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A Review of the Development and Functions of Cat Play

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A Review of the Development and Functions of Cat Play Applied Animal Behaviour Science 214 (2019) 1–17 Contents lists available at ScienceDirect Applied Animal Behaviour Science journal homepage: www.elsevier.com/locate/applanim A review of the development and functions of cat play, with future research T considerations ⁎ Mikel Delgadoa, , Julie Hechtb,c a Department of Medicine and Epidemiology, School of Veterinary Medicine, University of California at Davis, Davis, CA, 95616, USA b Department of Psychology, The Graduate Center, City University of New York (CUNY), New York, NY, 10016, USA c Department of Psychology, Hunter College, New York, NY, 10065, USA ARTICLE INFO ABSTRACT Keywords: Although attention to domestic cat (Felis silvestris catus) behavior and cognition has increased in recent years, Play behavior numerous questions remain regarding their play. Few studies have included play as a variable of interest, and to Play functions the best of our knowledge no behavioral studies focusing on cat play have been published in the last 15 years, Domestic cat and there is no recent review of our current understanding of its development, behavioral components, function, Felidae or outstanding research questions. This is despite the accessibility of the cat as a convenient model for more Predatory behavior difficult to study members of the Carnivora, as recognized by pioneering studies of cat play inthe1970sand Ethogram 1980s. We address this gap by reviewing and synthesizing the existing literature on play development, identi- fying and discussing eliciting factors and possible functions of play in cats. Additionally, we conducted an ex- tensive review of the literature to identify how play has been operationalized in peer-reviewed publications (N = 46). We identified 138 behaviors measured in these studies, with 84 of them unique behavioral labels.Our findings demonstrate the diversity—and sometimes commonalities—of descriptions of play behavior across these studies, while highlighting the challenge of inconsistent operationalization of cat play in the literature. We conclude by proposing and exploring several open questions and offering suggestions for future research, par- ticularly related to pet cats. “It would be quite absurd to assume that a cat, who in its daily cats. existence had already killed and eaten thousands of prey animals, Despite an increase in general empirical inquiry related to cats, and could not recognize the difference between a mouse and a ball of recent reviews of cat behavior and cognition (e.g., Vitale Shreve and paper; it knows that a ball of paper is not a mouse just as precisely as Udell, 2015, 2017), numerous scientific questions remain regarding cat it immediately recognizes the difference between a familiar anda play. Although a few studies have included play as a variable of interest new prey animal, or between a harmless and a potentially dangerous (e.g., Vitale Shreve et al., 2017; Ellis et al., 2017), to the best of our one.” – Paul Leyhausen, Cat Behavior, 1979 knowledge there have been no peer-reviewed behavioral studies fo- cusing on cat play in the last 15 years, and no recent review of our current understanding of the development, behavioral components, and 1. Introduction functions of cat play, framed within the context of their relationship to other wild felids and Carnivora. This is despite the accessibility of the Perhaps one of the most pervasive stereotypes about the predatory domestic cat as a convenient model for more difficult-to-study members behavior of domestic cats (Felis silvestris catus) is that they play with of Carnivora, as recognized by Bateson and Caro in their pioneering prey. This suggests that we believe we can recognize functional dif- studies (e.g., Barrett and Bateson, 1978; Bateson, 1981; Caro, 1979, ferences discriminating some interactions between cats and prey from 1980a; Caro, 1980b,c). This paper addresses that gap by reviewing the others—some behaviors appear strictly predatory, others not. This presently available research, by proposing several open questions and stereotype highlights the potential difficulty both in defining play and challenges related to cat play—including the problem of inconsistent in drawing a line between play and predation in a naturally predatory operationalization of play in the literature, and by suggesting future species, particularly given that cats display several forms of play out- research directions with a focus on pet cats. side the context of hunting, such as toward inanimate objects and other ⁎ Corresponding author at: VM: Medicine & Epidemiology, 1 Shields Ave., 2108 Tupper Hall, Davis, CA, 95616-5270, USA. E-mail address: [email protected] (M. Delgado). https://doi.org/10.1016/j.applanim.2019.03.004 Received 28 August 2018; Received in revised form 1 March 2019; Accepted 4 March 2019 Available online 08 March 2019 0168-1591/ © 2019 Elsevier B.V. All rights reserved. M. Delgado and J. Hecht Table 1 Operationalization of play behavior in 46 studies where play was an explicitly measured variable, exactly as defined in the referenced studies. Behavior Definition How measured Categorization References Allogrooming Licking the dog’s face or body. Duration and frequency Play behavior patterns (toward dogs) Feuerstein and Terkel (2008) Approach Movement of any sort (excluding canter) towards another cat. Frequency Social play Caro (1981b) Arch Each occurrence of a marked upward curving of the spine while standing still, Frequency Playful activity Barrett and Bateson (1978); leaping upwards, or moving sideways. The orientation is usually side-on in relation Bateson and Young (1981) to another cat or object. This combined West’s (1974) ‘Horizontal leap’ and ‘Side- step.’ Arch A marked upward bending of the spine while standing still, leaping upwards or Frequency Social play Caro (1979), 1980a moving sideways. Arch A marked upward bending of the spine while standing still, leaping upwards or Frequency Social play Caro (1981b) moving sideways, with or without piloerection. Attack Jump onto a cat and grasp it with forepaws or forelegs. Frequency Social play Caro (1981b) Bat Cat touches object with a front paw, claws retracted. Frequency Object play Hall and Bradshaw (1998) Bat Striking the mouse from above with the palm of the paw parallel to the ground. The Frequency Predatory behavior Pellis et al. (1988) mouse was moved or compressed by the strike, but the paw was quickly withdrawn. Belly up To move the back legs in a treading motion and to make reaching or pawing Frequency Social play West (1974) movements with the front legs. The mouth is held open and the teeth are exposed. In a social encounter, one kitten assumes the belly-up position and another kitten stands over it. Thus, the treading and pawing movements bring the kitten into contact with parts of the body of the standing kitten. Usually, these areas are the head, neck and ventral area. Bite Bringing jaws into contact with a cat and closing them. Frequency Social play Caro (1981b) Bite Bites are less forceful than killbites. Frequency Object play Hall and Bradshaw (1998) Bite The mouth was closed around the mouse. This could be subdivided into light bites, Frequency Predatory behavior Pellis et al. (1988) in which the skin was only grasped and the mouse uninjured, and heavy bites, in 2 which the skin was broken and a firm grip on structures beneath the skin was achieved. Canter Jerky running gait during which all paws repeatedly and simultaneously leave the Frequency Social play Caro (1981b) ground; head and tail often held high. Cat contact Each pat with a paw making contact with another cat and each bite of another cat. Frequency Playful activity Barrett and Bateson (1978); Bites could be given in the course of 'Wrestling' (see below) but were scored Bateson and Young (1981) separately. Frequencies of patting and biting were not scored by West (1974). Cat contact Each pat with a forepaw making contact with another cat (including the mother) Frequency Social play Bateson et al. (1981) and each bite of another cat. Cat contact Each pat with a forepaw, and each bite, making contact with another cat (mother or Frequency Social play Bateson et al. (1985a) sibling). Scored separately for each kitten. Cat contact The kitten paws, pats or bites a conspecific. Frequency Social play Bateson et al. (1990) Chase Running after a moving kitten. Frequency Social play Caro (1979, 1980a) Chase Running after a moving cat. Frequency Social play Caro (1981b) Chase Each bout of running after another individual/mobile object with the chased Frequency Social or object play Mendl (1988) Applied AnimalBehaviourScience214(2019)1–17 individual running away from the chaser for at least a distance of 1 m. Chasing Cat runs rapidly in pursuit of (modifier). N/A Solitary or social play Stanton et al. (2015)a Chase A chase involves a kitten running after or from another kitten. It could, perhaps, be Frequency Social play West (1974) differentiated into pursuit and flight. Chew Cat holds object in the mouth and chews it with rapidly repeated, small bites. Frequency Object play Hall and Bradshaw (1998) Chirping Making a sound similar to a high-pitched phone ring, tone often rises near the end. Duration and frequency Play behavior patterns (toward dogs) Feuerstein and Terkel (2008) Clutch Cat holds the toy close to the body with one or both front paws. Frequency Play Hall et al. (2002) Clutch Object is held close to the body with one or both front paws. Duration Object play Hall and Bradshaw (1998) Crouch Belly on the ground with all limbs by the side of the body, oriented and attentive to Frequency Social play Caro (1981b) a conspecific; back legs often treading. Exploration Visual regard, licking, sniffing and/or touching without manipulation of an object Duration Object play West (1977) with the paws or mouth while moving slowly toward or around an object. Exploratory and play Responses included looking at the string, approaching it, pawing or batting it, Frequency Object play Collard (1967) responses mouthing it, and pursuing and pouncing on it.
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