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ABSTRACT

TERRITORIAL BEHAVIOR IN PRIMATES

by Charles 0. Ellenbaum
Body of Abstract

This master's thesis is a review of the literature on territorial
behavior and an attempt to use this literature to analyze primate behav-
ior.” It attempts to determine the forms of primate territorial behavior
and the conditioning variables of this behavior. I attempted to find the
answers in the primate data to certain questions such as,what is the form
of territorial behavior, territory or home-range. I found that primates
territoriallbehavior is of the home-range type and is conditioned.by
group living and continuous sexual receptivity. Primate defense behavior

seems to be an expression of maintaining a group's dominance position.

TERRITORIAL BEHAVIOR

IN PRIMATES

by

Charles 0. Ellenbwmn

A THESIS

-Submitted to
Michigan State University
in partial fulfillment of the requirements
for the degree of

MASTER or ARTS

Department of Anthropology

1967

A CKNOWLEDGEMEN TS

I want to thank many people for helping me write this thesis-rm
committee, consisting of Dr. I... Ishina, Dr. D. Swindler, and Dr. S.
Parker, who spent many hours reading this manuscript and helping me in the
many rewritings. If it wasn't for Dr. Swindler, I never would have become
inspired with primate behavior. Drs. John and Ruth Useem gave me a great
deal of encouragement when it was sorely needed and in seminar listened
to av ideas on using animal behavior studies in studying human behavior.

I want, to thank my fiancee for giving me moral support and putting up
with me during the period of writing. I want to thank Mrs. G. Mattson
for proofreading and typing this thesis. I wrote it while studying under

an N'DEA’ IV fellowship.

ii

TABLE OF CONTENTS

Chapter I - Introduction . . . . . . o o . .

Reason for Thesis

Thesis Organization

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Relationships between Human and Primate Behavior

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Chapter II - Territoriality in Nonprimates

Territory and.Home-Range . o .

Instinctive Behavior . .

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Birds 0 o
Rodents o
Ungulates‘
Seals

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Chapter III - Territoriality
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Lungurs

Howler Monkeys . . . . . . .
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Chapter IV - Territoriality in

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O O O O O O

O O O C

O O O O

Gibbon‘........

Gorilla
Chimpanzee .
Hum...

Conclusions

Bibliography . ...

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Capuchin and Spider Monkeys .

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CHAPTER I
INTRODUCTION

Primate territorial behavior, as a major area of study, has never
been done. Concepts of territorial behavior have been developed from
studying birds for the last one hundred years and have been used extensive-

ly to describe and analyze the behavior of nonprimates, such as rodents and

 

fish, for years. Robert Ardrey's Territorial Imperative has been the most
ambitious attempt to study territorial behavior in many different animal
groups. .He makes many generalizations and.hypotheses about it. He says:
Man is as much a territorial animal as is a mockingbird singing in the

clear California night. 'we act as we do for reasons of our evolutionany
past, not our cultural present and our behavior is as much a mark of our
species as is the shape of a human thigh bone or the configuration of nerves
in a corner of the hwman brain. If we defend the title to our land, or the
sovereignty of our country, we do it for reasons no different, no less in-
nate, no less ineradicable, than do lower animals. The dog barking at you
from behind his master's fence acts for a motive indistinguishable from
that of his master's when the fence was built (Ardrey l966:5).

In my research of the literature on primates, I never read much evidence

to show that primates were aggressively territorial. Field reports tend

to minimize territorial behavior or not even mention it. This is in marked
contrast with the field data on nonprimates where territorial behavior plays
an important role in their lives, especially with regard to reproductive
behavior.

I first became interested in the possibilities of primate research
last year while doing a.paper on the field behavior of chimpanzees. Jane
Goodall (1963zhh) describes chimpanzees as "toolsusers," "tool-makers,"
and "transmitters of a primitive culture.“ She is implying that the discon-
tinuities in the behavior between man and at least the higher primates is
not'as great as one might think. This is an example of the current stress

1

 

 

 

2
on the idea that many traits of man which had been thought to be ex-
clusively his, e.g., culture, tool-making, may be found.in at least the
higher primates. The lower primates may actually be capable and, in
fact, exhibit behavior traits which up to now have not been recognized
by anthropologists. Kortlandt and.Kooij (1963:61-62) try to tie man and

the great apes together by classifying them'both as cultural primates.

‘ Harlow (1963:15h-158) discusses the basic process of culture (learning)

and shows that primates exhibit this trait to a large extent in a manner
which suggests cultural transmission. He says:

Culture is dependent upon the ability of an animal within a social group

to acquire from another group member information previously learned.by the
latter member. To obtain such information it is essential that the animal,
particularly the young animal, observe other members of its social group
and duplicate their behaviors whether explicitly or implicitly. . .If pri-
mates can solve formal laboratory problems when the only clues to the solu-
tion are those obtained from observing the experimenter, it seems only
reasonable to assume that they can solve such problems when the cues are
obtained'oy observing or imitating a fellow member of their species. . .
The existence of human-type culture is dependent upon learning capability
and the capacity to transmit this learning to subsequent generations.
“Without language, the capacity to transmit previous learning is dependent
upon either imitative forms of learning or on a similar type of learning

in which the infant follows the group, and the movement and behavior of the
group are such as to place the infant repetitively in the situations where
the infant learns by direct experience the danger responses or affectional
responses essential to individual and group survival.

In his laboratory work, Robert Yerkes (l9h3:52) observed this learning pro-
cess at work. Chimpanzees taught each other how to use a push button'water
fountain. He (l9h3:59) provided evidence that the young male must learn
hotho copulate and the female how to care for her infant. Carpenter (l96hg:
3h2) goes beyond the learning process and generalizes both on the similari-
ties between human and primate behavior and the uses of primate behavior in
studying human behavior. He says:

That human behavior characteristics has analogies and homologies in non—

H human vertebrate and invertebrate forms is evident. That consequently,

continuities, homologies, convergencies, variations, and specializations

3
of behavior and social relations can be discovered and systematically com-
pared among species of the phylogenetic series. That non-human types which
have the greatest degree of general morphological similarity to man, or
‘which show anthrOpoid specialization or convergencies when studied.from the
viewpoint of behavior or social relations, are likely to yield data most
useful in understanding and interpreting problems of human behavior. That
for the valid investigation of some problems in comparative behavior it is
obligatory not only to study animals as wholes but to observe whole animals
in natural, organized undisturbed groups living in that environment which
Operated selectively on the species and.to which the species is fittingly
adapted. That all behavior (or culture) has physiological substrate.
By studying a.particular behavioral pattern in primates, e.g., learning,
'we mey'be able to isolate the components of this behavior, and, with this
information, examine the same type of behavior in humans. Experimental
psychologists for years have constructed.behavioral models based on animal
research. Anthropologists have conducted cross-cultural research since
Tyl'or'. With appropriate safeguards, anthropologists should be able to use
cross-species research to study social processes.

I believe that human behavior should generally fit within the broad
context of primate behavior, and that primate behavior should help to pre-
dict the broad outlines of human behavior at the very least. One cannot
lose sight of the fact that the living primates and man are the contemporary
end.points of distinct evolutionary histories existing in different environ-
ments. Behavioral differences and divergencies are to be emected. Primates
are the closest living group to man suitable for cross-species research. The
results of such research could provide guidelines and models of behavior as
well as ideas for future research. However, one must avoid the trap of
appearing to formulate a rectilinear model of cross-species behavior and
behavior evolution.

I will first examine the literature on territorial behavior in non-
primates and then in the human and non-human primates. For each animal

population I am looking for the answers to certain key questions-~Is terri-

h
torial behavior of the home-range or territory type? Is it learned or in-
stinctive behavior? Is defensive behavior observed and is it part of
territorial behavior or some other behavioral system? . What functions does

territory serve and is it replaced by other behavior systems?

CHAPTER II
TERRITORIALITY IN NONPRIMATES

Territoriality in some form is found in virtually every animal group
from fish to primates. The very spread of this behavioral system.indicates
the adaptive significance it must possess to exist in such a diverse group
of animals. In many of the nonprimates, territoriality seems to rest on
an instinctive hormonal basis. It can and does serve a variety of adaptive
functions for each animal pOpulation. Carpenter (1958:229) feels that
variations in territorial behavior are related to differences in the species
and their habits, to season and climate, to population pressure, to social
organization, to fluctuations of food supply, to predation and other factors.

Because of the wide species range of territorial behavior, an inves-
tigator must be wary of interSpecies' similarities and must determine whether
they are homologous, homoplastic, or analogous before making generalizations.
According to Simpson (1958:533), homology is resemblance due to inheritance
from a common ancestry, homeplasy is purely convergent resemblance without
inheritance in common, and analogy is functional equivalence that is not
homologous and not necessarily homoplastic. Some behavioral examples would
be: (a) the homologous eye-hand manipulation in man and other primates, (b)

I homoplastic nectar feeding in hummingbirds and Sphinx moths, and.(c) ana-
logous locomotion in earthworms and.moles.

The terms "territorial behavior," “territoriality," and “territori-
alism" should be treated as different concepts although many authors do not
always do so. In general, territoriality is any behavior on the part of an
animal which tends to confine the movements of the animal to a particular

locality (Etkin l96ha:21-23). This behavior can take various general forms.
5

6
Etkin (l96ha:2h) defines home-range as a sense of locality which is charac-
terized'oy an element of attraction to an area, but without the element of
driving others of the same species off the home-range. Territory is a
defended area (Etkin 196ha:2l-23). Both home-range and territory confine
an animal to a.particular range, but only in a territory does the animal
fight to defend the area, usually only against members of the same species,
but sometimes also against species with similar food habits.

Two distinct forms of territory exist, object oriented and geograph-
ical. The defense is the defining variable according to Ratner and.Denny
(l96h:h31). In an object oriented territory, the defense occurs around a
particular object, e.g., buck deer fight around the doe. In a geographical
territory, the entire area is defended even though the nest often becomes
the center of this territory. Animals often mark the boundaries of their
territory in such a way that the boundaries are recognized by other members
of the species. Birds primarily use song, announcing their possession
audibly, while some mammals follow yet another method of setting marks with
scent. Gronefeld (1965:2h5-2h7) Showed that tiny traces of musk stick to
tree trunks, to bushes, and to rocks and stones that flank the most probable
entry ways to the territory. Bears, peccaries, red deer, and.lynmes are a
few of the animals who use scent to mark their territory,but as far as is
known, free-ranging primates do not use scent.

Defensive behavior in a territory may take various forms. Ratner
and Denny (l96h:h32) describe the forms of defense from territorial encroach-
ment as usually consisting of ritualized threats, threatening advances, and
actual physical fights. Specialized.body structures and/or postures are
frequently used'by the defending animal. Etkin (l96ha:29) believes there

is a definite tendenqy for an animal in a conflict situation to show increas-

7
ing dominance as he comes closer to the center of his territory. This
tendency places a limit on the reduction in size of a territory; for when
an owner‘s territory becomes small enough, he will fight to the death to
prevent further loss.

By far the commonest forms of nonprimate territory is the indivi-
dually held area necessary for reproduction, or the pair held area that is
used for reproduction and raising of offspring. But the key factor is re-
production. By confining the potentially reproductive individuals to a
small territory, their behavioral and physiological co-ordination is facili-
tated. By only allowing territory-holding individuals to mate, a theoretical
upper limit is placed on population increases in terms of the carrying ca-
pacity of the land (Etkin l96haz31-32). Odum (1965:222) believes that it
also helps to reduce competition, conserve energy during critical periods,
and to prevent overcrowding and consequent exhaustion of the food.supply.
The strongest, most aggressive and fit are selected to breed. Those who
are not fit enough to secure a territory do not breed.and, hence, do not
perpetuate themselves through offspring. Simpson (1958:532) believes that
the sole function of natural selection is reproductive success. Scott
(l96hz231) says that the most general function of behavior is adaptation‘
to environmental change which ensures reproductive success.

Simpson (1958:522) makes a distinction between immediate and long-
range adaptation of a particular behavioral aystem, i.e., territorial be-

havior, to the environment. Immediate adaptation tends to be narrower

 

(more specific in its correlation'with environmental factors) and less
variable, while long-range adaptation must be maintained in the face of
extremes of environmental fluctuation and of inevitable evolution of the

environment itself. The environment includes all biotic associations

8
and.not just physical factors. The Gombe Stream.chimpanzees ate meat whereas
the chimpanzees of the Budongo Forest do not (Goodall 1963zh53‘1965:hh3-hh5).
Meat eating is an immediate behavioral adaptation tied to the environmental
fact that the Gombe Stream area area is not as lush in vegetable food as is

the Budongo Forest. An example of long-range adaptation is the chimpanzee

 

adaptation to terrestrial life.

In general,according to Etkin (l96hb:279), territory correlates with
the nature of the reproductive pattern and defense is usually a male func-
tion unless a pair holds the territory then the pair defends it. Carpenter
(1958:229) believes that those processes which are called territorial are
actually higher order, complex behavior systems which are based on a plur-
ality of subsystems. For example, highly deve10ped territorial, behavioral,
and food—gathering techniques exist among birds for the protection and sus-
tenance of nestlings (Freedman and Roe l9S8:h7h). For animals who migrate,
territorial behavior relates to the beginning or terminal phases of migra-
tory'responses. Territorial behavior relates centrally centrally to the
complex behavior of possession and guarding areas of space; elements of
selective and discriminatory responses; complexes of reproductive behavior;
attack, encroachment, and defense; challenge, vocalization, song, and other
displays and/or signaling activities; foods and feeding; search, securing,
hoarding, and.protecting food; security, covers, lairs, dens, nests, burrows,
or even places providing merely space and distance. Territorial behavior
is a social phenomenon involving individuals, pairs, and groups (Carpenter
1958:229-230). Defense may be an individual or pair endeavor. It must be
'omxe in mind that territorial behavior is not a single simple phenomenon
but is part of many behavioral systems and fulfills certain critical func-

tions or it would be replaced.

9

Territorial behavior in the nonprimates seems to be intimately bound
up with reproduction. The onset of this territorial behavior seems to be
largely,but not wholly, dependent upon a combination of hormonal signaling
and instinctive behavior cued by outside stimuli. Perhaps I should define
instinctive behavior. Iorenz (1957b:290) says that instinctive behavior
consists of at least three components: (1) appetitive behavior motivated
by internal accumulation of readiness for a specific action, (2) activation
of an innate releasing mechanism (IBM) which disinhibits the instinctive
reaction, and (3) discharge of the consummatory act which is the purpose
of the behavior. He (1957a: 116) says that the IBM is the key combination
of perceived stimuli that the animal is ready to respond to and initiate
a certain behavior pattern. The IBM seems to be nothing more than a re-
leasing stimulus. It is called a mechanism but the nature of behavior
mechanisms is largely not known for sure. It is undoubtedly highly cor-
related with the genetic code of a species and the level of hormonal out-
put. It is described as either a behavioral trait (a song or posture) or
a physical trait (color of belly or breast). An animal raised in isolation
should be able to perform the behavior perfectly, without prior learning,
if the prOper stimuli are present.

Perhaps the male stickleback has been the fish most studied. Male
sticklebacks live in schools until spring when they establish territories.
Two male sticklebacks will chase one another until they reach an invisible
boundary where neither is dominant over the other. Tests by wunder (1930)
and ter Pelkwijk and Tinbergen (1937) (Carpenter 1958:230) have shown that
the male stickleback will suppress its own reproductive behavior when other
males are in close visual distance. This condition could.not occur if the

male were dominant in a territory. Territorial behavior is confined to the

lO
spring by a combination of differing hormonal output and.by the males only
having a red belly in the spring, according to Hinds (l966:h5). The IRM
or releasing stimulus of defensive behavior of the territory is the red
belly and not the shape of the male. A circle, if the lower half is red,
‘will trigger this type of behavior.

In an experimental study of the cichlid.fish, Breder (l93h) (Carpen-
ter 1958:230) concluded that the territorial behavior of fish closely re-
sembled that of birds. Territories are established, nests built and guard-
ed, and mating occurs within the territory. The male stickleback exhibits
the same behavior, but he chases the female away after she has laid her
eggs and thus no enduring male-female pair bond is formed. Noble (1938),
in his review of sexual selection among fish,reports that in different
Species, every male, or spawning couple, occupies a small territory within
which no other sexually active member of the same species are tolerated.

In experiments on factors influencing the size of territory in Chinese fish
(Cyprinidae), Fabricus (1951) (Carpenter 1958:231) found that males are
strongly territorial during spawning, stay in a small area, and defend it
by lateral display, but rarely by fighting. Fish size does not seem to be
correlated with the size of the territory. He also found that bream.males
in a lake are strongly territorial and that territories are defended by
splashing reactions. Territory sizes relate to the topography of the lake
bottom and the amount of vegetation. In another experiment, Fabricus (19Sh)
(Carpenter 1958:231) found that char males defend well defined territories.
The fish swims over its territory and vigorously attacks all trespassers.
Territorial males may, during the chase, cross territorial boundaries, and
they may leave their territory to join females and then return. Females

also occupy definite territories.

11

Bird territorial behavior has been the subject of many books and
articles. Altum (1868) (Carpenter 19582225) deve10ped a systematic descrip-
tion of territoriality for birds. He said that "territory is an area oo-
cupied by one male of a Species which it defends against intrusion of other
males of the same species and in which it makes itself conspicuous.“ He
introduced the idea that birds settle at fixed distances from each other
in order to ensure food for themselves and their young. His ideas on dis-
tance and food seems to be an early example of the'wynne-Edwards Hypothesis
which ties spacing and population size with the amount of the food.supply.
Moffat (1903) (Carpenter 1958:225-226) suggested that fighting serves to
parcel out territories which in turn limits the number of pairs. .Territor-
ies as a prerequisite to mating would prevent indefinite population increases
and would condemn less powerful individuals to unreproductiveness rather
than to death. Territory, by allowing only the most adaptively fit to
reproduce, ensures the viability of the species. .

Eliot Howard (l907-19lh, 1920, 1929) (Portmann 1961:25) made some
of the classic studies of territory in bird life. He expounded the theory
that brooding birds possessed territory for the purpose of preserving the
species. Howard (Carpenter 1958:226) showed that the male "isolates him-
self, makes himself conspicuous, becomes intolerant of other males (of his
Species) and confines his movements to a definite area." He believed that
territoriality serves the function of ensuring a food supply, ensuring
mating by providing a.focus for activity prior to nest building, and limit-
ing the number of bird pairs in a limited area.

Diebschlag (l9hl) (Carpenter 1958:23h) has Shown that house wren
territory plays an important role in establishing and maintaining peck

order dominance in the social hierarchies. Hochbaum (19hh) (Carpenter

12
1958:23h-235) observed that the boundaries of the canvasback duck's terri-
tory and tolerance range are the same and as one varies, the other varies.
The tolerance is not a species constant.

Carpenter (1958:235) believes that a near universality of some sort
of territorial behavior exists even though types and.patterns of their be-
havior Show great variance. Brown (1965:1h) believes that songbird terri-
tory is typically an allepurpose family area in which mating, nesting, and
food-seeking occurs. Some evidence exists to show that the average size
of a territory for a species varies with food abundance. The Alaskan
pemarine Jaeger feeds chiefly on lemmings. In 1952, there was a moderate
number of lemmings and four Jaeger pairs per square mile. In 1953, there
were many lemmings and eighteen Jaeger pairs per square mile. Brown (1965:
15) believes that this data shows that one of the functions of territory'
is to disperse pairs so that they can obtain a constant ideal amount of
food.per family.

Bird territory also functions in other areas. A territory gives a
subordinate animal a clear advantage. A subordinate animal in its own
territory will usually win a fight with a dominant animal. Hewever, a
dominant animal will usually defend a larger and.more choice territory and
will.usually attract more desirable females.r According to Noble (1939)
(Carpenter 1958:23h):

Territory should be sharply divided into different categories. Sexual
territory which is so characteristic of most birds is, however, found in
most egg-laying and.nest-building vertebrates. It arises from.the sexual
interest of the animal in an area suitable for nesting and it functions
primarily to test sexual readiness of the Opposite sex and.to make sexual,
bonds. Sexual territory is not to be confused with nesting territory which
has a different motivational basis, nor with an isolated retreat which is
defended by many vertebrates against intruders at any season.

Hinds (1966:237) states that in fighting over territory, it isn't more

proximity, but the presence of another bird within a particular area which

13
elicits aggression. A bird ignores a nearby rival outside his territory
but flies off to attack a more distant one who is within its boundaries.
The territory owner constantly patrols his territorial boundaries.

Just as in the male stickleback, there are certain IRMis for birds
'which trigger territorial defense behavior. Fighting seems to be elicited
by a fairly specific set of releasing stimuli. The red.breast is of pri-
mary importance in eliciting threatening behavior in territory holding
robins (Hinde l966:h0). The Chaffinch needs a red.breast and the Great
Tit needs a broad black ventral stripe to elicit fighting behavior. Fight-
ing occurs in the spring when the gonads are active and the sex hormones
are being released (Hinde 1966:237).

'Within the rodent group, Andrewartha (1961:112) has shown that mice
are intolerant of each other within a home-range. He (1961:11h) believes
that there are two important features in their territorial behavior. They
recognize a home-range within which they are more secure against predators
and other hazards because of their intimate knowledge of the area. They
exhibit a peculiar intolerance of crowding. Davis (19h8, 1953) (Carpenter

1958:236-237) defines home-range for the brown rat as "an area regulated
frequented by an individual.“ The scope of behavior or range of behavior
varies from season to season, with sex and with population density. He
concludes that: (l) the range of movement is related to food supply, (2)
rats occasionally shift their home-range, (3) the range of the brown rat
is seldom.more than one hundred.feet in diameter, and (h) radical changes
in the environment may lead to changes in locations occupied. Bradt (1938)
(Carpenter 1958:235) found that beavers build dams, construct houses, and
occupy localized areas. They mark the possessed territory by scent and use

various sound signals to defend the area.

1h

Gordon (l9h0) (Carpenter 1958:235) found that the Sciurus frementi,
living in the lodge pole pine forest, collects pine seeds and.places them
in the center of an area which is guarded by a single individual. This
object oriented (Seed pile) territory provides an alternative to a.random
burying of food seeds. Evans and Holdenried (l9hl, 19h3) (Carpenter 1958:
235) found that Citellus beecheyi (ground squirrel) adult males restrict
themselves to small ranges but there is seldom an overlap of ranges. Fe-
males also remain within their ranges but there is overlapping and sometimes
a complete overlapping of them. I would assume that the male ranges are
actually territories because of the lack of overlap while the female ranges
are actually home-ranges because of the overlap.

Territorial behavior is very evident among the ungulates. .In the
red deer, territory and sex are related mechanisms of their society. .Ac-
cording to Darling (1937, 1952) (Carpenter 1958:238), territory "occurs
within the range and is determined psychologically rather than physiologi-
cally. . .Defense of a group territory among ungulates is not known.“ Home-
ranges result from choice and are known by the occupants.l This learned
home-range has advantages for finding feed and for strategic purposes be-
cause of the intimate knowledge of the terrain. Herds move in response to
adverse conditions and they "may combine in the face of common need" (Car-
penter 1958:238-239).

Buechner studied the kob and came to several conclusions about ter-
ritorial behavior in the kob. The male kobs compete for territories (stamp-
ing ground) and never for females, but only kobs holding a territory are
able to breed. COpulation occurs nowhere but on the stamping ground. On
a kob stamping ground, the territorial prOprietor almost always wins until

the effort of continual defense and mating wear him down to the point where

15
he can be defeated. However the psychological advantages of the proprietor
reduces the incidence and severity of actual fighting. Antagonism between
male kobs is confined to the stamping ground and elsewhere their relations
are amiable. Populations of kob do not really mix, though to the eye they
may seem to. Each feeding pOpulation retains an identity with its own
stamping ground, making interbreeding unlikely. Genetic isolation is thus
achieved. For the male, the attachment to a piece of ground is stronger
than to the female herd (Ardrey 1966:60-62).

Other mammals exhibit forms of territorial behavior. Darling (19b?)
and Bartholomew (1952, 1953) (Carpenter 1958:238) observed that Seal Island
colonies represent spatial organization as one link in the annual cycle of
movement that includes migration. Bulls may occupy territories held and
fought for during previous years. According to Bonner (1955) (Carpenter
1958:238), "They may fight for their territory and no doubt there are
frequent changes of position." Carpenter (1958:238) believes that terri-
toriality has a central role as both a motive and a response in the dom-
inance order of breeding-age bull seals. For this largely aquatic mammal,
territory is a basic condition for reproductive behavior and colony organi-
zation. For seals the individual territory serves as a dueling ground and
as a breeding ground. Many other animals exhibit behavior that could tena—
tively be described as some form of territorial behavior. Lions hunt in
packs (prides) of up to sixteen individuals and though they periodically
change areas, they do cover a definite locality. Herds of grazing ungulates
in Africa such as the springbuck, wildebeest, hartebeest, and gazelle re-
main in definite areas. In another class entirely, various spiders, crabs,
and insects seemingly defend territories (Pearse 1939:221-222).

This data conclusively indicates that territory is held by an indi-

16
vidual or a breeding pair and is part of the species reproductive pattern. _
Territory is quite significant by conferring a biological advantage and by'
being adaptive by ensuring reproductive success and.food for the fit.
Reid (1962: 83) believes that territory is valuable in eliminating indiscrim-
inate and uselessly severe fighting. Territory saves time in the search
for food, makes it easier to find.mates, and may strengthen the male-
female bond. It directs and is a controlling influence on species density.
In summary, I would like to present some of what Carpenter (1958:
2h2-2h3) feels are the possible functions of territorial behavior. It
spaces or diSperses a species p0pulation and thus limits or regulates pop-
ulation size by limiting reproduction to territoryéholding males. By en-
airing.adequate Space, it prevents overpOpulation or widespread starvation
or undernourished offspring. It exposes nonterritorial elements to preda-
tion and affords protection against predators for the territorial members
of the population by an area of knowledge. It reinforces dominance and
selective breeding of the strong but is still advantageous for subordinate
animals. It regulates spawning in fish and facilitates and perhaps ensures
breeding for some species. It reduces sexual fighting and killing and
helps to protect the nest and young. For birds, it provides an attraction
area for females ready to mate and warns away any trespassing animals.
Territorial behavior then performs certain indispensible functions
for these nonprimates. Could another behavioral aystem(s) remove the
necessity for territorial behavior? I would.now like to examine the various
ferns that territorial behavior takes in primates and attempt to explain
the possible differences and absences of it.v Primates are clearly different
from these animals in terms of anatomy, physiology, mode of existence, and

behavior. Since territorial behavior is closely tied to the environment

17
and these are factors of environment, I would.expect to see some changes

in the form and.function of territorial behavior.

CHAPTER III
TERRITORIALITY IN PRIMATES

I am confining my data to that which was collected in the field and
not in the laboratory even though it is someWhat Skimpy for this type of
research. In general, approximately 2h0 primate species exist and only
twenty species have been reliably studied (Southwick 1963:5). In some of
the field literature, territoriality is assumed.but the supportive data is
either not presented or does not seem to conclusively prove the supposition.
In some instances, the authors are contradictory. Buxton (as quoted in
Haddow 1963) contends that the African redtail monkey troops do not exhibit
ady territorial behavior. Directly Opposite, Haddow (1963:63) says that
the larger groups of African redtail monkeys do exhibit definite territori-
alism. The gaps in the literature indicate that this particular area would
be very productive to study in both the field and.in the laboratory situa-
tion. Let us now turn to the literature and examine territorial behavior
in primates.

Lemurs live on the island of Madagascar. Lepi—lemur groups move
independently; each one in its own restricted area. According to Petter
(1965:307-308), they utter many cries which have been interpreted as cries
of defense. This points toward typical territorial behavior. Buettner-
Janusch (1966:231) agrees with Petter and says that lemur troops appear to
exhibit territorial behavior. "They recognized foreign troops of their own
species and gave warning calls when they appear, and either sit in trees
and make warning or threat gestures or retreat." He (1966:238) believes
that their behavior indicates that they are aware Of the boundaries of
their normal territory . They try to avoid other troops of the same species.

18

19

Their warning cries seem to function as a spacing mechanism.to minimize
the confrontation of different troops. An individually held, vigorously
defended territory linked to reproduction does not seem to exist. There
is also no evidence to suggest the working of IBM's which would trigger
territorial behavior. Though the data is not clear as to what type of
territory exists, or even if it is a territory rather than a home-range,
it does seem to be vastly different from.nonprimate territory.

Relations between langur trOOps are peaceful with no displays of
aggression. Phyllis Jay (1963:116) never observed fighting when several
troops were next to each other. 'TrOOps were observed to meet and mix
without aggressive behavior. Boundaries were not defended, instead, the
trOOps normally occupy their home-ranges. Home-ranges of adjacent troops
may overlap either slightly or extensively. TrOOps Often come together
in this overlap. Jay (l965b:536) observed that though the home-ranges
overlap, the core area do not overlap. .She defines a core area as an area
of intensive use within a home-range which are usually connected by tradif
tional pathways. Though the overlap areas are usually only occupied.by
one group at a time but when two groups are in the same area, they do not
threaten each other. The larger group usually takes precedence and the
smaller remains at a distance until the larger moves away. This is very'
similar to the action of the dominance hierarchy when an individual waits
for the one more dominant than he to get done, before commencing himself.
Jay (1963:116) observed that langur trOOps mix peacefully with rhesus
macaques. As with lemurs, territorial behavior is still largely confined
to the home-range variety.

Jay (1965bz537) found that only the rhesus macaques living in severe-

ly over-crowded conditions in Indian cities are aggressive and defend the

20
boundaries of their territory. It seems as though the only vigorously
defended territory is that of the "urban" macaques who live in the abnormal
conditions of the city and consequently under the pressure of overpopula-
tion. In this case, territory would seem to be a learned immediate adapta—
tion to a changed environment and not an inherent element in the animal's
behavioral repetoire. Itani (l965:hl2) found that macaque troops range in
size from h to 600 with 30 to 150 the most common size. Usually the trOOp
is an exclusive unit and repulses new individuals seeking entry into the
trOOp. Koford (1963:1h3) found that their home-ranges overlapped some 80%
to 90%. Group movements were so structured that groups usually avoided
contact with each other, as was the case on Cayo Santiago. Each band moves,
feeds, and rests as a unit. He believes that band movements are largely
influenced.by the location of food and encounters with other bands, which
may be hostile. , Southwick, Beg, and Siddiqi (1965:1142) found that behavioral
relationships between different social groups were characterized by marked
intergroup intolerance and a prominent intergroup dominance hierarchy ex-
isted. Dominance gave a group freedom of movement, but severe intergroup
fights occurred when a subordinate group would not retreat. Etkin (l96hb:
276) believes that the dominance of the groupas a unit depended.upon the
fighting.ability of the tOp males. I do not believe that this is territorial
behavior as seen in the nonprimates any more than when a dominant individual
'displlees.another less dominant individual for food or some other dominance
provoking them. Carpenter (1963:39) says that "an individual [Or group] is
said to be dominant over another when it has priority in feeding, sexual,
and colomotor behavior and when it is superior in aggressiveness and in
group control to another or other individuals [Or groups)." In any domin-

ance situation, the subordinate indicates his position by retreating, grim-

21

acing, presenting, or by some other Sign of subordination. Then why not
carry the dominance idea from intragroup to intergroup interaction? The
dominant group is defending their position in the hierarchy by defending
whatever it is they are close to at the particular time they meet with
another group. They do not patrol an entire area to defend it as do birds.
Within an overlap, a group is bound to meet other groups without looking
for them» A robin will patrol his entire territory and only attack those
who trespass and not those who are close. This dominance intergroup fight-
ing seems to be analogous too, but not the same as territorial defense.

Both Carpenter (l96ha:98) and Buettner-Janusch (1966:251-256) believe
that both capuchin monkeys and Spider monkeys live in troops which travel
specific routes in their daily round of activity through the forest. TrOOps
will try to drive other troops away when they meet. While the space of
their ranges may overlap, the time Of use does not. I believe that this
is the same sort of situation as is found with the macaques. The macaques
who live in the forest behave as do the capuchin and spider monkeys.

In areas where howler monkey groups are concentrated and do not have
normally dispersed territorial ranges, the frequency of intergroup conflicts
is increased according to Carpenter (196hd:h05). This situation seems to
be similar to the one involving urban macaques. Carpenter (l96he:87) ob-
served that groups of bowlers tend to occupy a definite and limited.range.
This range may be shared with other groups and from time to time, the range
may Shift somewhat. He (1958:2h1) found that the defense of their position
consists mainly of specialized patterns of vocalization and very rarely
.actual physical fighting. He(l965:273-27h) says that their vocalizations
consist of howling and roaring in bursts that may last for hours. "It

can be seen that the 'territory' concept holding that an animal or bird,

22
or a group of them, defends an exclusive space, den, or nest does not de-
scribe accurately the behavior of howler groups." Carpenter (1965:273)
also found that in the less familiar peripheral range areas, they are more
active, restless, excitable, and appear to be on the defensive. I believe
that this behavior is typical of any animal, from man.on down, that is in
a strange area. Unknown dangers abound and escape routes, if known at all,
are known poorly. This is illustrated by howler monkeys. Hooton (l9h2:
239) says that when a howler group gets out to the edge of its territory,
it becomes slowed down by unfamiliarity and a good deal of milling about
and frustration results. The range boundaries,according to Carpenter
(1965:273), are fluid and variable. The area covered is localized, but
shifts seasonally, largely becausecaf responses to ripening fruits or
preferred leaf trees. Occasionally a group will move for unusually long
distances.

The howler monkeys provide a useful comparison with the rhesus
macaque. The bowlers live in groups like the macaque but aggressive behavior
is much less evident. This is true even when taking into account the less
aggressive behavior of "rural" macaques as compared with "urban" macaques.
Howler males show little evidence of dominance and do not even compete for
the oestrus females nor do they exclude younger males from.the group.
Etkin (l96hb:277-278) found that sexual dimorphism.is not as marked. If
two howler groups come into close contact, a howling contest, not a physi-
cal battle, determine which group yields. The difference in settling
intergroup dominance is undoubtedly'partly due to the fact that macaques
are partly ground dwellers while bowlers stay in the tree teps. Etkin
(l96hb2278) believes that fighting in the tree tops is dangerous and in-

effective. Terrestrially adapted primates would face more predators than

23
arboreally adapted.primates and would need features such as sexual dimorph-
ism of canines and aggressiveness just to survive. However, both use a
form of spatial-temporal defense of the group behavior. They do not defend
a territory, but defend their present position in space and time against
'anofihef group for the same reasons, I believe, that individuals have dom-
inance interactions. I believe that if primate groups exhibit aggressive
behavior against other groups of the same species, that it is analogous
to dominance interactions between individuals and that this is the basis
for defensive behavior of primates rather than a defense of a set geographi-
cal territory. At most, they could have a mobile object oriented territory
which is the tr00p itself. But this is a far cry from the territorial be-
havior of nonprimates which is so closely connected.with reproduction and
the rutting season.

Each pack of baboons keeps to the same general area. Zukermann
(1932:196) never saw any two packs unite for any length of time. De VOre
and Hall (1965:3h-38) never saw any indication of defended boundaries.

This does not imply'that groups moved about without reference to any fixed
boundaries but that the boundaries were not defended. Baboon groups spend
most of their time in circumscribed areas. It was very difficult or im-
possible for De Vore or Hall to drive baboons outside the boundaries of
their area, therefore the boundaries seem to be psychologically real to
the baboons. Range size varies with trOOp size, concentration of food
plants, and the proximity of neighboring troops. TrOOps frequently use
only certain portions of their home-ranges such as sleeping trees, water
resources, resting places, and food sources. They tend to avoid each
other but even in areas of high population density, different groups may

be in close daily contact without displays of intergroup aggression. Hall

2h
(1963:1h) found that even Chacma.baboon home-ranges show varying patterns
in terms of day-ranging and sleeping places according to the nature of the
terrain, distribution of staple foods, and water sources. .They tolerate
other groups without aggressive displays unless for example, the others
attempt to steal females.
The great apes are quite close to man in terms of physical features

and.behavior. For this reason I want to analyze the territorial behavior

of both the great apes and humans.

CHAPTER IV
PRIMATE TERPITORIAL BEHAVIOR

Carpenter (l96hcz23h) found that gibbons restrict themselves to
limited areas and seemingly stayed in them fairly consistently. He (1963:
20-21) believes that gibbons acquire and defend sections of the forest
and generally confine their activities within these possessed.regions.
This territoriality is a fundamental characteristic of the behavior of
free-ranging gibbon according to him. He (l96hcz2h1) observed that com-
petition between groups occurs most frequently in the border regions of
"territories.“ I believe that Carpenter has misnamed these ranges as
territories and that the competition is actually a defense of the group
and that they live on home-ranges and not territories. As evidence to
support this supposition, Schaller (1965azh75) noted that neighboring
groups actually intermingle for short periods of time. Shifts in group
ranges occurred and he thought they were apparently the result of pressures
from shifts in the supply of food, surrounding groups, competiton, and
group splitting.

Gorilla groups restrict their activities to definite areas or home-
ranges on the order of 10 to 15 square miles according to Schaller (1965b:
3h1). “The recognition of certain physical features, coupled with a pre-
ference for familiar terrain is probably the most important factor which
confines the activity of groups to specific ranges." No evidence exists
to show that a section of the forest was defended directly by fighting or
-indirectly by aggressive displays against intrusion of another group ao-
icordingto Schaller (l965b:3hl). Schaller (1963:111, 120) observed that

though dominance was sometimes displayed.between dominant males of different
25

26
groups, groups were familiar with each other. Emlen and Schaller (1963:
133-13h) found that tolerance is extended to new members of the group.
Visiting males may join a tr00p for a few days without overt signs of an-
tagonism. Gorillas do not appear to defend a definite territory. When
one tr00p meets another, they may pass each other by, or they may join for
an hour, or even bed down together for the night. Schaller (1965bz3h2)
concludes that the almost complete overlap of some ranges and observations
of peaceful interactions between groups indicate that gorillas have no
territory in the sense of an exclusive area defended against others of the
same species.

The major field studies of chimpanzee behavior (Nissen 1931) (Good-
all 1965) (Reynolds 1965) indicate the same findings as are seen in the
gorillas.’ Chimpanzees exhibit no territorial behavior in the sense of an
exclusive area defended against the same Species according to Reynolds
(19652hOO-h03). No boundaries exist that a chimpanzee group would hesitate
to cross such as is found in baboons. Reynolds (1963:100) concludes that
chimpanzees are localized in'habitual ranges and.not territories. Goodall
(l965:hSS-h56) reports no territorial defense but a free mixing of bands.
Reynolds (l965:h15) found some evidence for status difference between in-
dividuals but no evidence of an unilinear hierarchy of dominance among the
males and females. No evidence of exclusive rights to receptive females
existed and there seems to be no prominent or permanent leaders of groups.
Reynolds (1965:398-h00) found that the smallest social unit seems to be a
rather unstable band of from two to six individuals whose association is
probably based on personality characteristics or similar age, sex, status,
or condition. He found four recurrent band types which are: (1) adult

bands containing adults of both sexes and occasionally adolescents but no

27
mothers with dependent young, (2) male bands containing only adult males,
(3) mother bands, and (h) mixed.bands which combine the elements of the
other three. Goodall (1965:153) found that aggressive and submissive
interactions between individuals are infrequent.

I think that the data clearly shows that what passes for territorial
behavior in primates is significantly different than it is in nonprimates.
The seemingly territorial defense behavior of primates, I believe, is the
defense of the band and not of a territory and is not linked to reproduction
as it is in nonprimates. On the basis of my data, I believe that the pri-
mate defense behavior is an outgrowth of dominance hierarchy interactions
from intragroup to intergroup spheres of action. The primate groups which
show almost no aggressive behavior and virtually no dominance system, 1.6.,
chimpanzee, gorilla, show almost no defensive behavior. Those groups which
show marked aggressive behavior to one another and in which a dominance
system is working show this defense behavior. This is a defense of group
integrity and dominance position and not of a territory. Before attempting
to explain why territory largely drops out of primate behavior, I would
like to briefly give a few behavioral examples of human territorial behavior.

Bartholomew and Birdsell (1962:2h-25) believe that territoriality
in man is created by the necessity for finding and maintaining environmental
conditions suitable for survival and reproduction. It is clear that terri-
toriality exists in all complex human societies and it is clearly establish-
ed that group territoriality is also important at the simplest levels of
human culture. Service (1966:?) observes that on the band level of social
organization, groups sometimes define themselves territorially as inhabit-
ants and owners of a foraging range. He says (1966:30-31):

There are variations in hunting-gathering societies in the degree of terri-

28
toriality-~that is, in the explicitness of boundaries-~but in all cases
there is some, and in most a rather strict definition of the band in terms
of the general locality even when boundaries are not specific. The terri-
tory is never occupied exclusively or defended against all outsiders.

There is of course a wide range of variation in the form of terri-
torial expression. Extreme jealousy regarding territorial rights exists
among the Vedda, Maidu, Shasta, and Thompson River Indians. In these ex-
amples, the group exists as a unit exclusive of kinship ties (Lowie 1961:
39h). A person was identified with a geographical location and not by his
position in the kinship network. For this identity system to work, the
allocation of territorial space would have to be rigorously'adherred to
or the system of identity would be dysfunctional.

Forde (1963:26-27) found that Bushman territory of the Kalahari
Desert serves to allocate permanent water sources. Trespassing across
tribal frontiers is dangerous unless previous relations were friendly and
the arrival is frankly announced. There is often a neutral zone between
the territory of groups to facilitate peaceful entry. According to Gearing
“almost all groups which act politically. . .possess or control territory;
but gypsies and EWestern] Shoshones do not" (Holmes 1965:228-229).

Human territorial behavior fits into the general context of primate
behavior. Human territory is present because of biological imperative of
food.but not because of reproduction, and it is also present because of
cultural directives. Culture and.not instinct is the deciding factor in
human territorialism. According to'Washburn and Hamburg (1965:615-617)
primate ranges commonly overlap and as far as is known, there are no de-
fended demarcated boundaries as is found in fish and birds. Familiarity
seems to be a major factor in limiting the group's activities to an area

of knowledge. The group learns the local conditions, the dangers, and the

Opportunities. De Vore (1963:307) says:

29
Clearly no definite areal boundaries are defended against groups of co-
species but organized groups do occupy distinct home-ranges and a strange
troop is rarely in core areas of another group which indicates spacing
mechanisms. Different groups are kept apart not by overt aggression and
fighting at territorial boundaries as by the daily routine of a group in
its own range, by rigid social boundaries of organized groups, and by loud
vocalizations.

In the nonprimates, territory is closely interrelated with the
breeding season. Territory is generally held by the male and its posses-
sion allows him to breed. It places a check on the possible number of
breeding pairs and is a selective factor weeding out the less fit. At
least in the fish and birds, behavior subsystems, e.g., defense and main-
tenance, seem to be largely instinctive behavior correlated with hormone
output. The territory possessor patrols and defends his territory but
only against actual invaders, proximity is not enough. There is no evidence
that primates exhibit any of this type of behavior in the field. I believe
that primates differ because of the combination of two factors, group life
and continuous sexual receptivity by both the male and female.

According to Ford and Beach (1951:13), sociologists recognize the
integrating, cohesive functioning of sex as contributing to the entire
structure of the social group. They (1951:199) have proved that primates
engage in a great deal of sexual play and showed many autoerotic, homo-
sexual, and heterosexual responses. Sexual games and erotic advances are
part of the prepuberal play of primates. Beach (1961:131-132) found that
in most primates, the adult male is sexually potent at all times. The
female primate may under special conditions exhibit sexual receptivity of
a low order at times when she is not physiologically in estrus. The non-
primates are characterized by distinct, usually brief, breeding periods.

According to Etkin (l96hdz76-87), a direct adaptation of ensuring

reproduction of a species is the almost universal limitation of spawning

30.

to one specific and limited time period. The integration of the mammalian
reproductive pattern into the seasonal and other environmental variables
is centered chiefly around the female. The larger mammals, particularly
in temperate or arctic climates, have only one brood per year or sometimes
one in two years, e.g., hear. The females and males are sexually active
only during a limited period (rut period). The estrus cycle consists of
short-termed physiological cycles containing a brief interval of sexual
excitement called heat or estrus. These cycles go on during the rut
period but cease during the rest of the year. According to Washburn and
Hamburg (1965:610), the commonest situation in primates is one in which
mating and births occur in all seasons, but in which there is a marked
peak time in a relatively restricted two or three month period. Beach
(l96h:39) sees three trends in sexual behavior when comparing rodents,
carnivores, monkeys, apes, and humans. There is a progressive decrease

in the extent to which sexual arousal and performance are dependent upon
gonadal hormones and an increase in the degree to which the cerebral cor-
tex influences and controls sexual expressions. The importance of learn-
ing and experience for the successful execution of the sexual pattern of
heterosexual union increases. The maintenance of a male held territory
for breeding would be destructive of the species if it were patterned
after either the kob or stickleback territory. Since the female and.male
can breed all year then this fighting for breeding territory would take
place all year. This would lead to the consequent exhaustion of the males
of the species and.make them easy prey for predators. But it would be
feasible if it were patterned after bird territory; However, the factor
of group life precludes that possibility.

Why does the group exist? The group is the locus of knowledge and

31
experience far exceeding that of the individual member. The group pools
experience and links generations. One biological adaptation that group
life confers is prolonged biological youth which gives the animal more
time to learn. During this period, the animal learns fromcather members
of the group and is protected by them. Slow develOpment in isolation
would simply mean disaster for the individual and extinction for the
species according to Washburn and Hamburg (1965:613). Ratner and.Denny
(l96h:h3h) believe that groups facilitate finding and.using large sap-
plies of food, function for the detection and defense against enemdes,
and enhance survival of the individual when the group is under severe
environmental conditions. The goup helps ensure protection and.surviva1
just as a territory does for nonprimates.

Free-ranging primates do not defend a locality. At most they de-
fend the integrity of their group in a way analogous to settling intra-
group dominance interactions that is similar to territorial defense
behavior in nonprimates. Those primates with the most aggressive intra-
group dominance system are also those who are most aggressive in either
physically or in some other way in repelling other groups of their Species.
Except for the urban macaque, which I contend is a Special immediate
adaptation to a severe environment,no primate defends boundaries but
rather defends his present location. In fact, the seeming defense of
boundaries by the urban macaque may just be the result that his territorial
space is so limited that its seeming defense is actually the defense of .
the group for food or some other vital commodity in a dominance situation.
The year around breeding season and the development of group life does not
permit holding the type of territory found among the nonprimates. The

localization of movement is a learned phenomenon resulting from the infant

‘32
learning from other group members.. Variations in this behavior are learn-
ed adaptations to a changed physical and social enviromnent.

The primate behavioral authors are not very precise in their use of
terminology. They use home-range and territory interchangeably. They
seemingly ignore the differentiating factor of defense of a territory or
they observe defense behavior but do not determine what is being defended.
This may indicate a lack of awareness with territorial theory or a lack
of data to decide which form existed.

I think that future research should attempt to rectify this condi-
tion. Laboratory work or observation under controlled conditions should
try to isolate the-variables which determine the form of territorial be-

havior in primates.

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