Feline Territorial Behaviour
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Home Range
The home range is the main area of territory that encompasses the resources that the cat needs for survival, inducing latrine locations, resting sites and hunting areas. Beyond this home range, the wider territory controlled by the cat, or cat group, may be very large. Feral and wild cats may hold territories that are more than 1-2 square miles. The territory may be formed from a patchwork of spaces that the cat can only access by traversing areas that are not its own territory. A cat may regularly pass close to, or through, another cat's territory in order to access parts of its own territory. Claw and urine marks are therefore used not only to signal the boundary of a territorial space but also the timing of the resident cat's presence in a particular location. In this way, cats can maintain distance from each other in time and space and thus avoid conflict. In an outdoor space this works well, as there is enough space for the different types of odour marks to be deposited in a meaningful way that allows the cats to avoid potential competitors. It does not work effectively inside a domestic home, where space is limited and physical barriers such as windows limit perception of deposited scent marks.
The size of the territorial range of cats (home range) varies between feral and pet cats, male and female cats and neutered and entire cats. Although evidence regarding the absolute size of home ranges is inconsistent between studies performed in different countries and habitats, intact male home ranges are, on average, three times larger than intact female home ranges [1]. The same author found that pet females in Sweden had home ranges of around 30-40 hectares and rarely roamed further than 600m from their homes[2], and feral females had home ranges that were about four times larger than those of pet females[3]. It appears that male home range size is determined by the availability of reproductive opportunities, whilst that of females is governed by the availability of food. Range size has been found to vary between 0.1 hectare in a Japanese fishing village to 170 hectares in the Australian bush[4], indicating that cats do not have a specific need to maintain a large territory, only to hold sufficient territory to satisfy specific survival needs.
The home ranges of neutered male and female domestic pet cats are likely to be smaller than their feral counterparts, and in one UK study were found to be 0.45 and 0.27 hectares respectively[4]. This reflects the high-density food resources, the close proximity of hunting, resting and latrine locations, and competition for space between the numerous cats in a typical suburban area. Home range size appears to be inversely proportional to population density[4].
Core Territory
Within the home range, the cat will also have a smaller core territory that is primarily used as a secure site for resting, feeding, and self-maintenance behaviour. In wild cats this core territory is typically around 100m in diameter and makes up 0.2 to 4% of the total home range[5]. This core territory is away from direct view or intrusion by other cats that are not part of the social group.
Facial and flank marks are deposited in core territory where a cat does not anticipate meeting unfamiliar cats. Scent marks are not only a signal to other cats, but also carry meaning for the depositor. In the core territory, cats are more likely to be in a parasympathetic state of arousal, in preparation for activities such as resting, feeding and grooming. Outside the core territory, some degree of sympathetic arousal is always required, as the individual is either vulnerable or engaged in hunting. Facial and flank marks may provide a chemical signal to the cat that favours a parasympathetic state in all individuals that are present, reducing the likelihood of conflict in colonies where several cats share a common core territory.
Territory Defense
The extent to which female cats will defend their home range may relate to the abundance of food resources. Work by Foley et al (2005)[6] and Driscoll et al (2009)[7] showed that the studied cats defended their territories, whilst a study by Corbett (1979)[8] identified a lack of competition between cats due to the reliable excess of food in the abandoned farms on the Scottish islands where the study took place.
Conflict is avoided by the use of scent marks and cats following strict timetables for their movements around their territories. Urine spray marks provide information about the time of day the area is being used by a specific individual so that other cats may traverse an area at times when the resident is not present, and claw marks are used to indicate firmer territorial boundaries that discourage intrusion by non-resident cats.
Scent marks contain information about the purpose of the mark, as well as about the depositor. This includes information about the health, sex and reproductive status of the depositor. Common chemical signatures between the claw, urine and facial/flank marks of an individual enable other cats to identify the at that deposited a particular make. When combined with knowledge of previous encounters with that individual, other cats are able to assess the level of risk associated with entering a particular area.
Domestic Cat Territory
The natural organisation of territory in cats poses some problems for domestic pet cats. In a domestic setting, the difference in area between the home range and core territory may be minimal, and for indoor-only cats that have a view of a garden, the boundaries of the indoor space are the absolute limits of territory, leaving no opportunity for conventional territorial organisation or distance maintenance from neighbourhood cats that are easily visible from indoors.
Pet cat groups are made up of unrelated and neutered males and females with widely differing rearing backgrounds. Some may come from a genetic and rearing background that does not favour sociable living in a group. Owners expect that the cat’s core territory will match the internal living space of the home, so that facial and flank marking are seen indoors and spraying or claw marking is only performed outdoors. However, instead of being one large contiguous area, each domestic cat’s territory may consist of several small patches that are distant from each other. Each cat is forced to travel across several other cat’s territories in order to get to a latrine or hunting site. This increases the amount of feline traffic through gardens and increases the likelihood that each cat’s core territory will be overlooked by cats outside. Underfed, despotic or intact male cats may enter the homes of resident cats, which further undermines the perception of the home as ‘core’ territory. These issues may lead to indoor marking behaviour that becomes problematic for the owners.
In urban areas the density of cat populations may be high, exceeding 50 cats per square kilometre. In a survey conducted by Bowen (2013), 81% of 734 UK cat owners whose cats were allowed outdoor access indicated that their neighbours also had at least one cat that was allowed outside, and 66% reported seeing a neighbour’s cat in their garden at least once a week. Owner reports of the number of different cats regularly seen in the garden and the frequency of cats visiting the garden were both correlated significantly with the frequency of injuries due to conflict with non-resident cats.
Forty-one percent of those households reported some level of home entry by neighbourhood cats, with 18.7% reporting that cats came in to fight with their cat and 25.6% reporting that cats came in to steal food. For households that had a plain cat door without security features (as opposed to a selective entry cat door that only allows resident cats in and out) the figures were significantly higher; 24.8% of cat owning households reported that neighbourhood cats came into their home to fight with their cats, and 39.4% reported that cats came in to steal food.
Regular experience of this kind of home invasion, and subsequent injuries from fights with non-resident cats, are a significant source of stress for domestic pet cats. The fact that cats commonly take the risk of entering each other’s core territory in order to get access to food indicates that there are serious problems with the way that pet cats are being fed.
Cats that had experienced injuries due to conflict with other cats showed 3.9 times that rate of indoor spray marking compared with cats that had not experienced injuries. They also waited by the cat door before going out, and became agitated and afraid when they saw another cat in the garden.
Only 45% of owners in the survey fed their cats ad libitum, indicating that a lack of availability of food may be a strong motivation for many cats to enter homes seeking sources of food, and thereby ending up in conflict with residents. In support of this, there was a strong, and significant, correlation between cats entering the home for food, and fighting with resident cats in the home.
Cats who had experienced home invasion of any kind showed 24% higher levels of claw marking behaviour in the home compared with cats that had not experienced home invasion. They also waited by the cat door before going out, became agitated and fearful when they saw other cats in the garden, and showed significantly higher rates of hair loss and skin disease.
Domestic gardens also often lack suitable landmarks for scent marking, such as posts for scratching. As a result, the boundaries between the necessarily small territories of domestic cats become blurred. There are few clear indicators of territory ownership to deter a cat from approaching another cat’s home.
Forty-one percent of cat owners that gave their cats outdoor access did not provide a cat door, making these cats are reliant on a person to let them in and out of the house. Cats follow a strict timetable of movement around their territories so that they can avoid contact with other cats, and deposit scent marks at the right times and intervals. By being tied to the owner’s routine, a cat that does not have free outdoor access cat is unable to follow this kind of pattern. This increases the risk of conflict and stress.
However, even though outdoor access is fraught with problems due to a lack of owner understanding of the actual needs of cats, there is evidence that a lack of outdoor access contributes to problem behaviour. Heidenberger (1997)[9] found that cats that were allowed to go outside when they wanted, or at least 2-3 times weekly, were less problematical to their owners.
References
- ↑ Liberg, O., Sandell, M., Pontier, D., Natoli, E. (2000) Density, spatial organization and reproductive tactics in the domestic cat and other felids. Pp. 119-148 In: D. C. Turner and P. Bateson (eds.). The Domestic Cat: the biology of its behavior. 2nd edition, Cambridge University Press, Cambridge, England.
- ↑ Liberg, O. (1980) Spacing patterns in a population of rural free roaming domestic cats. Oikos. 38, 336-349.
- ↑ Liberg, O. (1984) Home range and territoriality in free ranging house cats. Acta Zoologica Fennica. 171, 283-285.
- ↑ 4.0 4.1 4.2 Bradshaw, J.W.S (1992) The Behaviour of the Domestic Cat. CABI, Oxford, UK.
- ↑ Turner, D.C. & Bateson, P. (1986) The domestic cat; the biology of its behaviour. Cambridge University Press, Cambridge.
- ↑ Foley, P., Foley, J. E., Levy, J. K., Paik, T. (2005) Analysis of the impact of trap-neuter-return programs on populations of feral cats. Journal of the American Veterinary Medical Association. 227(11), 1775-1781.
- ↑ Driscoll, C. A., Macdonald, D.W., O'Brien, S.J., (2009). From wild animals to domestic pets, an evolutionary view of domestication. Proceedings of the National Academy of Sciences. 106(1), 9971-9978.
- ↑ Corbett, L.K. (1979) Feeding ecology and social organization of wild cats (Felis silvestris) and domestic cats (Felis catus) in Scotland. PhD thesis, University of Aberdeen, Scotland.
- ↑ Heidenberger, E. (1997) Housing conditions and behavioural problems of indoor cats as assessed by their owners. Applied Animal Behaviour Science. 52, 345-364.
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