J. Zool., Land.

(1980) 192, 531-542

The ecology, structure and functions of social play

in Bighorn sheep (Ovis canadensis)
JOELBERGER*
Department of Environmental, Population, and Organismic Biology,
University of Colorudo, Boulder, Colorado 80309, U.S.A.

(Accepted 8 January 1980)

(With 1 plate and 1 figure in the text)

Social play was studied in Bighorn sheep (Ovis canadensis) by examining its structure in
two natural populations and by inferring functions through sexual differences in behaviour.
Lambs discriminated the size of lambs with which they played. During contact play they
responded most often to similarly sized individuals, whereas in locomotor play they chased
and followed all sizes. Males engaged in more contact-oriented play than females. This
evidence supports the idea that contact-oriented play developed in part to provide training
for combatant skills in males and locomotor play as part of a general anitpredator strategy
in both sexes. Social play was severely restricted in a desert environment due to risks and
personal injury. The implications of lack of play in natural populations are discussed.

Contents
Page
Introduction .. . . . . . . .. .. .. .. .. .. 53 1
Methods . . .. . . . . .. .. . . .. .. .. .. 532
Study populations .. . . . . . . . . . . .. .. .. 532
Definitions . . .. . . .. . . . . . . .. .. .. 533
Data collection and analysis .. . . . . . . .. .. .. 533
Results .. .. . . . . . . . . .. .. .. .. .. 534
Familiarity and the development of playful interactions .. .. .. 534
Play partner preferences . . .. .. . . .. .. .. .. 534
Structure of play sequences . . .. .. .. .. .. .. .. 535
Play signals . . . . . . . . . . . . .. .. .. .. 535
Ecological aspects of social play . . . . . . .. .. .. .. 536
Discussion . . . . . . .. . . . . . . . . .. .. .. 537
Development and communication of play . . . . . . .. .. .. 537
Functional and evolutionary aspects of social play .. .. .. .. 537
Ecological aspects of social play . . . . . . .. .. .. .. 539
References . . . . .. .. . . . . . . . . .. .. .. 541

Introduction
Play behaviour is often called a paradoxical activity because the immediate contributions
to a player's fitness are not obvious while a great deal of energy and personal risk are
expended at an early age (Bekoff, 1976; Fagen, 1974, 1977). Play behaviour in mammals
has become the subject of increased attention by ethologists and zoologists and it has also
'Present address: Conservation and Research Center, National Zoological Park, Smithsonian Institution,
Front Royal, Virginia 22630, U.S.A.
531
0022-5460/80/120531+ 12 002.00/0 @ 1980 The Zoological Society of London
532 J . BERGER

generated interest among scientists investigating the evolution of behaviour (Alexander,

1977; Wilson, 1975). Recent studies of mating systems have contributed greatly to our
understanding of the functional significance of sex differences in behaviour (i.e. Brown,
1975; Fisher, 1930; Symons, 1978) and this knowledge may now be applied toward
understanding the evolution of social play.
In species of mammals where intrasexual selection has been great (i.e. some pinnipeds)
aggressive behaviour is more intense among adult males than adult females (Le Bouef,
1974). Since behaviours comprising an individual’s behavioural repertoire are established
and refined during its ontogeny (Gentry, 1974; Reiter, Stinson & Le Bouef, 1978; Symons,
1978), differences in behaviours used in accordance with male and female reproductive
strategies should be expected. Therefore, sex differences in infant playful and aggressive
encounters could also be predicted.
A group of mammals ideally suited to examine this prediction are sexually dimorphic
species of polygynous ungulates. Adult males have greater variance in reproductive success
and they behave differently than females (Geist, 1966, 1974; Walther, 1974). Males compete
intrasexually for mates, as inferred through the presence of secondary sexual characteristics
(horns, antlers, tusks) and indicated by the relatively high frequencies of butting, shoving
and outright fighting. Conversely, secondary sexual characteristics are not well developed
in females and dangerous fighting is not as frequent. Females are not in direct competition
with one another to be fertilized. Quite obviously the reproductive strategies and associated
behaviours of males and females differ. A comparison of sex differences in the behaviour
of adults and its ontogeny in juveniles may cast light on functional interpretations of social
play.
During a previous study (Berger, 1979a) of the social ontogeny of Bighorn sheep (Ovis
canadensis)it was found :(1) differences in the use of motor actions by male and female lambs ;
(2) these differences were accentuated as lambs grew older; and (3) the frequency of specific
motor actions (e.g. rotational) differed between individuals inhabiting desert and mountain
environments. In this paper the following hypotheses are examined in an attempt to explain
the evolution of social play in Bighorn sheep : (1) rotational movements communicate
play intention; (2) changes occur in the structure of play sequences in sheep of differing
ages and sexes; (3) males, at any age, engage in more contact-oriented play than females.
Lastly, the social ecology of play, or “where play occurs” is considered. Several authors,
(Altmann, 1956; Byers, 1977; Darling, 1937; Symons, 1978) have pointed out that play
occurs most often in certain physiographical locations. Since mountain (and northern)
habitats differ topographically from desert ones, the present author was interested in
determining whether, and how, habitat complexity affected the location(s) of social play.

Methods
Study populations
Sheep were studied in natural environments from May 1976 through August 1977. One popula-
tion was located in the Chilcotin-Caribooregion of the interior of British Columbia.The population
size of California Bighorn sheep (0.c. californiana) was estimated at 400 (Berger, unpubl.). Two
prominent features in this habitat are gradually sloping grassy hills (e.g. bunchgrasses) and several
large (c. 400 mz) sandbowls located at the base of these hills (see Demarchi & Mitchell, 1973 for
a further description). Sheep were observed in this environment for 896 hours. A second popula-
tion was studied in the Santa Rosa Mountains, an arid insular range in the Colorado Desert,
 SOCIAL PLAY I N B I G H O R N SHEEP 533

California. The present studies concentrated on about 80 desert Bighorn sheep (0.c. cremnobates)
in the Deep-Carrizo Canyon region. Prominent habitat features here included canyon walls (some
over 500 m high), broken rocky, craggy areas, and large abundant concentrations of cacti (Opuntia
spp.) and century plants (Agave spp.). Sandbowls and grassy slopes were completely absent from
this habitat. Climatological data were recorded directly in the Santa Rosa habitat based on
apparatus established at 900 m (see Ting & Jennings, 1976). Sheep were observed in the desert
habitat for 454 hours.
The play of captive Rocky Mountain (0. c. canadensis), California, Stone’s (0. dalli stonei),
and Dall(0. d. dalli) sheep were also studied for seven days at the Okanogan Game Farm, British
Columbia. Additionally, a newborn male desert lamb was captured and used for experimental
observations. At three weeks of age he was exposed to a two-and-a-half week old female desert
lamb for three consecutive days. The female lamb had been raised previously with its mother,
but both were isolated from all conspecific contact. The introductions were performed at the
Boyd Deep Canyon Desert Research Station, Palm Desert, California.

Definitions
Numerous terminological problems are inherent in any definition of play (see Bekoff, 1976;
Poole, 1978).For the sakeof clarity, however,a definition of play will be offered only as it relates to
Bighorn sheep. Social play will be operationally defined as erratic and often jerky (torso) body
movements that include components of butting and rotational behaviours. Running activities
were considered to be play if they occurred without obvious attempts to avoid conspecifics or
predators. This working definition encompassed activities of lambs, yearlings, ewes, and rams.
Contact patterns in play included butting, pushing, and touching heads or pushing other portions
of the body. Rotational body movements included neck twists, gambols. and heel kicks. Loco-
motor play included running and chasing movements without contact or rotational patterns. A
more thorough description and ethogram are found in Geist (1971) and Berger (1979~).In
contrast to play, aggressive interactions (which include the contact patterns mentioned above)
are attempts to supplant or dominate conspecifics. They lacked jerky (torso) body movements
and role reversals (see below: play signals).
During dyadic interactions between young lambs, I designated those from 1 to 10 days of age
as L1 lambs; 11 to 20 days old as L2 lambs; 21 to 30 days old as L3 lambs; and those older than
30 days as L + + lambs. Ages of lambs were estimated visually by their size in relation to ewes or
smaller known-aged lambs. During interactions in groups larger than dyads it became increasingly
difficult to define ages as precisely as stated above. Therefore, the identities of interactants used to
analyse the structure of play sequences (Fig. 1) were defined as follows: (1) lambs (=male and
female young less than three months of age); (2) male non-lamb groups (=male yearlings and
Class I rams; the latter being two to three years old: see Geist, 1971); and (3) female non-lamb
groups (=female yearlings and ewes). Only data from undisturbed natural populations were
analysed.

Data collection and analyses

Observations were made directly with a spotting scope and recorded directly into a cassette
deck. Fifty-five play sequences were recorded in the Chilcotin (N=27 for lambs; 16 for female
non-lamb groups; and 12 for male non-lamb groups); these ranged in duration from 20 seconds
to 28 minutes and from 10 to 401 acts. The latter occurred in the 28 minute sequence and it was
not continuous. Play sequences also occurred between mixed age groups and once it involved at
least 95 different individuals at the same time. It was far from possible to record verbally all
players and their subsequent actions when groups were so large. Consequently, 11 incomplete
sequences were omitted from the analyses.
534 J. BERGER

The structure of play sequences was analysed by recording the sequential order of motor
actions and the number, ages, and sexes of players. The frequency distribution of motor acts
during play sequences were analysed by casting them into those segments comprising the first
one-fifth of a sequence, the second one-fifth, etc. From this, it was possible to assess whether or
not various motor acts occurred randomly throughout play sequences (modified from Bekoff,
1976).
All statistical analyses of the differences between percentages (proportions) were performed
according to the Brandt and Snedecor method when there were at least two samples (Snedecor,
1956) and the arcsin transformation for testing the equality of two percentages (Sokal & Rohlf,
1969). This latter method generates a test statistic ( T , ) that may then be compared with the area
under the normal curve.

Frequency distribution of patterns

FIG.1 . Frequency distribution of behaviour patterns occurring during the play sequences of different aged Chilcotin
sheep. Horizontal axes represent the temporal distribution of acts from the beginning to the end of sequences.
Thus, 0.2 indicates those acts occurring during the first one-fifth of the sequence. Vertical lines indicate standard
errors. Each group has been staggered so that s.e.’s are non-overlapping. The total number of acts recorded were:
lambs (a), 1103; male non-lamb groups (w), 356; female non-lamb groups (o),460.

Results
Familiarity and the developrnent of playful interactions
Ewes in natural populations normally introduce their lambs into nursery bands when
the lambs are three to seven days old (Geist, 1971). During the first few days of association
between new lambs, the motor patterns most frequently observed were chasing and
following movements. The youngest new lambs that headbutted and pushed were about
two weeks old and they had already associated with “resident” lambs for about four days.
Familiarity with other lambs appeared to be a more important variable than age alone in
affecting the development of contact patterns. For instance, two captive three-week-old
desert lambs, that previously were unexposed to any other lambs, interacted for over 300
minutes over several days by chasing and following the other. Although these lambs used
locomotor patterns on cliffs and rested not more than five metres apart, they never engaged
in contact patterns. I n the study populations, lambs younger than three weeks of age
butted, pushed, chased, and mounted one another.
Play partner preferences
Lambs demonstrated age preferences during play depending upon the type of playful
 S O C I A L PLAY I N B I G H O R N SHEEP 535

activity. For instance, during contact play lambs initiated successfully more encounters
with equal-aged lambs than they did with disparately aged ones (Table l(a)). However, no
age or size preferences existed during locomotor and running play. In these activities lambs
responded similarly to the ambulatory movements of all aged lambs (Table I(b)).

T A B L EI
Interaction probabilities of different-aged Chilcotin lambs during contact and locomotor
Play *
A Contact (N=214) B Locomotor ( N =190)
Initiator Initiator

Recepient L1 L2 L3 L + + Recepient LI L2 L3 L + +
L1 0.58 0.10 0.18 0.07 L1 0.93 0.70 0.76 0.93
L2 0.06 0.50 0.28 0.18 L2 0.70 0.86 0.85 0.89
L3 0.25 0.21 0.56 0.41 L3 0.78 0.82 0.93 0.78
L+ + 0.06 0.24 0.25 0.54 Lf + 0.99 0.94 0.97 0.95

*Age classes of lambs are discussed in Methods. For each cell P = a / b ; where P= the
proportion of individuals interacting, either by engaging in contact behaviour or
following for all bouts; a=any lamb that interacts during a bout: and b=all lambs
obviously in the vicinity of a bout and capable of interacting in it. (Lambs resting were
judged to be capable of interacting.) Thus, the maximum value for any individual cell is
1.00. For contact patterns x*=280; df=9; P<O.Ool; for locomotor patterns, x2=8.39;
df= 9 : NS

Structure of play sequences

The frequency distribution of motor acts during play sequences are illustrated in Fig. 1.
For all groups, running and rotational movements occurred more frequently than contact
patterns when play sequences began. Running decreased throughout the sequences in each
group, but in lambs it increased significantly more toward the end of sequences than in
non-lamb groups (T,= 2.30; P <Om05 : Fig. 1). In contrast, rotational movements were
more evenly distributed for lamb and male non-lamb groups. However, in female non-lamb
groups these motor actions occurred significantly more in the middle of play sequences
than in the other groups (T,= 2-56;P <Om01 ; only one test was necessary since male rota-
tional play occurred less). Contact patterns occurred more in male non-lamb groups than
in the other two groups (T,= 1.99; P <0.05).

Play signals
The motor patterns that appeared to be play signals were rotational movements. They
occurred regularly throughout the play sequences of all sex and age groups (Fig. 1).
However, rotational movements were conspicuously absent from some dyadic interactions
among lambs, although these interactions lacked supplantations and other movements
associated with aggression. Consequently. it was difficult to interpret directly the presumed
function of these signals in lambs. Therefore, play signals were investigated in yearlings.
The frequency of yearling interactions was recorded when rotational movements were given
and supplanting occurred vs. the number of times that no supplantations occurred (i.e.
the interaction continued; Table 10.The rationale for analysing signals in this manner is
536 J . BERGER

as follows. (1) If play intention is transmitted, then players will continue to interact under
the auspices of a play “atmosphere” (see Altmann, 1967; Bateson, 1955; Bekoff, 1975).
(2) If play intention is not communicated, then subordinate individuals should retreat
quickly. (3) I n intermediate situations when dominance is not clear and play intention is
not transmitted. interactions should persist until a winner is established. Thus, the presence
or absence or rotational movements and supplantations may be used as a criterion of
making inferential judgements about the “meaning” of play signals. And, as is shown in
Table IT, rotational movements communicated play intention.

Ecological aspects of social plaji

Several variables affected the occurrence of social play. In the Chilcotin topographical
features were conducive for play and large bands of sheep (that included at the same time,
lambs, yearlings, ewes, and rams) played on grassy slopes and often terminated their playful
activities after running into sandbowls (Table 111). Play also occurred on cliffs. The desert
lacked grassy slopes and sandbowls and play was less frequent.
Lambs fron one to ten weeks of age were observed in the Chilcotin and desert environ-
ments for 4,706 and 1084 lambs-observation hours, respectively. During that time,
Chilcotin lambs played significantly more often than did desert lambs (x2= 12.64; df = 1 ;
P < 0.001 ; 424 vs 59 minutes; data were cast into frequencies by recording their occurrence
at 5 second intervals).

I1
TABLE
Tlic responses of’ Cllilrotin yearling Bighorns torational (“play”) signols rhot
occurred during playful and aggressive interactions.*

Supplanting occurs Supplanting does not occur

Signal present 9 78
Signal absent 38 10

*Chi square test: P<O.OOI ; Numbers indicate the frequencies of different

bouts between different individuals.

TABLE
111
Loctrriorr nrirlfiec~iiericie.sof play seqrretri.es in Chilrotin Bighorn sheep*
~ ~ ~ ~ ~

Locat ions
Sandbowls Hills Cliffs
Aggressive Play Aggressive Play Aggressive Play
Sex and age group N acts acts N acts act? N acts acts

Lambs 9 0 160 10 0 275 8 0 150

Female non-lamb groups 7 2 94 x 10 147 I 3 16
Male non-lamb groups 5 I 75 4 6 118 3 6 49

* N = number of playsequencesobserved. The numbers given for aggressive and playful acts indicate the number
of motor actions recorded.
 S O C I A L P L A Y I N B I G H O R N SHEEP 537

Ambient temperature was another variable that influenced play. In the desert no
instances of lambs play occurred when ambient temperature was greater than 32°C and
88% of lamb play occurred below 26°C. Of the 59 minutes that desert lambs played, 57
min occurred before 17 March. Nevertheless, mean maximum daily temperatures were
below 26°C until mid-April. Thus, most of the play in the desert was restricted to a period
of time prior to 17 March, although ambient temperatures did not appear sufficiently high
to preclude play for almost another month.

Discussion
Development and communication of pluy
Lamb familiarity with one another is an important factor influencing the development
of contact play. Geist (1971) found lambs in nursery bands butted and mounted, but the
prior duration of lamb associations were unknown. It appears, however, that unfamiliar
lambs must associate with, run, and chase their peers before they gain enough familiarity
(or “confidence”) to play by using contact patterns. For instance, newborn lambs that
accompanied their ewes to nursery bands were observed on 14 occasions. Yet, these new
lambs never played with resident lambs by using contact patterns. Only locomotor and
running play occurred at this young age. In contrast, young lambs that had associated with
resident lambs (see Results) used contact patterns. Additionally, a newborn lamb that
became imprinted on me, headbutted and mounted me after only four days of association.
It therefore appears that familiarity affects the development of contact patterns more than
age does.
Prior studies of ungulates have not emphasized quantitatively the role of familiarity nor
play signals in the development of social interactions (see Gilbert, 1968; Miller, 1975;
Muller-Schwarze, 1968; Walther, 1964). Rotational movements were clearly implicated
as play signals (Table 11) for Bighorn sheep yearlings, but they were not always present
when lambs played. These signals may be important during the playful interactions of
older, larger-horned sheep so that individual risks associated with injuries are minimized.
Conversely, play signals may not be as well developed in young lambs since they have no
horns and injuries resulting from butts are probably rare.

Functional and evolurionary aspects of socitrl play

The study of function in evolutionary biology may be approached in several ways
(Williams, 1966). Ideally, controlled experiments should be performed over several
generations so that differential gene survival may be documented and competing
hypotheses may then be rejected (Hinde. 1975). Unfortunately, this approach is usually
not possible and inferences about function are often drawn after making intra- or inter-
specific comparisons or by studying sex differences. These latter methods frequently fail to
separate cause from effect, but they nevertheless may provide data supportative of a
particular hypothesis. For instance, Synions (1978) recently addressed the problem of
function in his superb study of play in free-ranging Rhesus monkeys (Macaca mulatta).
Based on his knowledge of sex differences in behaviour and mating strategies, Symons
observed coincident differences in juvenile play. He then argued, quite convincingly, that
5.38 J. BER G ER

social play evolved to provide “practice” in fighting and to improve upon skills used i n
predator avoidance.
If combatant skills in adult male Bighorn sheep are developed through training, it could
be predicted that young (or infant) males would engage in more contact-oriented play than
females. Figure 1 showed that males in non-lamb groups participated more in contact play
than females. Elsewhere, I have demonstrated that male lambs, male yearlings, and Class
I rams from populations inhabiting desert, semi-desert, and mountain environments
engaged in more contact, threat, and display behaviours than females from similar age
groups and environments (Berger, 1979~).Geist (1968, 1971) found that rams of varying
ages also used more “overt” patterns than females, and that during playful “huddles”
subordinate rams clashed with dominant ones. The above evidence indicates clearly that,
regardless of age, sex differences exist in play structure. The occurrence of contact-oriented
play at significantly lower frequencies in females further suggests the relative unimportance
of these behaviour paderns in the mating strategies of females.
Existing data on other sexually dimorphic ungulates support the hypothesis that young
males engage in more behaviours related to dominance than females. General studies of
Reedbuck (Redunca arundinum) (Jungius, 1971), Pronghorn (Antilocapra anwricana)
(Auntenrieth & Fichter, 1979, Thomson’s gazelles (Gazella rhomsonii) (Walther, 1978),
Greater kudu (Tragelaphus strepsiceros) and Sitatunga (T. spekei) (Walther, 1964), and
Mule deer (Wachtel et al., 1978) have suggested that juvenile males interact more often
with aggressive behaviour patterns than females. Detailed studies of young Ibex (Capra
ihes) (Byers, in prep.), bovine calves (Bos indicus) (Reinhardt et a/., 1978), and domestic
sheep (Sachs & Harris, 1978) confirm that behavioural dimorphism occurs at a young age.
A number of other, and not necessarily competing, explanations have been offered for
the playful activities of young ungulates (see Fagen & George, 1977). Fagen (1976) lists
many of the presumed physiological functions of non-social play. Sachs & Harris (1978)
suggested recently that the temporal peaks of play in juvenile Domestic sheep (0. arks)
may be an ancestral trait coinciding with migration patterns in their wild congenerics,
Bighorn sheep. However, they failed to note that many populations of native desert and
mountain Bighorn sheep are non-migratory and their data were based on 20 minutes of
play per week. Nevertheless, Sachs & Harris demonstrated that male and female play
differed.
In Bighorn sheep lambs, locomotor and rotational movements are also important
components of play (Fig. 1) and they occur most often during group play. If locomotor
play in young sheep improves, in part, upon social cohesiveness or synchronized running
movements that are useful during predator avoidance behaviour, then components of
locomotory behaviour should also occur during adult anti-predatory activities. That is,
locomotory behaviour should occur in play or in actual flight without regard to age or sex
differences. This idea is supported by the fact that play partner preferences were uncommon
in the locomotor sequences of lamb play (Table I) and individuals at any age were likely
to follow any other infant. This clearly was not the case during contact play (Table I),
where lambs selected equal-aged or -sized partners for interactions. The lack of choice for
equal-aged individuals during locomotor play is quite similar to the flight of adults in
groups. When one member runs, many, if not all, follow (Berger, 1978). Meyer-Holzapfel
(1956) also suggested that running and following movements of young ungulates during
play serves a later function in terms of predator avoidance.
 SOCIAL PLAY IN BIGHORN SHEEP 539

Ecological aspects of social play

Darling (1937) first suggested the existence of “playgrounds” in Red deer (Cervus
elephus) fawns. Certain hills were described as areas where play was more likely to occur.
For Chilcotin lambs, sandbowls and grassy slopes also served as playgrounds. These areas
were sparsely located yet more play occurred there than in other area. In the Sierra Nevada
Mountains of California, where there are no sandbowls, Bighorn lambs played frequently
on or near glaciers and icefields (Wehausen, pers. comm.). Altmann (1956) also reported
juvenile Elk (C. canadensis) played in snowfields. It is plausible that certain physical
features may be attractive as playgrounds and that play may actually be facilitated in these
areas.
An interesting problem is that play sequences occurred infrequently in the desert whereas
they occurred significantly more often (see Results) in Chilcotin lambs. Geist (1971),
Horejsi (1976) and Shackleton (1973) argued that Bighorn sheep populations of low quality
are characterized by individuals that are stressed energetically and therefore they play less
than those individuals from higher quality populations or those found under better
nutritional regimes (see Baldwin & Baldwin, 1976). Since Chilcotin lambs played more than
desert lambs, it would follow that they may have been in superior physiological condition.
However, desert lambs consumed more milk at an early age, matured sexually earlier in
life, and almost certainly were better nourished (Berger, 1979b). Why could it be then, that
lambs in the Santa Rosa Mountains played less than lambs in the Chilcotin?
Play occurred frequently at early ages in lambs from both study populations, but it
then tapered off in frequency in the desert environment. In fact no locomotor play occurred
after mid-March in this environment. One reason for the decrease in play may be that
players in the desert faced certain hazards. For instance, not only did my study area in the
Santa Rosa Mountains lack playgrounds (e.g. sandbowls and grassy hills), but much of the
area was populated heavily with cholla cacti (Plate I). On 10 occasions, all before 17 March,
I observed playing lambs step on or into cacti. (Eight occurrences were between 10 and
16 March.) Not surprisingly, chunks of cholla inflicted great pain in lambs. Play ceased
immediately and lambs jumped frantically, kicked, and tugged at the spines that remained
in their skin on their heads, faces, and legs. Desert ewes and yearlings never engaged in any
locomotor sequences. In contrast, Chilcotin lambs sustained very few external injuries
during play.
A number of reasons other than hazards could explain why play occurred infrequently
in the desert. Perhaps heat stress (or the avoidance of it) caused desert sheep to minimize
energy expenditure through a reduction of play activity. This alternative seems unlikely
though, because (as mentioned previously) lambs tenninated playing by mid-March, a
time period after they had been exposed to cacti, and before ambient temperatures pre-
sumably affected play activities. Another possible explanation for the lack of observed
play may be that it occurred at night. Again, this explanation would seem unlikely since
my observations included crepuscular periods. If sheep played at night it seems reasonable
to expect that some of their play would carry over into crepuscular periods.
It appears, then, that lambs in the desert experienced negative feedback (painful wounds
from cacti) when playing. No obvious restriction limited play in Chilcotin lambs. In
essence, suitable playgrounds were in short supply in one environment and not the other.
If social play is an important element of training or practice, then a logical prediction
would be that individuals in populations that play more socially will also be behaviourally
SJO J . BERGER

more diverse. That is, those individuals with greater play experience should utilize different
behaviour patterns from their repertoire more often than individuals playing infrequently.
This prediction was supported for Chilcotin sheep (Berger, 1979a). Greater H values
(measures of behavioural diversity) existed for Chilcotin lambs, male and female yearlings,
ewes. and Class I rams than for individuals in the desert population.

PLATEI . (a) Example of Lower-Sonoran habitat of desert bighorn sheep in the Santa Rosa Mountains, California.
(b) Note the abundant amounts of cholla cacti and agave, which reduced play behaviour.
 SOCIAL PLAY I N BIGHORN SHEEP 541

I thank Marc Bekoff for advice and support throughout this project. I also thank the following
people for their comments on prior copies of this paper: David Armstrong, Ruth Bernstein, Carl
Bock, Valerius Geist, Ingrid Porton, Melody Serena, Peter Stacey, and Fritz Walther. Logistical
aid was offered by: Vernon Bleich, Eldon McLaury, Harold Mitchell, Stephen Walker, and John
and Ann Walsh. The Boyd-Deep Canyon Research Station (University of California), California
Department of Fish and Game, United States Fish and Wildlife Service, American Museum of
Natural History, British Columbia Branch of Fish and Wildlife, Sigma Xi, and the University of
Colorado provided financial assistance.

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