AN ETHOGRAM FOR BEHAVIOURAL STUDIES OF THE DOMESTIC CAT (F elis silvestris catus L.) by UK Cat Behaviour Working Group UNNERSITIES FEDERATION FOR ANIMAL WELP ARE ( UFAW ANIMAL WELFARE RESEARCH REPORT NO 8 AN ETHOGRAM FOR BEHAVIOURAL STUDIES OF THE DOMESTIC CAT (Felis silvestris catus L.) by UK cat Behaviour working Group Published by UNIVERSITIES FEDERATION FOR ANIMAL WELFARE _ The Old School, Brewhouse Hill, Wheathampstead Hertfordshire AL4 8AN England ISBN 0 900 767 90 1 ISSN 0956-1137 ( © UFAW 1995 workshop Participants (in alphabetical order) Dr. John w.s. Bradshaw Department of Biology, School of Biological Sciences, University of Southampton, Bassett crescent East, Southampton SO9 3TU Dr. sarah L. Brown Department of Biology, School of Biological sciences, university of Southampton, Bassett crescent East, Southampton SO9 3TU Mrs. Sarah E. cook Department of Biology, School of Biological Sciences, University of Southampton, Bassett crescent East, Southampton SO9 3TU Miss Katherine J. Durman Department of Biology, School of Biological Sciences, University of Southampton, Bassett crescent East, Southampton SO9 3TU Dr. Hilary N. Feldman sub-Department of Animal Behaviour, University of Cambridge, Madingley, Cambridge CB3 8AA Present address: Department of Integrative Biology, University of California, Berkeley, CA 94720, U.S.A. Dr. Gillian Kerby Wildlife conservation Research Unit, Department of zoology, University of Oxford, south Parks Road, Oxford ox1 3PS Dr. David w. Macdonald Wildlife conservation Research Unit, Department of zoology, university of Oxford, south Parks Road, Oxford ox1 3PS Dr. Sandra Mccune sub-Department of Animal Behaviour, University of Cambridge, Madingley, Cambridge CB3 8AA Present address: Waltham centre for Pet Nutrition, Freeby Lane, Waltham-on-the-Wolds, Melton Mowbray, Leics. LE14 4RT 2 Dr. Michael Mendl Department of Clinical veterinary Medicine, University of Cambridge, Madingley Road, Cambridge CB3 OES Present address: SAC, Genetics & Behavioural Sciences Department, Bush Estate, Penicuik, EH26 OOE Mr. warner c. Passanisi Wildlife conservation Research Unit, Department of zoology, University of Oxford, south Parks Road, Oxford ox1 3PS Dr. Ian Robinson Waltham centre for Pet Nutrition, Freeby Lane, Waltham-on-the-Wolds, Melton Mowbray, Leics. LE14 4RT Miss Deborah F.E. Smith Department of Biology, School of Biological Sciences, university of Southampton, Bassett crescent East, Southampton SO9 3TU 3 contents Acknowledgements 3 Preface 4 Definitions of Solitary Behaviour Patterns 5 Definitions of Social Behaviour Patterns 10 References 16 List of Plates 18 Acknowledgements We thank the Waltham Centre for Pet Nutrition for financial assistance with the three cat behaviour workshops held at the Universities of Southampton, Oxford and California at Davis. We are also grateful for their funding of the cost of producing the illustrations. We thank UFAW for publishing this document and in particular thank Roger Ewbank and Trevor Poole for their encouragement during the writing of this ethogram. 4 Preface This document arose from the co-operative efforts of ethologists from three British universities, Cambridge, Oxford and Southampton, and the Waltham Centre for Pet Nutrition, all of whom wished to provide a standard set of descriptions of domestic cat behaviour for their own use, and for those who might study this species in the future. The primary focus of many, but not all, of those ethologists was the recording of social behaviour, and this is undoubtedly reflected in the degree of "splitting" and "lumping" of behaviour categories of different kinds. Many of the descriptions are extremely broad, and some could even form the basis for ethograms of their own, for example those concerned with play and with feeding behaviour. None of the group who drew up this document is likely to use it without modification, nor should any future user feel that the categories cannot be moulded to suit the objectives of a particular study. The list is intended only to be a common starting point, but as such should help to alleviate the problems that can be caused by research workers using the same words to indicate different behaviour, and vice versa. The task of categorisation is, of course, left to the individual researcher. However, we have subdivided the full list of behaviour into either "SOLITARY", where the behaviour does not generally involve another cat, or "SOCIAL", where the behavioural act involves at least two individuals being present at the same time, or is observed only in a "social" context. The authors of this document are all those who participated in the workshops which were held on October 25th and November 30th 1990 and on July 24th 1993. All are (or have been until recently) actively involved in the study of cat behaviour and ecology, observing cats in environments ranging from artificially restricted areas, such as cages in animal shelters, through the whole range of cat-human associations, to the more natural free-ranging setting. However, it is acknowledged that the basis for many of the descriptions comes from Gillian Kerby's D.Phil. thesis (University of Oxford, 1987). 5 Definitions of Solitary Behaviour Patterns POSTURE ELEMENTS: LIE ON SIDE -(Plate 1) - Positioned fully on side, one side of the cat in complete contact with the ground. Head on side or extended. Paws extended. LIE HALF SIDE - (Plate 2) - Positioned on side with body but not head in complete contact with the ground. CROUCH - (Plate 3) - Positioned ·with ventrum and legs in contact with the ground, paws unfolded. LIE VENTRAL - Positioned with ventrum and legs in contact with ground, paws folded. LIE DORSAL - Positioned flat with back in contact with the ground. LIE CURLED - Positioned with body pulled into a compact circular formation; this can be further subdivided based on the degree of tightness. SIT - The pads of the front paws are on the ground with the front legs straight and the rump squarely on the ground. STAND - Positioned with just four paws in contact with the ground. TAIL MOVEMENTS: SWISH - A cat moves its whole tail rapidly from side to side. TWITCH - A cat abruptly moves part of its tail from side to side or up and down. SLAP - A cat quickly strikes its tail on the ground. QUIVER - A cat vibrates its tail while raising it vertically. EAR POSITIONS: BACK - (Plates 4 and 5) - Ears are held at the rear of the head. FORWARD - (Plate 6) - Ears are held at the front of the head. FLAT - (Plates 4 and 7) - A cat flattens its ears to its head, such that they tend to lie flush with the top of the head. ERECT - (Plates 5 and 6) - A cat points its ears upward. SLEEP Cat immobile, eyes closed and not easily disturbed. 6 REST Cat remains generally inactive with eyes closed but occasionally opens them to scan the area; ears flicking regularly. ALERT Cat remains generally inactive with eyes fully open and flicks its ears occasionally as it scans its surroundings. GROOM SELF Cat grooms itself by licking its body or by licking its paw and passing the paw over its head. (Subdivisions are possible according to region of body groomed, e.g. GROOM GENITALS). Grooming can include scratching and brief chewing of the body. OBJECT SCRATCH (Plate 8) Cat repeatedly scrapes its extended claws against a rough surface, e.g. wood. OBJECT COVER Cat makes movements with its front paws to cover, or as if to cover, any object apart from urine or faeces; e.g. food. DRINK Cat laps water or other liquid. LICK OBJECT Cat licks an inedible object. FEED Cat eats. FORAGE Cat hunts or scavenges for food (includes eating). (Hunting behaviour can be considerably subdivided - see Leyhausen 1979 and Panaman 1981). 7 DEFECATE Cat either scrapes a shallow pit or deeper hole in the ground or other substrate, or selects a spot on the surface of the ground by sniffing around, and then defecates. Faeces are either COVERED (cat covers, or acts as if to cover, faeces with soil, vegetation or any other suitable material) or UNCOVERED (cat makes no attempt to cover faeces). URINATE Cat either scrapes a shallow pit or deeper hole in the ground or other substrate, or selects a spot on the surface of the ground by sniffing around, and then urinates. Urine is either COVERED (cat covers, or acts as if to cover, urine with soil, vegetation or any other suitable material) or UNCOVERED (cat makes no attempt to cover urine). SPRAY (Plate 9) Cat directs a jet of urine backwards against some object. The tail is raised vertically and quivers as the urine is discharged (see Leyhausen 1979). PSEUDOSPRA Y Cat backs up to an object, tail is raised vertically and quivered, but no urine is emitted (subdivision of SPRAY, when distinction can be made reliably). SCRAPE IDND LEGS Cat (usually an entire male) scrapes its hind legs backwards rapidly, one after another, displacing vegetation and soil or other material. WALK Cat travels without obviously investigating its environment. EXPLORE Cat travels slowly, sniffing at objects and investigating its surroundings. 8 PASSIVE EXPLORE Cat moves its head repeatedly while sitting or standing, investigating its environment. PACE Cat repeatedly walks to and fro along the same route. CRAWL Cat moves with body in a crouched position. RUN Cat moves rapidly in a gait that is faster than walking. FREEZE Cat suddenly becomes immobile with body tensed. CLIMB Cat ascends an object, e.g. a tree or a fence. OBJECT REAR Cat stands up on its hind legs with its forelegs against an object. SNIFF Cat raises and twitches its nose, as if to smell. Subdivisions include SNIFF AIR (the nose is not near any particular object), SNIFF OBJECT (the cat smells/investigates an object with its nose), and SNIFF CAT SCENT (the cat investigates with its nose placed where cats are known to have scent marked [urine sprayed or cheek rubbed], urinated or defecated). PANTING Cat breathes deeply and quickly, with mouth open.