-Luong, mammals 5 7 5 2 1
'78) % meat from mammals in the diet <0-5 <0-5 <1 <06 35
m & Brain (1968) Components of the hunting pattern
nagh, 1978 Kill and eat vertebrates
'ew et al., 1978 Search for and pursue vertebrates ? -
saker, 1967 Practice scavenging; males hunt more
er-Hion, 1978 than females
ski, in press Cooperative hunting; stalk prey;
no at al. (1976) share meat
m, 1978 Kill prey by ailing; carry meat ____..._______._>
Ion, 1971 Kill large prey; complex vocal
:r-Hion (1973) communication; weapon-use ;
1965 bipedalism ._ ._ _)
------.____._
3 (pers. comm.)
" References as in Table l.
n (1971)
tma (1971) 1' Includes simians other than baboons, chimpanzees and man.
(1974) I Contemporary hunter-gatherers.
Invertebrate portion not included.
426 'r. M. BUTYNSKI
clearly show that the hi
Forest Pri m a t e s a n d Ve rt eb ra te H u n t i n g activity. Data on this 5:
t e s a p p e a r s to b e m o s t f r e q u e n t a m o n g
x t a n t p r i m a t e s , t h e h u n t i n g o f v e r t e b r a
Am o n g e w e l l i n g a d a p t a t i o n s .
i m p a n z e e s a n d m a n , a n d m a y b e r e l a t e d t o g r o u n d - d Hunting Pattern qf Babom
b a b o o n s , c h e r t e b r a t e s , 2 6 ( 6 8 %
o n - h u m a n p r i m a t e s o b s e r v e d to h u n t v
However, of the 38 species of n a u s e a h i g h e r p r o p o r t i o n o f t h e Baboons (Harding, 19
at le as t 11 o f t h e s e h u n t m a m m a l s . B e c
are a r b o r e a l a n d
te ns iv el y t h a n t h e a r b o r e a l (Lawick-Goodall, 1968
h a v e b e e n st ud ie d, a n d s t u d i e d m o r e in
te rr es tr ia l p r i m a t e s d i f c u l t to (Washburn & Lancastc
o b s e r v a t i o n c o n d i t i o n s , th is n d i n g is
pri m a t e s , a n d b e c a u s e o f d i f f e r e n c e s in 1978), and share meat.
r e a d o c c u r r e n c e o f v e r t e b r a t e h u n t i n g
, i n d i c a t e a w i d e s p
interpret. It does, nonetheless s s u g g e s t s t h a t th is b e h a v i o r chimpanzees and even :
p r o s i m i a n s a n d s i m i a n s . T h i
among arboreal, forest-living n g b e f o r e t h e y b e c a m e te rr es - Chimpanzees (Lawic
r st p r i m a t e s a n d . th er ef or e, l o
pattern was present in the (Washburn & Lancast'

' vowre?-
trial or colonized savannas. prey by ailing it again
No primate other tha
Search f o r a n d Pu rs ui t q f Ve rt eb ra te Pr ef Chimpanzees and babo

L
1
h fo r, p u r s u i t , c a p t u r e a n d c o n s u m p t i o n 8: Altmann, 1970; Ha
e t e p r e d a t o r y p a t t e r n i n v o l v i n g s e a r c
T h e c o m p l a n d m a n is n o t w e l l upper limit (Teleki, 19'
t e s o t h e r t h a n b a b o o n s , c h i m p a n z e e s
of vertebrate prey by prima p u c h i n s ( C e b u : ap el la ) ri p o p e n size.
o c c u r . F o r e x a m p l e , b l a c k - c a p p e d c a
docu m e n t e d b u t d o e s er co pi th ec u: While chimpanzees
fr og s ( I z a w a , 1 9 7 8 ) ; v e r v e t m o n k e y s (C
s t e m s o f B a m b u s a q u a d u r in s e a r c h o f
the
6 7 ; K a v a n a g h , 1 9 7 8 ) a n d c h a s e , ki ll a n d chimpanzee was obsen
e a r c h t h e ne st s o f b i r d s ( S t r u h s a k e r , 1 9
aeth io ps ) s e r , 1 9 6 7 ) ; a b l u e m o n - This may be a case of a
i c k s (F ra nc ol in ix le uc os ce pu s) ( S t r u h s a k
eat yellownecked spurfowl ch

A
g a l a g o ( G a l a g n sp .) ( B u t y n s k i , i n pr es s) , see Eaton, 1978).
mi ti r) c h a s e d , ki ll ed a n d at e a

key (C er ca pi th ec us a n d f e d u p o n The hunting pattern
e y (E ry th ro ce bu s pa ta s) p u r s u e d , ki ll ed
o c c a s i o n s a p a t a s m o n k
ing complexity (Table i

M
and o n t w o
a st uc ga lm zs ix ) ( R o s t e r , pe rs . c o m m ) .
laughing d o v e s (S tr ep to pc li tern and associated bel

M
It seems that comple:

I
tial for primates huntir

Scavenging
e s w e r e l o c a t e d . T h e r e h a v e b e e n basic predation patterr

to s c a v e n g i n g b y n o n - h u m a n p r i m a t
Only si x r e f e r e n c e s s h b u c k
a n z e e s s c a v e n g i n g f r e s h l y k i l l e d b u bilities, the complete hc
r

r v e d a n d v e s u s p e c t e d c a s e s o f c h i m p
three o b s e o n e o b s e r v e d ( S h o r t e r , hunt and utilize large
r i s 8: . G o o d a l l , 1 9 7 7 ) ,

e l a p h u r sc ri pt us ) f r o m b a b o o n s ( M o r
(Tra g n b a b o o n s s c a v e n g e d ga ze ll es , primates and contemp<
t e r , 1 9 7 6 ) o c c a s i o n s w h e
WW

1981) an d t w o p r o b a b l e ( H a u s f a
c h e s ( W ' r a n g h a m , 1 9 7 7 ; O l i v e r , 1978), of these three compone
e a t i n g d e a d s h o n b e a
several instances of baboons r o m a h o u s e c a t ( S c h l i c h t e , consumption of meat 0:
u e m o n k e y s a p p r o p r i a ted a bird f
and one case in which bl years ago (Fagan, 1971
1978).
, 7"

t e d t h a t
t

9 ) a n d S z a l a y ( 1 9 7 5 ) h a v e s u g g e s
( 1 9 6 9 ) , S c h a l l e r & L o w t h e r ( 1 9 6
Pfeiff er a t t e r n m a r k i n g a sh if t f r o m
p a r t o f a n i n t e r m e d i a t e s u b s i s t e n c e p
scav e n g i n g f o r m e a t w a s
n t i n g i n e a r l y h o m i n i d s . Predation on vertebrat
plant-eating to prey-hu e x t a n t n o n - h u m a n p r i m a t e s . I f a
e n c e f o r a s c a v e n g i n g p h a s e a m o n g among non-human prii
There is n o e v i d t e s b y h o m i n i d s , it
r e q u e n t h u n t i n g o f v e r t e b r a
scaveng i n g p h a s e d i d p r e c e d e t h e sh if t t o f
D e V o r e & from the basic predato
v o l u t i o n . P e r h a p s , a s s u g g e s t e d b y
may h a v e b e e n a p h a s e u n i q u e i n p r i m a t e e
d t o u s e Hunting of vertebra!
t a n t t o m a n o n l y af te r h e l e a r n e
( 1 9 6 3 ) , s c a v e n g i n g b e c a m e i m p o r and overlap in pattern,
Washburn
weapons to drive car n i v o r e s f r o m t h e i r ki ll s. ism, complex communi
enable him to underta
Prehistoric man was
Division qf Labor important component
d a l l , 1 9 6 8 ; S u z u k i , 1 9 7 1 , 1 9 7 3 ;
, c h i m p a n z e e s ( L a w i c k - G o o
Data for m a n ( T e l e k i , 1 9 7 5 )
o n s ( A l t m a n n & A l t m a n n , tions not shared with
N i s h i d a et al ., 1 9 7 9 ) a n d b a b o
Teleki, 1975; M c G r e w , 1 9 7 9 ; 1 9 7 8 ) hominid ancestor.
H a u s f a t e r , 1 9 7 6 ; M c G r e w et al .,
1 9 7 5 ; H a r d i n g & S t r u m , 1 9 7 6 ;
1970; Hardi n g ,
 427
vanmmrs PREDATION BY rnmxrns

a m m a l s by th es e th re e sp ec ie s is pr ed om in an tl y a ma le
clearly show that th e hu nt in g of m
r ot he r sp ec ie s of pr im at es is sc an t.
activity. Data on this subject fo
most frequent among
dwelling adaptations. Hunting Patter n of Ba bo an s, Ch im pa nz ee : an d H u m a n :
vertebrates, 26 (68%) 19 76 ; Ha us fa te r, 19 76 ), ch im pa nz ee s
Baboons (H ar di ng , 19 75 ; Ha rd in g & St ru m,
gher proportion of the le ki , 19 75 ; M c G r e w , 19 79 ) an d m a n
(Lawick- Go od al l, 19 68 ; Su zu ki , 19 71 , 19 73 ; Te
y than the arboreal , hu nt co -o pe ra ti ve ly (b ut se e Bu ss e,
(Washburn &. La nc as te r, 19 68 ) al l st al k th ei r pr ey
s nding is di'icult to e in fr eq ue nt an d no t we ll de ve lo pe d in
1978), an d sh ar e me at . Th es e th re e be ha vi or s ar
of vertebrate hunting de ve lo pe d in ba bo on s.
chimpanzees and even more infreque nt an d po or ly
:sts that this behavior ki , 19 71 , 19 73 ; Te le ki , 19 75 ) and man
Chimpanzees (LawickGoodall, 1968; Su zu
:e they became terres-
- . v

th e on ly pr im at es k n o w n to oc ca si on al ly ki ll th ei r
(Washbur n 8t La nc as te r, 19 68 ) ar e
re .
... -, .

su rf ac e an d to ca rr y me at fo r di st an ce s of 1 k m or mo
prey by ailin g it ag ai ns t a ha rd
en ob se rv ed to pr ey up on an im al s la rg er th an it se lf .
; 3a, a; Rn-Kasfwgh : ,. r 4,; n j

No primat e ot he r th an m a n ha s be
ki ll wi ld m a m m a l s we ig hi ng mo re th an 6 kg (A lt ma nn
Chimpanz ee s an d ba bo on s se ld om
Te le ki , 19 75 ) an d 10 kg is ap pa re nt ly cl os e to th e
ture and consumption & Altmann, 19 70 ; Ha rd in g, 19 75 ;
ra st , m a n fr eq ue nt ly ki ll s m a m m a l s m a n y ti me s hi s
s and man is not well upper limit (T el ek i, 19 75 ). In co nt
; (Cebu: apella) rip open Size. ad ul t ma le
we re hu nt in g bu sh pi gs (P ot am or ho em : po ro us ), an
, monkeys (Cercopitheru: While ch im pa nz ee s
w a ro ck an d st ri ke an ad ul t bu sh pi g (P lo oi j, 19 78 ).
78) and chase, kill and chimpanzee wa s ob se rv ed to th ro
n- hu ma n pr im at e us in g aw ea po n wh il e hu nt in g (b ut
.er, 1967) ; a blue mon- This may be a ca se of a fr ee -l iv in g no
- u

,) (Butynski, in press), see Eaton, 1978). of in cr ea s-

ex ta nt pr im at es ca n be ar ra ng ed al on g a co nt in uu m
:1, killed and fed upon The hunting pa tt er n of
in uu m m a y re e ct th e ev ol ut io n of th e hu nt in g pa t-
ing complexity (T ab le 2) . Th is co nt
tern and as so ci at ed be ha vi or s in pr im at es .
un ic at io n, bi pe da li sm an d we ap on -u se ar e no t es se n-
It seems th at co mp le x vo ca l co mm
te s, in cl ud in g ma mm al s. Ne ve rt he le ss , w h e n th e
tial for primates hu nt in g sm al l ve rt eb ra
pr im at es is su pp le me nt ed wi th th es e un iq ue ca pa -
ated. There have been basic predation pa tt er n of no n- hu ma n
g pa tt er n em er ge s an d wi th it th e ab il it y to ef c ie nt ly
Freshly killed bushbuck bilities, the co mp le te ho mi ni d hu nt in
ab le 2) . Ev id en ce fo r me at -e at in g by no n- hu ma n
)ne observed (Shorter, hunt and ut il iz e la rg e m a m m a l s (T
er -g at he re rs in di ca te s th at , du ri ng th e ev ol ut io n
)ns scavenged gazelles, primates and cont em po ra ry h u m a n hu nt
ni d hu nt in g pa tt er n, a 30 to 35 -f ol d in cr ea se in th e
rn, 1977; Oliver, 1978), of these three co mp on en ts of th e ho mi
in cr ea se pe rh ap s al re ad y ev id en t mo re th an 2- 5 mi ll io n
1 house cat (Schlichte, consumption of me at oc cu rr ed a n
ye ar s a g o (F ag an , 19 74 ; Ki ng , 19 76 ).
3) have suggested that
4. S u m m a r y a n d C o n c l u s i o n s
rn marking a shift from
in g m a m m a l s , is a wi de sp re ad bu t in fr eq ue nt be ha vi or
Predation on ve rt eb ra te s, in cl ud
i-human primates. If a es . T h e co mp le x ho mi ni d hu nt in g pa tt er n pr ob ab ly de ve lo pe d
among non-h u m a n pr im at
ebrates by hominids, it y pa tt er n st il l ex hi bi te d b y m a n y ex ta nt n o n - h u m a n pr im at es .
from the ba si c pr ed at or
suggested by DeVore & te s by m a n a n d n o n - h u m a n pr im at es in di ca te s si gn i ca nt si mi la ri ty
Hunting of verteb ra
titer he learned to use li ke n o n - h u m a n pr im at es , ho we ve r, m a n in co rp or at es bi pe da l-
and overlap in pa tt er n, Un
o n us e in to hi s hu nt in g pa tt er n. Th es e ad ap ta ti on s
.; ism, complex co mm un ic at io n a n d w e a p
th e hu nt in g of ve rt eb ra te s as a m a j o r ac ti vi ty .
enab le h i m to un de rt ak e
w h o m hu nt in g b e c a m e a w a y of li fe a n d m e a t a n
Prehistori c m a n w a s a pr im at e fo r
rt ic ul ar pa tt er n of pr ed at io n co ns is te d of ad ap ta -
important compone n t of th e di et . H i s pa
.3; Suzuki, 1971, 1973; th a n y ex ta nt n o n - h u m a n pr im at e no r, pr es um ab ly , wi th a n y pr e-
(Altmatm & Altmann, tions not shared wi
i; McGrew at al., 1978) hominid ancestor.
428 T. M. BU'IYNSKI
Gaulin, S. J. C. & Konne
to w h i c h wi ll he lp us be tt er (R.J. Wurtman & J. J. V
qu es ti on s h a v e e m e r g e d , th e an sw er s
Two fundamental h u m a n be ha vi or . Gautier-Hion, A. (1973).
ca nc e of m a n th e hu nt er in th e ev ol ut io n of Gabonica 7, 295-391.
understand the sign i d
h of m a n s b e h a v i o r w a s d e v e l o p e d be fo re , a n Gautier-Hion, A. (1978). l
Why did man beco m e a hu nt er ? H o w m u c
Chivers &J. Herbert, Eds
how much duri ng , hi s hi st or y as a hu nt er ? Academic Press.
Goodall, J. (1977). Infant
259282.
bt ed to J a n Ka li na , M a r g i e Bl ak e, Ly sa
For their comm e n t s on th e ma nu sc ri pt I a m in de Hall, K. R. L. (1965). Beh:
n d Dr s G e o r g e Pe tr id es , T h o m a s St ru hs ak er , Journal of Zoology, London 1
Leland, Stanley Koster, Jo se ph S k o r u p a a
D r Pe te r M a r l e r ki nd ly pr o- Hall, K. R. L. (1966). Dis
M c G r e w , a n d Ph il ip T h o m p s o n .
Michael Rose, William iv er si ty , w h e r e th is London 17, 4973.
Fi el d R e s e a r c h Ce nt er , T h e Ro ck ef el le r Un Hamilton, W. J. III & Bu:
vided facilities at Th e
d g e th e n an ci al su pp or t of th e N e w Y o r k Bioscienoe 28, 761766.
paper was written. I gr at ef ul ly a c k n o w l e
Harding, R. S. O. (1975). 1
Zoological Society. Psychology of Primates, pp. 1
Harding, R. S. O. & Strum,
85 (3), 46-53.
Hausfater, G. (1976). Fred:
References Hladik, A. & Hladik, C. M
of Barro Colorado (Panama
Fi el d Re se ar ch . Ch ic ag o: Un iv er si ty
n, J. (1 97 0) . Ba bo on Ec ol og y: Af ri ca n Hladik, C. M. (1977). Chix
Altmann, S. A. & Altman
the diet. In (T. H. Clu1
Chicago Press. um .
in g Hy po th es is . N e w Yo rk ; At he ne Lemurs, Monkey: and Apes,
Ardrey, R. (1976). The Hunt 13 , 46 3 46 8.
A. & Ha dr ia n N. (1 97 7) . P y g m y ch im pa nz ee s. Or yx
k e y Hladik, C. M. (1979). Die
Badrian, in th e ev ol ut io n of re d co lo bu s m o n
e pr ed at io n as a po ss ib le fa ct or Study of Prosimian Behavior
Busse, C. D. (1977). Chimpanze
ga ni za ti on . Ev ol ut io n 31 , 90 7 91 1. te s; Hladik, C. M., CharlesDar
social or ni ) pr ed at io n on ga la go s. (P ri ma
n k e y (e er co pi th ec us mi ti s st uh lm an ians in dry forest of the i
Butynski, T. M. (in press). Blue mo re (L on do n) 23 8, 41 0 41 1.
nn ib al is m a m o n g wi ld ch im pa nz ee s. Na tu Malagasy Primates: Ecology
Bygott, J. D. (1972). Ca d so ci al re la ti on s of th e gi bb on (H yl o-
ld st ud y in S i a m of th e be ha vi or a n Homewood, K. M. (1976).
Carpenter, C. R. (1940). A e
ar at iv e Ps yc ho lo gy Mo no gr ap hs 16 , 1 21 2. Dissertation. London: I
hater lar). Comp at es . N e w Yo rk : C o l u m b i a Un iv er -
Ec ol og y an d Be ha vi ou r of No ct ur na l Pr im Hrdy, S. B. (1979). Infanti<
Charles-Dominique, P. (1977).
ic al tions for the reproductive
sity Press. Me th od ol og
fl:1
s o n lo ri si d be ha vi or :
r, S. K . (1 97 9) . Fi el d st ud ie Izawa, K. (1978). Froge:
CharlesDominique, P. & Boarde e St ud y of Pr ox im ia n Be ha vi or , pp . 56 7 62 9. N e w
rt in , Ed s) , Th 633642.
aspects. In (G. A. Doyle 8: R. D. Ma
Jones, C. (1972). Natural (
York: Academic Press. in fe ed in g a n d ra ng in g be ha vi ou r in
, P. H . (1 97 7) . Sp ec ie s di ff er en ce s Simian Primates. Basel: S.
Clutton-Brock, T. H. & Harvey ie s of Fe ed in g an d Ra ng in g Be ha vi ou r in
Ed .) , Pr im at e Ec ol og y: St ud Kavanagh, M. (1978). Th:
primates. In (T. H. Clutton-Brock, a d e m i c Pr es s.
58 4. N e w Y o r k : A c 30, 3063.
Lemon, Monkeys and Apes, pp. 557 n d ec ol og y of th e g e n u s Le on to pi th ec u:
er me ie r, R . A. (1 97 3) . Di st ri bu ti on a King, G. E. (1975). Sociott
Coimbra-Filho, A. F. & Mitt
Br az il . Pr im at es 14 , 47 6 6. Research 31, 6987.
Lesson, 1840 in at io na l An th ro po lo gi ca l an d Li ng ui st ic
y tr an si ti on f r o m a p e to m a n . In te rn King, G. E. (1976). Societj
Dart, R. A. (1953). The predator
Kingdon, J. (1971). East A
Review 1, 201208. bo on s. S y m p o s i u m of th e Zo ol og ic al So ci et y q u o n d o n 10 ,
ou s pr op en si ty of ba Press.
Dart, R. A. (1963). The carnivor
ic hi da e, Kortlandt, A. & Kooij, NI.
4956. n, Sa gu in u: oe di pu : (C al li tr
ec ol og y of th e Pa na ma ni an ta ma ri Symposium of the Zoological
Dawson, G. A. (1976). Behavioral ty .
es ). Di ss er ta ti on . Ea st La ns in g: M i c h i g a n St at e Un iv er si
o w e l l 8: Kummer, H. (1968). 5001';
Pr im at a n d h u m a n ev ol ut io n. I n (F . C. H
W a s h b u r n , S. L. (1 96 3) . B a b o o n ec ol og y Chicago Press.
De V o r e , I. 8: Ch ic ag o: Al di ne .
s) , Af ri ca n Ec ol og y an d H u m a n Ev ol ut io n, pp . 33 5 36 7.
on . Lait, L. (1967). Notes on
F. Bourliere, Ed m o n k e y , M a c a o a si ni ca . Di ss er ta ti
P. J. (1 97 4) . T h e ec ol og y a n d be ha vi or of th e to qu e gymnumr (Rafes) (moon
Ditt us , W .
Pa rk : Un iv er si ty of M a r y l a n d . (R . D . Journal of Zoology, London
Universi ty s h r e w Tu pa ia mi no r. I n
ar y fi el d re po rt of th e le ss er tr ee Laughlin, W. S. (1968). H
DSouza, F. (1976). A prelimin , Pr os im ia n Be ha vi ou r, pp . 16 7 18 2. L o n d o n : D u c k w o r t h .
In (R. B. Lee & I. Dth
e r , Ed s)
Martin, G. A. Doyle & A. C. Walk w e a p o n r y ? Ca rn iv or e 1, 10 3 10 6.
z e e s us e w e a p o n s or mi ni c h o m i n i d Lee, R. B. (1968). What hu
Eaten, R. L. (1978). Do chimpan
Ea rt h. Bo st on : Li tt le , B r o w n . 1. DeVore, Eds), Man th.
Fagan, B. (1974). Men of the n :1 . J a n u a r y A p r i l , 19 67 . Fi el di an o
at es co ll ec te d in we st er n T h a i l a McGrew, W. C. (1979). 1
Fooden, J. (1971). Report on prim , tool use. In (D. A. H:
i4 1' 'rVrg.

Zoology 59, 162. ra ph ic 15 9, 50 0 52 3.

h e im pe ri le d m o u n t a i n go ri ll a. Na ti on al Ge og Benjamin/Cummings.
Fossey, D. (1981). T n m o n k e y s in Se ne ga l. In (D . J. Ch iv er s & J. H e r b e r t ,
Martin, R. D. (1972). Ade
& Ga la t- Lu on g, A. (1 97 8) . Di et of gr ee s.
Galat, G . N e w Y o r k : A c a d e m i c Pr es
ol og y. Vo l. I (B eh av io ur ), pp . 25 7 25 8. ions of the Royal Society of
Eds), Recent Advances in Primat in Ce re op it he cu s ae th io p: a n d C. mi ti x.
. E c o l o g y a n d so ci al va ri ab il it y McGrew, W. C., Tutin, C.
Gartlan, J. S. & Brain, C. K. (1968) ti on an d Va ri ab il it y, pp . 25 8- 29 2. N e w Y o r k : Ho lt ,
upon vertebrates: New r
St ud ie s in Ad ap ta
In (P. C. Jay, Ed.), Primates:
to n. , .
Rinehart and Wins

ch will help us better
0 of human behavior.
developed before, and
13., Margie Blake, Lysa
;, Thomas Struhsaker,
er Marler kindly pro-
University, where this
tort of the New York
h. Chicago: University of
ion of red colobus monkey
0. gm... gp.......;~, t1
m) 238, 410411.
ations of the gibbon (Hylo
v York: Columbia Univer-
behavior: Methodological
ehovior, pp. 567629. New
; and ranging behaviour in
ing and Ranging Behaviour in
,1 of the genus Leontopithecux
Anthropological and Linguistic
laologicol Society quondon 10,
tinus oedipur (Callitrichidae,
.tion. In (F. C. Howell &
.go: Aldine.
Wacaca :inica. Dissertation.
Tupaia minor. In (R. D.
182. London: Duckworth.
)nry? Carnivore 1, 103106.
naryApril, 1967. Fieldiona
, 500-523.
D. J. Chivers & J. Herbert,
I York: Academic Press.
pithecus oethiops and C. mitis.
53292. New York: Holt,
1
VERTEBRATE PREDATION BY PRIMATES 4-29
Gaulin, S. J. C. & Konner, 1V1; (1977). On the natural diet of primates, including humans. In
(R.J. Wurtman & J. J. Wurtman, Eds), Nutrition and the Brain, Vol. I, pp. 186.
Gautier-Hion, A. (1973). Repertoire comportemental du talapoin (Miopithecus talapoin. Biologica
1.,r Gabonica 7, 295391.
Gautier-Hion, A. (1978). Food niches and coexistence in sympatric primates in Gabon. In (D. J.
Chivers & J. Herbert, Eds), Recent Advances in Primatology. Vol. I (Behaviour), pp. 269286. New York:
Academic Press. ,
Goodall, J. (1977). Infant killing and cannibalism in free-living chimpanzees. Folio primotalogico 28,
f 259282.
Hall, K. R. L. (1965). Behaviour and ecology of the wild patas monkey, Erythrocebus patas, in Uganda.
Journal of Zoology, London 148, 1587.
Hall, K. R. L. (1966). Distribution and adaptation of baboons. Symposium of the Zoological Society of
London 17, 4973.
Hamilton, W. J. III & Busse, C. D. (1978). Primate carnivory and its signicance to human diets.
Biorcience 28, 761-766.
Harding, R. S. O. (1975). Meat eating and hunting in baboons. In (R. H. Tuttle, Ed.). Socioccology and
.
g Psychology of Primates, pp. 245257. The Hague: Mouton.
Harding, R. S. O. & Strum, S. C. (1976). The predatory baboons of Kekopey. Natural History (New York)
85 (3) , 4653.
Hausfater, G. (1976). Predatory behavior of yellow baboons. Behaviour 56, 4468,
Hladik, A. & Hladik, C. M. (1969). Rapports trophiques entre vegetation et primates dans la fort de
Barro Colorado (Panama). La Terre et la Vie 23, 25117.
Hladik, C. M. (1977). Chimpanzecs of Gabon and chimpanzees of Gombe: Some comparative data on
l the diet. In (T. H. Clutton-Brock, Ed), Primate Ecology: Studies of Feeding and Ranging Behaviour in
Lemurs, Monkeys and Apes, pp. 481501. New York: Academic Press.
Hladik, C. M. (1979). Diet and ecology of prosimians. In (G. A. Doyle & R. D. Martin, Eds), The
Study of Prosimion Behavior, pp. 307357. New York: Academic Press.
Hladik, C. M., CharlesDominique, P. & Petter,J. J. (1980). Feeding strategies ofve nocturnal prosim-
ians in dry forest of the west coast of Madagascar. In (P. Charles-Dominique et al., Eds), Nocturnal
A/lalagary Primates: Ecology, Physiology and Behavior, pp. 4173. New York: Academic Press.
Homewood, K. M. (1976). Ecology and behaviour of the Tana mangabey Cercoccbu: galm'tus galeritus.
Dissertation. London: University of London.
Hrdy, S. B. (1979). Infanticide among animals: A review, classication, and examination of the implica-
tions for the reproductive strategies of females. Ethology and Sociobiology 1, 1340.
'1 Izawa, K. (1978). Frog-eating behavior of wild black-capped capuchin (Cebu; apella). Primates 19,
633642.
Jones, C. (1972). Natural diets of wild primates. In (R. N. T-W-Fiennes, Ed.), pp. 5877. Pathology of
Simian Primates. Basel: S. Karger.
Kavanagh, M. (1978). The diet and feeding behaviour of Cercopithecw aethiops tantalus. Folio primotologico
30, 3063.
King, G. E. (1975). Socioterritorial units among carnivores and early hominids. Journal ofAnthrapologicol
Research 31, 6987.
King, G. E. (1976). Society and territory in human evolution. Journal of Human Evolution 5, 323332.
f Kingdom, J. (1971). East African Mammals: An Atlas of Evolution in Africa, Vol. I. New York: Academic
Press.
Kortlandt, A. & Kooij, M. (1963). Protohominid behavior in primates (preliminary communication).
it Symposium of the Zoological Society of London 10, 6188.
I Kummer, H. (1968). Social Organization of Homodryo: Balloons: A Field Study. Chicago: University of
Chicago Press.
1. Lait, L. (1967). Notes on the food habits of Ptilocercu: lowii Gray (pentail tree-shrew) and Echinororex
gymnurus (Rafes) (moonrat) in Malaya with remarks on ecological labelling" by parasite patterns.
Journal of Zoology, London 152, 375379.
Laughlin, W. S. (1968). Hunting: An integrating biobehavior system and its evolutionary importance.
l In (R. B. Lee & I. DeVore, Eds), Man the Hunter, pp. 304-820. Chicago: Aldine.
Lee, R. B. (1968). What hunters do for a living, or, how to make out on scarce resources. In (R. B. Lee &
I. DeVore, Eds), Man the Hunter, pp. 3048. Chicago: Aldine.
McGrew, W. C. (1979). Evolutionary implications of sex differences in chimpanzee predation and
tool use. In (D. A. Hamburg & E. R. McCown, Eds), The Great Apex, pp. 441463. London:
Benjamin/Cummings.
Martin, R. D. (1972). Adaptive radiation and behaviour of the Malagasy lemurs. Philosophical Transact-
l ions of the Royal Society of London (B. Biological/Sciences) 264, 295-352.
McGrew, W. C., Tutin, C. E. G., Baldwin, P.J., Sharman, M.J. & Whiten, A. (1978). Primates preying
upon vertebrates: New records from West Africa. Carnivore l (3), 4-145.
 430 'r. it. 13mm

McGrew, W. C., Tutin, C. E. G. & Baldwin, P. J. (1979). New data on meat eating by wild chimpanzees. C. B. Stringer
Current Anthropology 20, 238239.
Mizuno, A., Kawai, M. & Ando, S. (1976). Ecological studies of forest-living monkeys in the Kibale Department of Palaeontology,
Forest of Uganda. Kyoto University African Studies 10, 135. British Museum (Natural Hi.
Morris, K. & Goodall,J. (1977). Competition for meat between chimpanzees and baboons of the Gombe Cromwell Road,
National Park. Folia primatologioa 28, 109-121. London SW7 58D,
Moynihan, M. (1976). The New World Primates. Princeton: Princeton University Press. England
Neyman, P. F. (1977). Aspects of the ecology and social organization of free-ranging cotton-top tamarins
(Saguinus oedipus) and the conservation status of the species. In (D. G. Kleiman, Ed.), The Biology and Received 24 May 1981 am
Conservation of the Callitriehidae, pp. 39-71. Washington, D.C.: Smithsonian Institution Press. accepted 22 December 198
Niemitz, D. (1980). Outline on the behavior of Torsius baneanus. In (G. A. Doyle 8; R. D. Martin, Eds),
The Study of Prosimian Behavior. New York: Academic Press. Keywords: Neanderthal, Fo
Nishida, T., Uehara, S. & Nyundo, R. (1979). Predatory behavior among wild chimpanzees ofthe Mahale Hominid, Evolution,
Mountains. Primates 20, 120. Saint-Csaire.
Oliver, J. (1978). Feeding behaviour of olive baboons, Papio anubis, at Gombe National Park, Tanzania.
Dissertation. Cambridge: Cambridge University.
Oppenheimer, J. R. (1968). Behavior and ecology of the white-faced monkey, Cebus eapucinus, on llztrro
Colorado Island, (2.2. Dissertation. Chicago: University of Illinois.
Pages, E. (1980). Ethoecology of illicroeebus eoquereli during the dry season. In (1. Charles-Dominique
et al., Eds), Nocturnal Malagasy Primates: Ecology, Physiology and Behavior, pp. 971 )6. New York:
Academic Press. The discovery of a N4
Pfei'er (1969). The Emergence ofMan. New York: Harper and Row.
Plooij, F. X. (1978). Tool-use during chimpanzees bushpig hunt. Cannon" 1, 103-106.
Saint-(Isairc, France
Quris, R. (1975). Ecologie ct organisation sociale tle Cereoeebus galeritus ngilis dan 1e nerd-est (lu Gabon. journal Nature (ApSimt
La Terre et la Vic 29, 337-398. is unable to devote furt
Read,.C. (1925). The Origin qfhlan (2nd edition). London: Cambridge University Press.
Rose, M. D. (1977). Positional behaviour of olive baboons (Papio anuhix) and its relationship to mainten-
(Jsairc discovery, but
ance and social activities. Primates 18, 59-116. to it (1981) are too in
Rose, M. D. (1978). The roots of primate predatory behavior. Journal of Human Evolution 7, 179189. discussion below will c
Rowcll, T. E. (1966). Forest living baboons in Uganda. Journal Q/Zoolog). London 149, 344-364.
Rudran, R. (1978). Socioccology of the blue monkeys (Cereopithecus mitis sluhlrnann of the Kibalr Forest.
archaeological evidence
Uganda. Smithsonian Contributions to Zoology No. 249. Washington. D.C.: Smithsonian Institution Press. R. Kruszynski and mys-
Saayman, G. S. (1971). Behaviour of the adult males in at troop 01 free-ranging chacma baboons (Pupil) At all times it must be
ursinus). Folio primatologico 15, 36-57.
Schaller, G. B. & Lowther, G. R. (1969). The relevance of Carnivore behavior to 1110 study of early
are necessarily prelimin
hominids. Southwestern Journal of Anthropology 25, 307341. by the French workers
Schlichte, H. (1978). The ecology of two groups of blue monkeys. Certopithecus milis stuhlmanni. in an The Neanderthals ar
isolated habitat of poor vegetation. In (G. G. Montgomery. Ed). The Ecology olerboreal Folit'ores,
pp. 505517. Washington, D.C.: Smithsonian Institution Press. they may be the closest
Shorter, C. (1981). Samburu-Bu'alo Springs Game Reserve. Swara 4, 20. Neanderthal problcm
Struhsaker, T. T. (1967). Ecology of vervet monkeys (Cercopithecus arlhiops) in the Masai Ambosrli game Neanderthals to model
reserve, Kenya. Ecology 48, 891-904.
Struhsaker, T. T. (1977). lnfanticide and social organization in the redtail monkey (Crroopithecus asconius 1981), there would seem
schmidti) in the Kibale Forest. Uganda. Zeitschriftfu'r Tierprrchologir 45, 7584. hominid evolution. Th
" Sugiyama, Y. (1971). Characteristics of the social life of bonnet macaques (Macaw radium). Primates evolutionary studies sin
12, 247266.
Suzuki, A. (1971). Carnivority and cannibalism observed among forest-living chimpanzees. Journal oft/1e and mode of hominid ('
Anthropological Society of Nippon 79, 3048. et al., 1981).
Suzuki, A. (1975). The origin ofhominid hunting: A primatological perspective. In (R. H. Tuttle, Ed.), The manner in whic.
Socioecology and Psychology of Primates, pp. 259278. The Hague: Mouton.
Szalay, F. S. (1975). Hunting-scavenging protohominids: A model for hominid origins. [Man 10, 420-429. deftly incorporated into
Teleki, G. (1975). Primate subsistence patterns: Collector-predators and gatherer-hunters. Journal of evolution (Wolpo', 198
Human Evolution 4, 125184. to the Neanderthal prc
van Lawick-Goodall,J. (1968). The behaviour of free-living chimpanzees in the Gombe Stream Reserve.
Animal Behaviour Monographs 1, 16131 1. more critical discove
Waser, P. (1977). Feeding, ranging and group size in the mangabey Cereocebus albigeua. In (T. H. Glutton- some of the recent work
Brock, Ed.), Primate Ecology: Studies of Feeding and Ranging Behaviour in Lemurs, [Monitors and Apes, pp. 183- a different approach by
222. New York: Academic Press.
Washburn, S. L. & Lancaster, J. B. (1968). The evolution of hunting. In (R. B. Lee & I. DcVore, Eds), Wolpo (1981) now :
' Man the Hunter, pp. 293803. Chicago: Aldine.
Wrang ham, R. W. (1977) . Feedi ng behav iour of chimp anzee s in Gomb e Nation al Park, Tanza nia. In
(T. H. Clutton-Brock, Ed.), Primate Ecology: Studies of Feeding and Ranging Behaviour in Lemurs, Alonkejs 00472484/82l050431 + 08
and Apes Pp. 504538. New York: Academic Press.
Young, J. z. (1971). An Introduction to the Study of Man. Oxford: Clarendon Press.