Applied Animal Ethology, 2 (1976) 181-185 181 o Elsevier Scientific Publishing Company, Amsterdam - Printed in The Netherlands

Short Communication

INTER-SPECIES INTERACTION DIFFERENCES IN PLAY ACTIONS IN CANIDS

MICHAEL W. FOX

Department of Psychology, Washington University, St. Louis, MO. 63130 (U.S.A.)

(Received July 27th, 1975)

ABSTRACT

Fox, M.W., 1976. Inter-species interaction differences in play actions in canids. Appl. Anim. Ethel., 2: 181-185.

Species-typical action patterns were identified in wild and domesticated canids during play in dyads of the same or different species. While these actions are under relatively rigid genetic control (i.e. are inherited), other dimensions of the play sequence may be more easily modified contextually. Thus not only species and context determine the frequency of occurrence of certain actions, but also the “compatibility” of the interactee, be it of the same or of a different species. The long-term consequences of inter-species interactions under conditions of captivity may lead to more permanent changes in the occurrence, frequency and amplitude of certain species-typical,“fixed”action patterns; this may be an additional variable to consider in investigating the complex influences of domestication upon behavior and warrants further investigation.

INTRODUCTION

Most studies of social behavior have focused upon intra-species interac- tions. This study involves a little explored area of inter-species social inter- actions, where the frequency of occurrence of clearly id.entified discrete action patterns may be modified in relation to the degree of dyadic compa- tability within the selected context of playful interaction.

Yves Rouget (personal communication, 1970) raised a red fox (Vulpes uulpes) with a domesticated dog; the impressive, but unquantified conse- quence which he recorded on film was the high frequency of face-oriented pawing manifested by the fox. Such an action is rare in foxes but is a com- mon action pattern in domesticated dogs. This kind of observation opens the question of the role of experience in the development and reinforcement of species-typical action patterns.

An incompatibility of reciprocal actions has been reported by H. Blauvelt (cited personal communication, 1960, by Davis, 1964) between sheep raised

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with goats: no effective fights were seen because the sheep tried to butt the goat and the goat tried to jump on the sheep. Interestingly, whom the ani- mal attacked was determined by early social experience, but the manner in which it attacked remained unchanged.

A consistent feature of social play in C. familiaris, F, C. familiaris X C. lupus and F, C. familiaris X C. latrans is the occurrence of the play leap (Fig.1). This action may be preceded by a direct stare, a play-soliciting bow or an incomplete leap intention which may be repeated three or four times. The leap itself varies in amplitude and may be repeated as the subject moves in on its congenor. The latter may also rear up and meet its partner with a reciprocal leap and both animals briefly stand and wrestle with the forelimbs and make face-, cheek- and scruff-oriented bites or incomplete bites. Most often, the leap is followed by an incomplete bite or bite intention or by a nose stab which may be interpreted as a bite-intention movement. Occasion- ally this may be followed by a backward leap.

In the coyote, C. latrans, such sequences involving the play leap are not observed. In this species, a play-soliciting bow is followed by side-to-side head flexions which move fluidly down the body, followed by rolling over, “spinning”, running around the partner, “diving” or exaggerated approach or withdrawal (Fox, 1971). From the play-soliciting bow, the head may be twisted upwards toward the forelegs, throat or cheeks of the congenor and bite-intention lunges or incomplete bites are then executed (Fig.2).

Fig.1. (A) Vertical leap of beagle during play which may be reciprocated (B) and both rear up. Leap may be followed by face or cheek-oriented bite. (D) playful standing over by F, coyote x beagle; (E) clasping by coyote during play fighting.

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An action resembling the play leap occurs when one coyote is chasing another; the action is of low amplitude and is followed by a bite or bite intention on the rump or shoulders of the partner, or a forelimb clasp (Fig.1). This clasp, which is often never fully executed, consists of seizing the part- ner around the waist and is frequently followed by a scruff-oriented bite. The partner may attempt to avoid being clasped by executing a hip-slam or twisting around to face its congenor. When facing, rearing up may occur, followed by face-oriented bites or bite intentions and pushing with the forelimbs.

MATERIALS AND METHODS

With these species-characteristic actions in mind, it was decided to evaluate the compatibility and reciprocity of such action sequences in variously com- bined pairs of subjects, by recording the frequencies of play leaps. All dyads were of opposite sex, each approximately 12 months of age. Each pair (of non-cage mates) was observed for 10 min play in an 8 X 8ft soundproofed arena equipped with one-way observation windows. Two observers recorded the frequency of play leaps for each pair, dyad combinations consisting of beagle X beagle, beagle X coyote, beagle X coyote-dog, coyote X coyote, coyote X coyote-dog, and coyote-dog X coyote-dog. A total of 20 dyad combinations were tested in 12 subjects (four beagles, four coyotes, four F, coyote-dogs, with two females and two males per species).

Fig.2. Although scruff-oriented biting in coyote (D) often occurs during play fighting, the coyote usually approaches low (A and B) and makes an upward-directed bite from the ‘C’ play-soliciting posture and rarely directs vertical leaps at its partner.

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RESULTS

The findings are summarized in Table I. It should be added that both coyote and coyote X dog showed hip-slams during play, but the frequency of this action was not recorded. Also several of the low amplitude play-leaps recor- ded in the coyote were difficult to distinguish from the incomplete action of rearing up to clasp the partner with the forelimbs.

The high frequencies of play leaps in pairs of beagles, coyote-dogs and beagle X coyote-dogs reflects the reciprocal nature of this action. In all dyads where one individual scored over 15 play-leaps, then its partner like- wise had a comparably high score.

TABLE I

Summary table of play-leap scores; average per individual and range in parentheses I_._

Subject With beagle With coyote With coyote-dog

Beagle 8.5 (2-16) 16.3 (l-37) 15.7 (2-35)

Coyote 1.0 (o-2) 2.5 (o-7) 0 (0) Coyote-dog 12.7 (1-31) 5.7 (4-9) 16.0 (7-22)

The beagles were more active when paired with a coyote or coyote-dog than with its own species and this may account for the lower scores in beagle pairs.

An intriguing finding was that the coyote-dogs had high frequencies of play leaps when paired with a conspecific or beagle, but lower frequencies when with the coyote. This was attributed not to a lowered activity or mo- tivation, for play was intense, but to the possibility that the hybrids had a more flexible or adaptive play repertoire which could be reciprocally matched with a beagle or coyote partner. In contrast, the beagle when paired with a coyote showed the highest average frequency of play leaps which was not reciprocated. No significant differences attributable to sex were evident in any of the three canid types.

DISCUSSION

These data support the general impression that the action sequences of beagle and coyote during social play are incompatible. The bow followed~ by upwardly directed bites and bite intentions of the coyote may be regard- ed as low ventral play attack, while the play leaps of the beagle as a high dorsal play attack. The coyote, in responding to the ventral low attack orientation, twists away and usually executes a shoulder or hip-slam to block or deflect the attack: such actions were not observed in the beagle, but were evident in the coyote-dogs and most probably contributed to their ability to maintain reciprocal synchrony with the coyotes during play bouts. These

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hybrids also showed the play-leap action which enabled them to sustain reciprocal play actions with the beagles.

The role of early experience and of social factors should also be consid- ered, and this point is emphasized here rather than earlier in this paper, with reference to the prior experiences of our canids. The criticism that might be leveled at this study to the effect that the beagles had no prior opportunity to interact and modify their actions with respect to coyotes (and vice versa) is untenable. All beagles and coyotes used in this study had been reared together from approximately 8 weeks of age, although tests were conducted using individuals from different cages. It might be expected that coyotes with such early social experiences might acquire the play-leap action of the beagle. But this was not evident, thus supporting the notion that the play actions studied are relatively fixed species-characteristic action patterns, the frequency and temporal sequencing of which may be modified contextually as in certain dyadic interactions.

Given that the interaction with a different species may modify the occur- rence and frequency of certain action patterns, more profound and long-lasting effects on inter-species interactions under conditions of sustained/enforced contact in captivity remain to be evaluated. While there may be little or no change in the “structure” (movement patterning) of certain action patterns, their occurrence and frequency of elicitation may be altered, especially within the broader context of domestication, as when different species (cat and dog, goat, sheep and cattle, cattle and horses, etc.) are raised and con- fined together. Similarly, the influence of human behavior on such actions remains to be evaluated. One classic example is the canid analog of the human “greeting grin” (which is quite distinct from the submissive grin and open- mouth play face of canids (Fox, 1971)). This facial expression is mimicked by the dog and has only been observed in man-dog interactions and not between dog and dog. There is also evidence that the capacity to elaborate this facial display is inherited since certain lines of dogs will never develop this behavior inspite of training, while others from the same kennel whose parents have this facial expression, will quickly develop it either spontaneous- ly or with little human reinforcement.

ACKNOWLEDGEMENTS

This research was supported by PHS Grant PlO-ES-00139, awarded through the Center for the Biology of Natural Systems, Washington Universi- ty, St. Louis, Missouri, and NSF grant GB-34172.

REFERENCES

Davis, D.E., 1964. The physiological analysis of aggressive behavior. In: W. Etkin (editor), Social Behavior and Organization Among Vertebrates. University Chicago Press, Chi- cago, Ill., pp. 53-74.

Fox, M.W., 1971. Behavior of Wolves, Dogs and Related Canids. Harper & Row, New York, N.Y., 220 pp.