A cross-species analysis of carnivore, primate, and hominid behaviour

Philip

A Cross-species Analysis of Carnivore, Primate, and Hominid Behaviour

R. Thompson*

Department of Psychology, Uniocrsi~ of Arizona, Tucson, Arizona, U.S.A. Received 31 December 1973 and accepted 14 September 1974

The traditional assumption that the origin of human behavior is found within the higher primates rather than the social carnivores is based on failure to adequately define primate and carnivore behavior. Phyletic classification takes no account of convergence and divergence; the behavior of a species is not necessarily characteristic of its order. Of 8 behavior variables that distinguish the order primates from the order carnivora, preagricultural man resembles the carnivores on 7 items: food sharing, food storing, cannibalism, surplus killing, interspecies intolerance, feeding of young, and division of labor; resembling the order primates only in group defense. The original form ofmuch human behavior is found within the carnivores.

1. Introduction In attempting

to deduce

have frequently

the origin

offered

for the relevance

hunter

(Dart,

1953;

partial

species.

answer.

carnivore parisons

behavior

is behavior

this behavior

behavior

characteristic

of species

and have acquired central

However,

and hominids

their similar behavior

1969).

behavior

the order

of a

are only a

the order primates;

carnivora.

For example,

Teleki

When

com-

(1973)

com-

with human hunting as evidence for deriving the predatory

behavior

but is in fact

are both converging through

with carnivore

long,

as a result of divergence

of chimpanzees

deviant.

convergence

is not

A more

with terrestrial evidence

likely

carnivores

on a common

thesis of this study is that there is incontrovertible

of human

as a

approaches

and the behavior

which characterizes result.

has also been evolution

Both

however

can vary drastically

in the order primates,

theory is that chimpanzees

vergence

of chimpanzees

from the primates.

species,

characterizes

anthropologists

of common

human behavior

is behavior

which

because

between

species behavior behavior

behavior,

Some evidence

& Lowther,

between

are made at a specific level ambiguities

pares the predatory

The

Schaller

1961;

lists of similarities

Primate

of human for clues.

carnivores

lists of similarities

But

Individual

and convergence.

primates

of terrestrial

Ardrey,

have usually provided particular

and evolution

looked to the nonhuman

niche. of the con-

behavior.

2. Methodology The

approach

as to which

outlined

posed in such a manner variable

is presumed

throughout primates.

above uses the following

of two paired

behaviors

that one of two variabIes

to be independent,

this study

procedures.

is characteristic

is taxonomic

Since the dependent

is presumed

one dependent. inclusion

behavior

variable

A hypothesis

of carnivores.

within

Each

to be causal;

The independent the order

is proposed hypothesis

carnivora

is also a dichotomy,

is

i.e. one

variable and

used order

usually A and .&

the reIationship of the two variables is expressed in a 2 x 2 contingency table. A hypothesis is considered confirmed if more carnivore species exhibit A than A and more * The author was a student at the University of Arizona during the academic years 1972-74. Requests for reprints shouldbe sent to Professor Wilbur R. Thompson, Department of Economics, Wayne State University, Detroit, Michigan 48202, U.S.A. Journal of Human Evolution (1975) 4,113-124

114

P.

R.

THOMPSON

primate species exhibit A than A and the probability that this relationship could have occurred by chance alone is less than 0.05. Because more primate and carnivore species are known biologically than behaviorally, it is not possible to select a random sample of species within these orders for study. Instead, a representative sample of primate and carnivore species was selected. Table 1 gives these species with their references. There is a noticeable bias in the sample toward higher species in the orders. This is because the study deals with convergent and parallel Table

1

Sample species and sources

Onnan Primates FAMILY

Cebidae

Alouattapalliata *** howler (Carpenter, 1965; Chiven, 1969) Callicebus moloch *** titi (Mason, 1968) Saimiri oersfedi *** squirrel monkey (Baldwin & Baldwin, 1972) FAMILY

Cercopithecidae

CercojnWwusaethiops * +* vervet (Hall & Gartlan, 1964; Gartlan & Brain, 1968; Struhsaker, 1967) Erythrocebuspatas *** patas (Hall, 1968) Macacu mulatta *** rhesus (Hausfater, 1972; Koford, 1963, 1965, 1966; Lindberg, 1971; Southwick, Beg & Siddiqi, 1965) Mucaca radiata l ** bonnet macaque (Simonds, 1965) Papio anubis * * * Savannah baboon (DeVore & Hall, 1965; Hall & DeVore, 1965) Papio hamadryas *** desert baboon (Kummer, 1968a,b) Presbytis entellus *** langur (Jay, 1965; Yoshiba, 1968) FAMILY

Colobidae

Colobus guereta * * * (Marker, 1969) FAMILY

Hylobatidae

Hylobates lar *** white handed gibbon (Be&son, FAMILY

Carpenter,

1940;

Ellefson,

1968)

Pongidae

Gorilla gorilla (Schaller, 1965) Pan troglody6es*** chimpanzee FAMILY

1971;

(Goodall,

1965;

Reynolds & Reynolds,

1965;

Teleki,

1973)

Horninidae

Homo sapiens (Birdsell, 1967)

1968, 1972;

Dart,

1953, 1959;

Davie,

1929;

Lee 8r DeVore,

1968;

Martin,

oxnxx Carnivora FAMILY

Canidae

Canis lup~r *** wolf (Cowan, 1947; Jordan, Shelton & Allen, 1967; Mech, 1970; Murie, 1944; Pimlott, Shannon & Kolenosky, 1969; Wolfe & Allen, 1973; Young, 1944) Canis aureus +** golden jackal (Schaller, 1972; van Lawick-Goodall, 1970) Lycaon pitus * * * wild dog (Ester & Goddard, 1967; Kuhme, 1965; Schaller, 1972) Vulpesfulva *** red fox (Burrows, 1968; Kruuk, 1966; Tinbergen & Ennion, 1967; Vincent, 1958) FAMILY

Hyaenidae

Crocuta crocuta *** FAMILY

spotted hyena (Kruuk,

1972;

Schaller,

1972)

Felidae

Acinonyxjubatus l ** cheetah (Abbott & Fuller, 1971; Schaller, 1972) Fe& concolor*** puma (Homocker, 1969, 1970; Young, 1946) Panthera lo *** lion (S&&r, 1972) Pantherupardus *** leopard (E&&erg, 1970; Lydekker, 18957; Schaller, Panthera tigris *** tiger (Lydekka, 1895?; Schaller, 1967) Lynx rufw l ** bobcat (Bailey, 1972; Young, 1958)

1972; Turnbull-Kemp,

Scientific names are italicized, sources are in brackets.

1967)

CARNIVORE,

PRIMATE

AND

HOMINID

BEHAVIOR

115

Figure 1. Summary of Tertiary fossil cebids. (Provincial ages after Patterson& PascuaI,1968). EPOCHS

PROVINCIAL AGES

FOSSIL FORhIS Neosaimiri fieldsi

Pliocene

Miocene -Pitheculus nustralis

Oligocene Eocene

Dolichocebus gaimansis

Paleocene

Branisella boliviana

behaviors and these are more likely to be associated with advanced species than primitive ones. This bias limits the universe to which the findingj may be legitimately generalized, which becomes the primate families Cebidae, Cercopithecidae, Hylobatidae, and Pongidae, and the carnivore families Canidae, Hyaenidae, and Felidae. The family Colobidae is probably not adequately represented by its one species, Colobus guereza. The variables selected were ones that were mentioned in the literature dealing with naturalistic behavior of carnivores and primates, were available in the field studies of the sample species, and could be formulated in a manner suitable for dichotomous classification. Since the statistics used in this study operate most efficiently when variables are dichotomized close to the median, a characteristic was also required to be present in at least 20 % of the species in a 2 x 2 table. 3. Carnivores Abstraction

and Primates

at the First Level of

Diet

Meat is a high calorie food relative to vegetal matter, requiring less bulk consumption. Meat eaters often share food among themselves; herbivores do not, since the calorie content is so low the effort is scarcely justified. Even in carnivores which exist as solitary adults, except for brief periods of mating, food is shared between a mother and her young. Moreover, a carnivorous diet requires a highly developed weapon system to hold and kilI prey. A carnivore must capture his food by careful stalk or long chase; the food of a herbivore cannot escape. Sharing of food acquired only with difficulty could be instrumental in maintaining sick or immature animals in carnivora, much less so in primates. Given the practice needed for successful capture of prey, it is doubtful that the young of predators could survive if they were not allowed to feed from the kills of aduits well after being weaned. With the exception of the chimpanzee and Savannah baboon, which on occasions kill and eat smaller mammals, the primates are noncarnivorous and even in the chimpanzee and baboons meat eating represents only an infrequent supplement to a basic vegetarian diet (Teleki, 1973). These observations can be phrased

116 Table

P. R. THOMPSON

2

Food shoring

Order No. Carnivora

0

Primates

13

Total

13

No Species

Total

YeS No.

Species

11

Acinonyxjubatus Canis aureus Canis lupes Crocuta crocuta Fe&s concolor Lycaon pictus Lyre rufu Panthera lea Panthera pardus Panthera tigris Vu&esfulva Pan troglo&tes

1

Alouatta palliata Calliccbus moloch Cercopithecusa&o& Colobus guerciz E@hrocebus patas Gorilla gorilla Hyiobates lar Macaca mu&& Mauaca radiata Papio anubis Papio hamadryas Presbytis entelIus Saimiri oerstedi

12

11

14

25

+ = o-92, x3 = 17.72, P < 0401.

into hypothesis form. Meat has both higher calorie value than vegetal matter and requires more skill to obtain, therefore it would be adaptive for meat eaters to share food within their social group, whereas primates, being vegetarian, would find the abundant supply of vegetal matter makes sharing unnecessary and the bulk of comparable vegetal calories makes it very difficult. Table 2 shows that food sharing is significantly positively correlated with carnivores (# = 0.92, xa = 17.72, P < O-001). The only primate to share food is the chimpanzee, and it shares food only when it is meat (Teleki, 1973). Carnivores also diverge from primates in food storing. Leopards frequently suspend their kills in trees to keep them from hyenas and return for them later; hyenas submerge kills in water; lions drag kills into thickets; and foxes bury them. There are no comparable instances of caching food among primates. A second hypothesis on primate and carnivore behavior is thus offered in food storing. Since meat is difficult to procure and relatively compact, carnivores would evolve methods of storing it; however, for primates to store bulky vegetal matter would be difficult and of doubtful adaptive value given the easy access. Table 3 shows food storing is positively correlated with carnivores (4 = 0.85, xe = 14.52, P < O-001). The two exceptions, cheetah and wild dog, share the highest hunting success rate. Among primates, cannibalism is nonexistent. However, it does occur among carnivores, although the factors which trigger it remain unknown and its occurrence is less frequent than pure opportunism predicts. Nevertheless, it can be hypothesized that cannibalism is associated with carnivorism.

CARNIVORE,

PRIMATE

AND

117

HOMINID BEHAVIOR

Superfamily-Ceboidea (Simpson, 193 1) Family-Cebidae (Swainson, 1835) + Subfamily-Homunculinae (Bordas, 1942) + Homunculus hatingtoni (Rusconi, 1935) + Homuncufus patagonicus (Ameghino, 189 1) (= Homunczdw umeghini, Blunt&i, 1931) + Dolichocebusgaimansis (Kreglievich, 1951) + Anthropus pcr&tus (Ameghino, 1891) + Pitheculus australis (Ameghino, 1891) Subfamily-Alouattinae (Elliot, 1894) + Stirtoniu tatucoensis(Hershkovitz, 1970) (= Homunculus tatacoensis,Stirton, 1951) Subfamily-Pithecinae (Mivart, 1865) + Cebuf@theciasarmientoi (Stirton, 1951) Subfamily-Cebinae (Mivart, 1865) + NeosaimiriJieldsi (Stirton, 1951) Family-Uncertain + Branisellu boliviana (Hoffstetter, 1969)

Figure 2. Classification of Tertiary South American monkeys.

Table 4 shows cannibalism xs = 11.81, P < O-001).

is positively

correlated

with carnivorism

(r$ = O-77,

Interspecies aggression During the Pleistocene, preagricultural man hunted a huge number of large mammals into extinction. Martin (1967) calling the phenomenon “Pleistocene Overkill,” estimated the loss at 200 genera. Some evidence strongly suggests that other predators are capable of hunting prey to extinction. Kruuk (1972) using the term “surplus killing” described how 19 hyenas took advantage of a dark night with heavy rain to kill or badly injure 109 Thompson’s gazelle, eating only 13 of 59 dead ones. There was no evidence of selection for healthy or mature animals. In commenting on a fox which had killed 239 gulls during a night storm, Tinbergen & Ennion (1967) stated that “a fox must have an ingrained habit to kill on sight because, clearly in lean times it cannot afford to lose the chance of a meal by hestitating even for a split second.” The following hypothesis is suggested by these observations. Surplus killing is a characteristic of carnivores.

Table 3

Food storing No Species

Order No. Carnivora

2

Primates Total

14 16

No.

Acinonyxjubatus Lycaon pictus

all

r$= 0.85, x’ = 14.52,P < 0.001.

Yes Species Canis aureus Canis iupes Crocuta croci& Felis concolor Lynr rufw Panthcra lea Panthcra pardus Panthera tigris Vu&es fulva

Total

11

14 25

118

P.

Table 4

Cannibalit3m

R. THOMPSON

No Species

Order No. Carnivora

3

Primates Total

14 17

Total

YeS No.

Acinonyxjubatus Lynx rufus Lycaon fiictus

8

all

0 8

Species Canis aureus Canis @es Crocuta crocuta Felis concolor Panthera Ceo Panthera pardus Panthera tigris Vulpesfulva

11

14 25

+ = 0.77, x* = 1 l-81, P < O-001.

Table 5 shows surplus killing is positively correlated with carnivores (4 = 0.77, X* = 11.81, P < O-001). Among primates there is little evidence, exclusive of natural enemies, of interspecific aggression. Primates are tolerant of other species (nonpredators) and even derive benefits from coexistence by using other species communication signals to warn them of approaching predators they have themselves not yet detected (Sarel & DeVore, 1965). Among carnivores there is, in addition to predator-prey aggression, a general animosity toward other carnivores. Lions make unprovoked attacks on leopards and cheetahs and kill them (Schaller, 1972). Wild dogs have attacked lions and tigers and eaten them (Schaller, 1968, 1972). Kruuk (1972) reports that lions account for 55 ‘A of hyena mortality. Relations are similar in the New World. Schaller & Lowther (1969) believe this intracarnivore intolerance is the result of competition for the same food resource, particularly the tendency of predators to usurp one another’s kills. The following hypothesis is thus offered. Carnivores manifest interspecific aggression not related to predator-prey; primates manifest no aggression other than predator-prey. Table 6 shows that nonpredator-prey mortal interspecies aggression is positively correlated with carnivores (q5 = 0.86, X* = 14.86, P < 0.001). The chimpanzee and Savannah baboon will prey on other primates, but even this hostility has not caused a complete intolerance. Teleki (1973) reports that even baboon troops that haveexperienced Table

5

Surplus killing No Species

Order No. Carnivora

Primates Total

3

14 17

No.

A&onyx jubatus Canis aureus Lycaon fiictus

8

all

0 8

c#,= O-77,x’ = 11.81,P < 0401.

YeS Species Canis Lupus Crocuta crocffta F&s concolor Lynx rufw Pantfwa lea Pan&a pardus Panthera tigris Vu&esfulva

Total

11

14 25

CARNIVORE,

Table 6

Mortal

Order No. Carnivora Primates

0 12

Total

12

PRIMATE

intempecics

AND

HOMINID

aggression

No Species

Alouatta palliata Callicebus moloch Cercopithecw aethiops Colobusguereza Evthrocebus patas Gorilla gorilla Hylobates lar Macaca mulatta Macaca radiata Papio hamadryas Presbytis entellus Saimiri oerstedi

No. 11 2

13

119

BEHAVIOR

not preditor-prey Yes Species all Pan troglodytes Pagio anubis

Total

11 14

25

#I = O-86, xa = 14.86, P < O-001.

losses to predatory chimpanzees commonly allow these same chimpanzees to pass among them. Therefore, there is some doubt that the above hypothesis has any exceptions at all. The traditional enmity between cat and dog is as apparent in the largest carnivores as it is between tabby cat and village cur. Groupdefense In baboons and macaques the males have evolved large canines to defend the troop against predators, while the canines of the females are by comparison much more modest and are not typically used in troop defense. Male-dominant defense occurs in other primates as well, but is scarce in carnivores. Carnivores need to maximize the number of food providers and for that function a male is as useful as a female, the lion being a notable exception. Nonhuman primates lack a scarce food supply and their social system often contains harems, similar to those found in ungulates species which are commonly preyed upon. The following hypothesis is thus suggested. It is adaptive for nonhuman primates to raise the maximum number of young, for which the role of the female is more active than that of the male; it is adaptive for carnivores to maximize the number of food providers, and for that there is no sexual difference. Therefore, males are more expendable for group defense in primates, but not in carnivores. Table 7 shows that males are used more for group defense in primates than in carnivores ($ = O-58, xa = 6.43, P < 0.025). The male lion assumes the primary burden for pride defense because the lioness assumes the primary burden for pride economy (Schaller, 1972). Lions are, therefore, a dubious exception. Feeding of young The young of solitary carnivores are entirely dependent on their mother. The young of social carnivores and primates get some added care from other adults, except in

P. R. THOMPSON

120 Table

7

Group

defense

No.

Primarily male Species

Order

Carnivora

1

Primates

10

Total

11

Panthera leo

No. 10

4

Alouutta palliata Colobus guereiz E@hrocebus patas Gorilla gorilla Hylobates lar Macaca mulatta Macaca radiata Papio anubis Pa&o hamadryas Presbytis entellus

Both sexes Species Acinonyxjubatus Canis aweus Canis lupes Crocuta crocuta Felis concolor Lycaon pictus Lynx rufu Panthera pardus Panthera tigris Callicebus moloch Cercopithecus aethiofx Pan troglodytes Saimiri oerstedi

14

Total

11

14

25

+ = O-58, xa = 6.43, P < O-025.

feeding. Carnivore young are vulnerable to the feast or famine economy of a hunter; primate young are not. Consequently, it can be hypothesized that carnivores would minimize the severity of lean seasons by dividing the feeding of young among more than one adult whenever possible, whereas primate young would be dependent wholly on their mother. Since multiple-adult feeding of young depends logically rather than empirically on the presence of two or more adults, only social species will be considered. Table 8 shows that among social species multiple-adult feeding of young is positively correlated with carnivores (4 = O-86, P < 0.001, using Fisher’s exact test). For wolves, wild dogs, and jackals multiple-adult feeding includes both sexes, who will carry food in pieces or regurgitate it to den young. Male lions will not feed pride cubs, but lionesses feed and suckle cubs communally. For hyenas, cannibalistic tendencies of stranger hyenas are probably responsible for delegating feeding young wholly to their mother and, according to Kruuk (1972), making the females bigger and stronger than the males.

Table 8

Number

Order No. Carnivora

Primates Total

1

14 15

of adults feeding one youug One, Mother Species

No.

Crocuta crocuta

4

all

0 4

More than one Species Canis aureus Canis lupus Panthera lea Lycaon pictus

# = 0.86, P < O@Ol using Fisher’s exact test.

Total

5

14 19

CARNIVORE,

PRIMATE

AND

121

HOMINID BEHAVIOR

4. The Carnivore Sophistication Evolution of Hominid Behavior

Scale and The

Seven variables dealing with carnivore and primate behavior have been described in the previous section. For each variable a theoretical argument was derived, based when possible on existing literature on carnivore and primate behavior. The hypothesis resulting from theoretical argument was tested, using taxonomic affinity within the order carnivora and order primates as the independent variable and a behavior dichotomy as the dependent variable. The findings of this analysis are presented in Table 9. The similarity and dissimilarity of hominid behavior to carnivore and primate behavior can now be established by comparison matching. Table 9 shows that preagricultural man resembles the carnivores in food sharing, food storing, cannibalism, surplus killing, interspecies intolerance, and feeding of young. Only the male-dominant group defense of preagricultural man resembles the primates. A scale of carnivore sophistication was constructed from the characteristics listed in Table 9. For a behavior variable to be included in this scale it must meet two criteria: first, the behavior had to be associated with the order carnivora. Second, the behavior had to be solely dependent on the independent variable; i.e. a variable was not included if it was confounded with two or more independent variables. Thus, solitary carnivores were not coded for multiple-adult feeding of young because this variable couId as likely depend on their social structure which lacks two or more adults as on their classification as carnivores. The degree of carnivore sophistication of any species is computed by totaling the number of characteristics which are listed under carnivore then dividing this number by the total number of behaviors which are reported for that species. In this analysis social species have seven variables and solitary species six. For the example of preagricultural man this statistic is + or O-86. Since each behavior variable used to construct this scale is associated with order carnivora, it is expected that high scores should be carnivores and low scores primates. This expectation can be formulated as a testabIe hypothesis. The higher the carnivore sophistication score the more likely the animal is classified as a carnivore. Table 10 shows that this prediction is strongly confirmed (rDb= O-94, t = 13.38, P < 0*0005). The validity of this scale as a continuous measure of carnivore characteristics can also be verified by using it as the independent variable to test a hypothesis about carnivore and primate behavior, e.g. one for which the xa cannot be used owing to low marginal frequency. Table 9

Comparison

Dependent variable Carnivore Shares food Stores food Cannibalism Surplus killing Interspecies intolerance Group defense No. of adults feeding one young

4

Yes Yes YeS Yes Yes Both sexes More than one

of carnivo~,

primate,

Independent variable Primate Preagricultural No No No No No Male One

and hominid

man

Ye.5 + Yes + Yes + Yes + Yes + Male More than one +

behavior

122

P.

Table

10

R.

The

THOMPSON

carnivore

Order carnivora

sophistication

scale

Order primates

Canis lupus Felis concolor Panthera pardus Panthera tigris Vulpesfulva Canis aureus Crocuta crocuta Pan&a leo Lynx rufus Lycaon pictus Acinonyxjubatus

0.43 0.14 0.14 o-14 0.14 0.00 o-00 0.00 0.00 0.00 0.00 0.00 0.00 0.00

Pan troglodytes Callicebus moloch Cercopithecusaethiops Papio anubis Saimiri oerstedi Alouatta palliata Colobus guereza Erythrocebuspatas Gorilla gorilla Hylobates lar Macaca mulatta Macaca radiata Papio hamadtyas Presbytis entellus

1.00 I.00 1.00 1.00 1.00 O-86 0.86 0.86 0.83 o-57 0.50

r 9) = 0.94, t = 13.38, P < 0*0005.

The young of carnivores care

from

primates ranges

their

mother.

unprotected

mobility

while the mother

social carnivores cats.

some wild dogs guard mobility

and more

calling

sophistication

often

restricted

results

mobility

The larger

in leaving

a division

Schaller

of labor

while others

the den while others to guard

fetch cubs.

hunt. their

home Some (1972)

not available

(1965)

The following young

reports

hypothesis

at all times;

greater

can guard their young at all times only by performing

for a division

of labor.

Three

species manifest

wild dogs O-57, and preagricultural

score

Kuhme

of this group

is significantly

score of all social species without

man O-86. greater

than

a division of labor

of

the young

their safety to no mean extent.

to cope with this dilemma.

in the lion made possible

allows primates

lions 0.86,

sophistication

carnivores

hunts, jeopardizing

a carcass

means social carnivores labor:

ranges

A lioness often serves as a guard for small cubs while others hunt.

pride lion may protect

concurrently,

of solitary

have evolved mechanisms

that group existence

to solitary

Lesser

are born largely helpless and would die without home

smaller

allows the adults to keep the young with them at all times. and greater

argued

and primates The

One that

is offered. mobility two tasks

such a division

of

The mean

carnivore

the

carnivore

mean

(rsb = O-52, t = 2.59,

P < O-01). The male-dominant general

divergence

group defense has been retained from

the primates.

The

reason

in hominid

might

evolution

be attributed

despite a

to the char-

acteristics

that are associated with it. In preagricultural man, the male dominant defense is present with the division of labor in which men serve as hunters and women as

gatherers.

The division of labor is associated

male-dominant division

Since evolution

with carnivore

defense may have been retained

Lophistication,

because it is associated

therefore

the

with the adaptive

of labor.

behavior

itself does not fossilize,

the behavior

exhibited

it is difficult

by preagricultural

(1959) reports cannibalism and intraspecific among fossil australopithecines at Taung, sites :

to know how early in hominid

man was attained.

However,

Dart

intolerance (against contemporary carnivores) Sterkfontein, Makapan, and Makapansagat

“with early universal cannibalism . . . this common bloodlust differentiator, this predacious habit . . , separates man dietetically from his anthropoidal relatives and allies him rather with the deadliest of Carnivora”

(Raymond

A. Dart,

1953).

CARNIVORE,

PRIMATE

AND

HOMINID

BEHAVIOR

123

I am grateful to Dr J. E. King and Dr C. A. Rogers for their critical reading of the manuscript. References Abbott, S. & Fuller, T. (1971). A fresh look at the cheetah’s private life. Smit/t.ronian2(3), 34-43. Ardrey, R. (1961). Afticun Genesis. New York: Dell Pub. Co., Inc. Bailey, T. N. (1972). The elusive bobcat. Natural History 81(8), 42-49. Baldwin, J. D. & Baldwin, J. I. (1972). The ecology and behavior of squirrel monkeys (Saimiri oerstcdi). Primates 11(3), 271-284. Be&son, G., Ross, B. A. & Jatinandara, S. (1971). The social behavior of gibbons in relation to a conservation program. In (L. A. Rosenblum, Ed.), Prbate Behavior. New York: Academic Press, pp. 225255. Birdsell, J. B. (1968). Some predictions for the Pleistocene based on equilibrium systems among recent hunter-gathers. In (R. B. Lee & I. DeVore, Eds), Man the Hunter. Chicago: Aldine Pub. Co., pp. 22940. Birdsell, J. B. (1972). Human Evolution. Chicago: Rand McNally and Co. Burrows, R. (1968). WildFox. New York: Taplinger Pub. Co., Inc. Carpenter, C. R. (1940). A field study in Siam of the behavior and social relations of the gibbon (Hylobates lar). In (C. R. Carpenter, Ed.), Naturalistic Behavior of Nonhuman Primates. University Park, Pennsylvania, Pennsylvania State University Press, pp. 144-271. Carpenter, C. R. (1965). The howlers of Barr0 Colorado Island. In (I. DeVore, Ed.), Primate Behuvior. New York: Holt, Rinehart and Winston, pp. 250-291. Chivers, D. J. (1969). On the daily behavior and spacing ofhowling monkey groups. Foliu Primatologku 10, 48-102. Cowan, I. (1947). The timber wolf in Rocky Mountain national Parks of Canada. Canadian Journal of Research 25, 139-74. Dart, R. A. (1953). The predatory transition from ape to man. International Anthropological and Linguistic Review l(4), 201-18. Dart, R. A. (1959). Adventures with the Missing Link. New York: Harper and Brothers. Davie, M. R. (1929). The Evolution of War. Port Washington, N.Y.: Kennikat Press, Inc. DeVore, I. & Hall, K. R. L. (1965). Baboon ecology. In (I. DeVore, Ed.), Primate Behavior. New York: Holt, Rinehart and Winston, pp. 20-52. Eisenberg, J. (1970). A splendid predator does its own thing untroubled by man. Smithsonianl(6), 48-53. Ellefson, J. 0. (1968). Territorial behavior in the common white-handed gibbon, Hylobates lar. Linn. In (P. Jay, Ed.), Primates. New York; Holt, Rinehart and Winston, pp. 180-99. Estes, R. D. & Goddard, J. (1967). Prey selection and hunting behavior of the African wild dog. Journal of Wild&e Management 31, 52-70. Gartlan, J. S. & Brain, C. K. (1968). Ecology and social variability in Cerojnthecus nethiops and C. mitis. In (P. Jay, Ed.), Primates. New York: Holt, Rinehart and Winston, pp. 253-92. Goodall, J. (1965). Chimpanzees of the Gombe Stream Reserve. In (I. DeVore, Ed.), Primate Behuvior. New York: Holt, Rinehart and Winston, pp. 425-73. Hall, K. R. L. (1968). Behavior and ecology of the wild patas monkey. In (P. Jay, Ed.), Prim&es. New York: Holt, Rinehart and Winston, pp. 32-l 19. Hall, K. R. L. & Gartlan, J. S. (1964). Ecology and behavior of the vervet monkey, Ceropithecus acthiofis, Lolui Island, Lake Victoria. Proceedings of Zoological Society of London 145, 37-56. Hall, K. R. L. & DeVore, I. (1965). Baboon social behavior. In (I. DeVore, Ed.), Primate Behavior. New York: Holt, Rinehart and Winston, pp. 53-l 10. Intergroup behavior of free-ranging rhesus monkeys (Macaca mulatta). Folk Hausfater, G. (1972). Primatologica 18, 78-107. Hornocker, M. (1969). Winter territoriality in mountain lions. Journal of Wildlif Management 33,457-64. Hornocker, M. (1970). The American lion. Natural History 79(g), 40-9, 69-7 1. Jay, P. (1965). The common langur of North India. In (I. DeVore, Ed.), Primate Behavior. New York: Holt, Rinehart and Winston, pp. 197-249. Jordan, P. A., Shelton, P. C. & Allen, D. L. (1967). Numbers, turnover, and social structure of the Isle Royale wolf population. American Zoologist 7(2), 233-52. Koford, C. H. (1963). Rank of mothers and sons in bands of rhesus monkeys. Science141,35657. Koford, C. H. (1965). Population dynamics of rhesus monkeys on Cuyo Santiago. In (I. DeVore, Ed.), Primate Behavior. New York: Holt, Rinehart and Winston, pp. 160-74. Tulane Studies Koford, C. H. (1966). Population changes in rhesus monkeys: Cuyo Santiago, 1960-1964. on Zoology 13, l-7. Kruuk, H. (1972). The Spotted Hyena. Chicago: University of Chicago Press.

124

P. R.

THOMPSON

Kuhme, W. (1965). Communal food distribution and division of labor in African hunting dogs. Nature 205,443-4. Kummer, H. (1968a). Social Organization of Hamad~as Baboons. Chicago: University of Chicago Press. Kummer, H. (19686). Two variations in the social organization of baboons. In (P. Jay, Ed.), Primates. New York: Holt, Rinehart and Winston, pp. 293-312. Lee, R. B. & DeVore, I. (1968). Problems in the study ofhunters and gatherers. In (R. B. Lee & I. DeVore, Eds), Man, The Hunter. Chicago: Aldine Pub. Co., pp. 3-12. Lindberg, D. G. (1971). The rhesus monkey in North India: an ecological and behavioral study. In (L. A. Rosenblum, Ed.), Primate Behavior. New York: Academic Press, pp. l-106. Lydekker, R. (1895). A Handbook to the Carnivores, Part I, Cats, Civets, and Mungooses. London: W. H. Allen and Co. Marler, P. ( 1969). Colobus guereza : territoriality and group composition. Science 163, 93-5. Martin, P. S. (1967). Pleistocene Overkill. In (P. S. Martin & H. W. Wright, Eds), Pleisticene Extinctions. New Haven: Yale University Press, pp. 75-120. Mason, W. A. (1968). Use of space by callicebus groups. In (P. Jay, Ed.), Primates. New York: Holt, Rinehart and Winston, pp. 200-16. Mech, L. D. (1970). The Wolf. Garden City, N.Y.: The Natural History Press. Murie, A. (1944). The Wolves of Mount McKinley. Washington: U.S. Government Printing Office. The Ecology of the Timber Wolf in Algonquin Pimlott, D. H., Shannon, J. A. & Kolenosky, G. B. (1969). Provincial Park. Ontario: Dept. of Lands and Forests. Reynolds, V. & Reynolds, F. (1965). Chimpanzees of the Budongo Forest. In (I. DeVore, Ed.), Primate Behavior. New York: Holt, Rinehart and Winston, pp. 368-424. Sarel, E. & DeVore, I. (1965). The Primates. New York: Time Inc. Schaller, G. B. (1965). The behavior of the mountain gorilla. In (I. DeVore, Ed.), Primate Behavior. New York: Holt, Rinehart and Winston, pp. 324-67. Schaller, G. B. (1967). The Deer and the Tiger. Chicago: University of Chicago Press. Schaller, G. B. (1972). The Serengeti Lion. Chicago: University of Chicago Press. Schaller, G. B. & Lowther, G. (1969). The relevance of carnivore behavior to the study of early hominids. South Western Journal of Anthropology25, 307-41. Simonds, P. E. (1965). The bonnet macaque in South India. In (I. DeVore, Ed.), Primate Behavior. New York: Holt, Rinehart and Winston, pp. 175-96. Southwick, C. H., Beg, M. A. & Siddiqi, M. R. (1965). Rhesus monkeys in North India. In (I. DeVore, Ed.), Primate Behavior. New York: Holt, Rinehart and Winston, pp. 11 l-59. Struhsaker, T. T. ( 1967). Social structure among vervet monkeys (Ceropithecusaethiops) . Behaviour 29,83-l 2 1. Teleki, G. (1973). The Predatory Behavior of Wild Chimpanzees. Cranbury, N. J.: Associated Presses, Inc. Tinbergen, N. & Ennion, E. A. R. (1967). Trucks. London: Oxford University Press. Turnbull-Kemp, P. (1967). The Leopard. London: Bailey Bros. and Swinfen. van Lawick-Goodall, H. & van Lawick-Goodall, J. (1970). I nnocentKillers. London: Collins. Vincent, R. E. (1958). Observations of red fox behavior. Ecolou 39, 755-7. Wolfe, M. L. & Allen, D. L. (1973). Continued studies of the status, socialization, and relationships of Isle Royale wolves, 1967 to 1970. Journal of Mummology 54(3), 61 l-35. American Wildlife Institute. Young, S. P. (1944). Wolvesof North America. Washington: Young, S. P. (1946). The Puma. Washington: American Wildlife Institute. Young, S. P. (1958). The Bobmt of North America. Washington: American Wildlife Institute.