Exploring Behavioural Plasticity of Cats

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This entry is adapted from the peer-reviewed paper 10.3390/ani12131717

Understanding behavioural plasticity and other recently evolved traits of domestic cats may lead to management strategies that maximise health and welfare of cats, wildlife, and humans—otherwise domestic cat behaviour may be ‘misunderstood’. Importantly, interdisciplinary research using expertise from biological and social sciences, and engaging human communities, should evaluate these management strategies to ensure they maintain optimal welfare of free-ranging domestic cats while preserving biodiversity and protecting wildcats.

domestic cat behavioural plasticity evolution

1. Density of Unowned Domestic Cats

The domestic cat (Felis s. catus L.) is now present on all continents, with the exception of the poles and certain islands. It lives in a wide variety of environments where the spatial and temporal distribution of resources vary widely, so that the species includes individuals with very different lifestyles (reviewed in ^[1], this Special Issue). This ubiquitous nature of the domestic cat is a sign of high behavioural adaptability ^[2], as will be seen below. It is now generally accepted that behavioural studies of domestic cats, and other opportunistic animal species (opportunistic species—a species that can quickly exploit new resources as they arise, for example, by rapidly colonising a new environment ^[3]) (Norway rats Rattus norvegicus, Berkenhout 1769, ^[4]; house mice Mus musculus L., ^[5]; red foxes Vulpes vulpes L., ^[6]; starlings Sturnus vulgaris L., ^[7]; yellow-legged gulls, Larus michahellis, Naumann 1840, ^[8]), have shown that the most important environmental variable in determining the density and spatial distribution of the animal population is the quantity and distribution of trophic resources ^[9]. But substantial variation in the density of the descendant species, compared to the ancestor, can affect all aspects of life for these animals, from spacing patterns to social organisation to the mating system, from the dynamics of prey–predator interaction to the dynamics of parasite–host interaction. Among domesticated species, the domestic cat represents a highly valuable and unique model to study these phenomena because it can live a strictly solitary life or in socially structured groups, depending on environmental conditions. This allows comparison of population characteristics at the intra-specific level to identify factors that predispose species to evolve sociality, a privilege for students of behaviour that few mammalian species allow.

Studies carried out in contrasting ecological conditions have shown that in the sub-Antarctic island environment, the unowned domestic cats, which can be defined as ‘feral cats’ (feral cats—domestic cats that, after being domesticated, returned to a feral situation and are not dependent on humans for trophic resources.), live at a very low density (less than a cat/km², ^[10]) in individual territories (Figure 1).

Figure 1. Domestic cat photographed at the Ratmanoff penguin farm zone (Grande Terre, Kerguelen Archipelago). The cats eat the carrion of young penguins or rabbits that have dug burrows in the vicinity of the penguin farm zone (credit Dominique Pontier).

They therefore live a solitary lifestyle, very similar to that of the European wildcat (Felis s. silvestris Schreber, 1777) ^[11].

At the other extreme are the urban unowned free-ranging domestic cats (FRCs), living in colonies in the urban environment at a very high density (more than a thousand individuals per km²) and within a social spatial organisation ^[12]^[13]^[14]^[15]^[16]^[17] (Figure 2).

Figure 2. Adult cats living in group in a Sicilian town; they are eating pasta (credit Eugenia Natoli).

The rural cat lives in an intermediate situation, typically within a human dwelling or property, with a relatively low density (about 250 individuals per km², ^[13]) in small groups.

2. Spacing Pattern of Unowned Domestic Cats

Home range and territory should not be taken as synonyms; the former indicates the area of activity for an animal, patrolled but not necessarily defended against trespassing conspecifics. In contrast, the territory is an exclusive area actively defended by the resident cat (or by a group of resident animals), smaller than the home range and enclosed within it ^[15]. There are few studies in which this distinction is made, and, they are in the urban context. In fact, in most cases, the research focused only on the home range, since it is easier to assess the area in which a cat’s activities occur (i.e., home range) than the actively defended part of this area (i.e., territory). As Liberg and colleagues emphasise ^[13], the utilisation of the term ‘territoriality’ requires active defence of the range (p. 133); of course, ‘There is a large asymmetry between the data needed to show range overlap and exclusive range’ ^[13] (p. 133). Intuitively, considering the small size of the spaces colonised by FRCs in the urban environment compared to, for example, the rural environment, it is easier to notice the difference between the home range and the territory of a resident male in the urban cat colonies. In every study that the first author (E.N.) has conducted on cat colonies in an urban environment, she has noticed that cats belonging to different neighbouring social groups tolerated each other, although they threatened each other in the overlapping marginal zones (one can assume, the marginal parts of the home range). But if an unfamiliar cat intruded into the innermost part of the group’s territory, it was attacked by the adult individuals (males and females) of the group (^[15] p. 301), and the intruder aroused the curiosity/fear of the young. These behavioural observations apply to unneutered domestic cats.

Therefore, researchers use the term that the author(s) use for each of the studies researchers cite, which may be either ‘home range’ or ‘territory’.

The domestic cat is a territorial animal, whatever environment it lives in. This means that, when it lives as a solitary animal, it conducts its life within a large area containing resources such as food, shelters, and sexual partners, which the cat defends from the intrusion of conspecifics. Therefore, if sufficiently competitive, an adult male can occupy an area in an exclusive way, with the size of the area determined by the richness of prey species. The more resources available, the smaller the area and the lower the costs of controlling it. The hypothesised original spacing pattern is one in which the home range of an adult male includes two or three smaller home ranges of adult females ^[18].

In general, tomcats have a significantly larger home ranges than females (but see ^[19]), regardless of habitat and type of resources—natural prey and/or food distributed by humans, as also reported for the African wildcat ^[18].

In sub-Antarctic islands, individuals of both sexes are solitary, living in large non-overlapping home ranges ^[20]^[21]^[22], although there are some indications that, in Kerguelen islands, male home ranges can overlap in space, but not in time ^[23].

The spatial distribution pattern of rural unowned domestic cats resembles the pattern of African and European wildcats ^[18]^[24]^[25], consisting of a male owning and monitoring a relatively large home range, which includes the home ranges of two or three females. In this environment, males and females are solitary but females may also form small groups of closely related individuals associated with human dwellings ^[13]^[26]^[27].

In contrast, in the urban environment, when unowned domestic cats live socially, the territory belongs to the group and is defended by all adult male and female members of the group ^[15]. At the centre of the territory is the core area, the area of most intense and regular use, which represents the most important place in the life of a social group of mammals, where mostly high-ranking adult and young cats live.

3. The Social Organization of Unowned Domestic Cats

In environments where unowned free-ranging cats live as solitary animals—depending exclusively on prey for their survival—there is no social organisation and therefore no point in using the term ‘dominant’ male; that is, the concept of dominance hierarchy is only relevant to a social group ^[28]^[29]. The same is true for the rural environment as described in the previous section.

In urban ecosystems where cats depend on food provided by cat caretakers ^[30], cats organise themselves around feeding sites, forming highly structured, stable multi-male/multi-female groups called colonies ^[31]. Male and female home ranges and territories overlap ^[32]^[33]. Cats are fed by people who distribute the food in specific locations where all cats eat at the same time. In a rural environment, each cat (or several cats attached to the same farm) has its own feeding location provided by humans (26), i.e., different to that of the cat(s) living on the neighbouring farm. In contrast, in an urban environment, all cats, even dozens of cats, belonging to the same group share the same feeding location. This provisioning may have a negative impact (e.g., spread of viruses ^[34]), but also highlights the resilience of domestic cats who can adapt and survive in seemingly hostile environments. Provisioned cats live in areas around hospitals, in public gardens, and in historical ruins, as these places become favourable islands with shelters, surrounded by roads with heavy traffic. In a study conducted in a hospital in the city of Lyon (France), the home ranges of adult females were organised around a single feeding site. Adult males’ home ranges included up to three of five permanent feeding sites ^[33], although all cats were able to meet their energy requirements by visiting only one site—supporting the hypothesis that in domestic cats, spacing between males is a response to the distribution of breeding females ^[14].

Moreover, these multi-male/multi-female groups show signs of sociality ^[35]. Adult individuals defend the territory against intruders belonging to different groups ^[15], and friendly behaviour has been observed among adult females and among adult males and females of the group ^[35]. In contrast, tolerance with moderate intra-specific aggressive behaviour, but no affiliative behaviour, was observed among adult males, to allow group living ^[32]. A linear dominance hierarchy, based on the outcome of agonistic interactions (behaviours that manage conflictual relationships between individuals: threat, aggression, submission), develops between males and, independently, between females ^[32]^[36]^[37]^[38]. When the group of cats is well-established, both males and females live ‘in harmony’ if each cat respects its place in the hierarchy; namely, its own rank, which is a measure of dominance ^[28]. To speak of rank has meaning only when referring to a group of individuals that stay together long enough to allow a dominance hierarchy to emerge.

Dominant social status is maintained primarily through a set of ritualised cues, rather than through aggressive interactions or even fights ^[39]. When encountering dominant cats, subordinate cats engage in behaviours such as looking away, lowering their ears slightly, or turning their head away. If the dominant cat walks the same path as the subordinate, the subordinate will deviate from the path to let the dominant cat pass. Dominant cats signal their status through other signals. When approaching a subordinate cat, they will fix their eyes on it, stiffen their forelimbs and hind limbs, raise and rotate their ears so that the opening is lateral, and raise the base of the tail while leaving the rest of the tail low—the tail is comma-shaped ^[40].

To reinforce their dominance status within their group, dominant cats very often circle their home range, making themselves visible to all cats, maintaining their home range size throughout the year ^[33]. However, socially dominant cats do not have priority access to food over adult females or kittens until they are one year old ^[36]. Interestingly, when the dominance hierarchy was measured in different contexts (in the absence of resources and in the presence of food), all adult females of the social group gained in rank ^[36], and the same occurred in cat colonies where all cats were neutered ^[37].

The dominant males mark the territory with urine sprayed throughout the year, more than subordinate individuals ^[37]; the higher the rank, the more frequent the territorial marking behaviour of urine spraying ^[13]. This is a ‘testosterone-dependent’ manifestation; in fact, it decreases dramatically, and in some individual ceases altogether, following castration ^[37].

Within the colony, cats show frequent friendly (affiliative) interactions, including greeting ceremonies (nose–nose contact with tails held high), reciprocal rubbing, allogrooming, and passive contact ^[35]^[37]^[38], behavioural patterns reported also at dyadic level ^[38] (Figure 3).

Figure 3. An adult female (on the right) and an adult male (on the left) belonging to the same social group. They are showing the affiliative behaviour ‘touching nose’ with their tails up ^[35] (credit Eugenia Natoli).

Cats also show olfactory recognition of group members over strangers ^[41], while females cooperate extensively in rearing their offspring ^[15].

Finally, cats recognise colony members and non-members. Most or all colony members show aggression towards unfamiliar cats. Thus, as is the case for most social species, non-members of the group are not allowed to approach and enter the group ^[13]^[15].

In summary, domestic cats can live in groups that, under certain conditions, have true organisation which is one of the characteristics of social groups.

4. The Mating System of Unowned Domestic Cats

In rural cats attached to human dwellings and in unowned domestic cats living in the urban environment, females have between one and two oestrus (ovulation) periods per year, averaging four to five days ^[27]^[32], with the number of oestrus periods depending on environmental conditions and food resources. Thus, in the Kerguelen archipelago, where the favourable climatic season is very short (a few months), females will only have one oestrus period.

As the cat is an induced ovulator, repeated coitus is necessary for ovulation to occur in most females ^[42], and ovulation occurs 24–50 h after copulation. Moreover, cat sperm require 24 h to capacitate ^[43]. Thus, they need to stay within the genital tracts of the female for about 24 h before they can fertilise the eggs. These characteristics of both sexes favour multiple copulations to ensure the fertilisation of all the eggs and, depending on the spatial and social organisation of populations, in some cases favour the mixing of sperm from different males that, in turn, can lead to multiple paternity (i.e., one litter sired by more than one father) ^[44]^[45].

It has been observed that the mating system changes between low-density islands and rural environments, as well as high-density urban unowned domestic cat populations. Males, whether social or solitary, compete for access to receptive females regardless of the social organisation of the females.

In the Kerguelen archipelago, no multiple paternity has been found ^[46], suggesting that the potential for sperm competition is very low there. The lack of behavioural data means it is impossible to determine whether the monopolisation of entire litters by females living in the territory of males is due to a system of polygyny without extra-pair copulations or to monogamy (Figure 4).

Figure 4. Kittens born at the Ratmanoff penguin farm zone (Grande Terre, Kerguelen Archipelago) (credit Dominique Pontier).

In rural areas, behavioural observations ^[26] have shown that a few males are able to control access to receptive females and perform essentially all copulations, defining a polygynous mating system. The outcomes of fights in males are largely influenced by their phenotypical characteristics ^[26]: age, experience, physical condition, and body size are all important variables in determining the behavioural tactics adopted by a cat in each encounter ^[26]^[47]. Generally, heavier males overpower lighter males in agonistic encounters during the oestrus period of females and have a higher mating success. One example is a male with an all-white coat, homozygous for the W (for White) allele—a dominant allele over any other coat-colour allele ^[48]. Near the house where this male lived, there were three other houses and two farms with a total of five males and eleven females, none carrying the W allele. This white male, monitored between 1990 and 1992 ^[49], in 1991 sired 63 (all white) out of 66 kittens delivered by 10 of the 11 females in 18 litters—clearly demonstrating that he had successfully monopolised these 10 females during the breeding season. This male sired no kittens in 1990 and only six kittens in 1992, suggesting that the costs of monopolising females may limit the ability of males to keep competitors out for several consecutive years.

Behavioural observations showed that other males may copulate with females in the absence of the territorial male ^[26], as the three kittens not sired by the white male suggest. In another rural cat population, Barisey-la-Côte (eastern France), 22% of litters with more than one sire, as well as litters of kittens sired by ‘stranger’ males from outside the population, were identified ^[44].

In large urban multi-male/multi-female social groups of domestic cats, males and females mate with several partners ^[16]^[32]^[44]^[47], adopting a promiscuous mating system. Males are not very aggressive during the breeding season, which allows subordinate males to remain in the social group and reproduce ^[32]. It has been shown ^[16] that males’ breeding decisions are flexible and depend on the degree of synchronisation of female oestrus; males must choose between defending each courted receptive female until the end of her oestrus period (which can last several days) or leaving her to find a new receptive female. Oestrus of females in colonies usually occurs synchronously ^[47], but female oestrous may become desynchronised in relation to environmental factors (such as poor weather conditions or prolonged absence of feeders). When many females enter oestrus simultaneously, a single male—even at the top of the hierarchy—can no longer monopolise each female, with males fighting to monopolise females therefore losing breeding opportunities. Rather than defending the females they have just mated with, males adopt a search strategy to gain access to as many females as possible ^[16]^[33], regardless of their social rank or body weight ^[50]—in contrast to cats in rural environments ^[44]. They spend less time with each female, which considerably decreases the probability of being the father of the kittens, leading to a high rate of multiple paternity. In an unowned domestic cat population living in the neighbourhood of a hospital in the city of Lyon, it has been shown ^[44] that the proportion of litters sired by at least two males was as high as 80%; in some litters, the numbers of the kittens coincided with the number of fathers.

Conversely, when the oestrus of females is asynchronous, the number of males per receptive female is greater, which increases the competition between them to mate with females, and only the most competitive males (at the top of the social hierarchy) have the highest reproductive success—as occurs in low density rural populations. These same males were unable to monopolise females when their oestrous cycles were synchronous ^[16]. The variance in male reproductive success was four times greater in years when females bred asynchronously, and dominant males sired most kittens produced. Although the percentage of multiple paternity was the same between the two situations, the proportion of litters where the top males sired at least one kitten was much higher when the females were asynchronous ^[16]^[45]. However, no males, even of the highest rank, were able to monopolise an entire litter ^[16]^[45].

5. Infanticide in Unowned Domestic Cats

Rare cases of infanticide have been observed in rural areas only, a strategy that may have developed to increase reproductive success of wandering males ^[51]. The method of killing kittens is generally the same as that described for lions ^[52]: the male bites the kittens on the neck, holding them while shaking them vigorously. All females observed by the authors (D.P. & E.N. ^[51]) reacted aggressively but were unable to prevent infanticide. One female had two litters attacked where the same male killed some of her kittens. On the second occasion, the female defended the litter in cooperation with the resident male (three years old). The couple managed to deter the attacking male after he had killed the first kitten, and both the resident male and the female showed injuries. No cases of infanticide were recorded in high-density urban colonies where the opportunities appear to be greater, but also where collective defence of litters by females could plausibly be a sufficient deterrent against male attacks. However, this divergence may also be explained by paternity uncertainty in large colonies, where most females mate with several males ^[53].

6. Domestic Cat Behavioural Plasticity as a Gatekeeper of Evolution

The relationship between behavioural plasticity and natural selection has attracted the attention of scientists for a long time ^[54]. On the one hand, behavioural plasticity allows individuals to respond to ecological pressures by modifying their interaction with new environmental conditions (e.g., exploiting a new food resource) without modifying the genetic diversity of the population. On the other hand, it slows down or even prevents the action of natural selection ^[55]^[56]^[57].

Zuk and collaborators ^[58]^[59] suggested that it can be the other way round—that behavioural plasticity might have the greatest influence in supporting new traits.

The intensity of selection for behavioural plasticity depends on the balance between a positive effect it may confer, and a negative one, such as the cost of plasticity on fitness. This balance can lead to different levels of genetic diversity within an adapted population.

The case of the unowned free-ranging domestic cat is particularly interesting because it is possible to compare the characteristics of different populations at an intra-specific level, and allows testing of hypotheses related to the effects of behavioural plasticity and natural selection on evolution of novel traits. There is a large array of traits that can be influenced by behavioural changes: morphological, physiological, and immunological. In fact, all levels of organisation within an organism are integrated by behavioural traits ^[60]^[61]. An individual who exhibits behaviour that differs from the original one must change other traits to cope with new abiotic, biotic, or social environments ^[58]. The latter point is of particular interest in the case of unowned domestic cats in the urban environment, since they face a social environment unknown to ancestral populations.

Therefore, the question researchers are asking is whether there are indications that new traits (behavioural, morphological, physiological, immunological) are evolving in the domestic cat and what role behavioural plasticity and/or natural selection (allele frequency changes) may have had or have in this process.

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