Development of Behavior: Evolutionary Background - Normal Cat Behavior
Karen L. Overall, MA, VMD, PhD, DACVB, ABS Certified Applied Animal Behaviorist
The story of canine domestication is the story of work and work-related tasks. The story of feline domestication is the story of rodent and vector borne diseases and their prevention. These 2 divergent paths to domestic life-styles have been shaped by, and in turn have continued to shape factors like reproductive schedules, fecundity, age at first reproduction, age at sexual and social maturity, composition of family or group units, and social interactions within these units.
Outline of feline social systems:
1. female (matriarchal) family packs in large cats
2. solo or paired animals in smaller cats
3. lone bachelor males or male groups
4. one primary male in breeding group
a. frequent copulations in a short period
b. short tenure
c. potential for demographically structured infanticide
5. communal breeding / care
a. females related
b. induced ovulator / polyestrous
6. gestation to 120 d. in jaguars
7. females as primary protector and hunter
8. more obligatory carnivore (but consider margays)
9. marking behavior: urination and defecation
a. open areas: spraying
10. shifts in this in solitary (mountain lions) vs. lions
11. social development - concept of risk assessment and reduction of cost of error
a. born blind
b. eyes open 10-14 d.
c. 3 weeks of age: queens begin teaching kittens predatory behavior
d. 5 weeks of age: independent kitten predatory behavior
e. early weaning hastens above periods
f. 4- 12/14 weeks of age: social play
1. 2-5 weeks of age: early period
2. 5-7 weeks of age: mid period
3. 7-10 weeks of age: late period
4. 14 weeks of age: post period
g. response to visual / olfactory threats: 6-8 weeks
h. object play: 7-8 weeks of age (depends on eye-paw coordination)
i. social fighting: 14 weeks of age
j. "temperament" mediated by:
1. father (determinant of friendliness)
2. coat color? (aggressive propensities)
a. 50 % maternal ration decrease - less mothering
b. decreased brain development for kittens
4. early exposure and total absence thereof
5. genetically shy / unfriendly litters
k. small periods of very early social exposure
1. earlier periods than dogs, possible
l. innate development of elimination behaviors
n. play (rough) as predatory behavior rather than social integrator
Feline olfactory / pheromonal cues:
1. vomeronasal organ - Flehmen
2. chin glands
3. role of kneading
5. tail quiver and rubbing
6. consider role of evolutionary social dynamics
Domestic and wild cats use both scrapes and elimination products to mark their territory. Cats are motivated to respond to olfactory cues in their environment, and are well equipped with a variety of devices, including glandular secretions, by which they can lay their scent in the area. When cats rub against someone and knead, or mark with their head, chin or cheek whiskers, they are leaving behind sebaceous gland secretions that can be detected by another cat. Cats have a superb senses of smell. It has been estimated that the size of the olfactory epithelium in cats can be up to 20 cm2; this contrasts dramatically with the 2-4 cm2 of olfactory epithelium estimated for humans. It has been postulated that cats' olfactory abilities run on a parallel order with those of dogs, who are able to detect compounds of thresholds a thousand times lower than those detectable for humans. Given this, it should not surprise us that feline social systems are mediated in part by olfactory mechanisms, and that these in turn may be a factor in elimination disorders. Accordingly, it is important to characterize patterns of spraying and non-spraying marking that are involved in feline aggression (Table 1).
Data collected on free-ranging domestic cats has shed some light on the importance of the social system in both feline elimination disorders and the potential for feline elimination disorders involved with aggression. Male cats range further than do female cats; the home-range size of males extends from 0.4 ha to 990 ha, while the home-range size for females extends from 0.02 ha to 170 ha). Average home range varies from 0.1 to 0.45 ha. The average apartment is anywhere from one to three orders of magnitude smaller than that range. It is no wonder that when multiple cats converge in small territories there may be some social jockeying.
Patterns of spraying that are involved in feline aggression and can be part of "normal" behavior include: 1) active aggression including overt fighting and visual threats (e.g., stares, rubs, posturing), 2) passive aggression including overt signaling and posturing and covert olfactory challenges (e.g., rubbing in absence of observer), 3) and status-related spraying. Active aggression usually involves a confident cat spraying. Confident cats can spray because of a) sexual advertisement in competition, b) advertisement not associated with sex, c) post-aggression or as a "victory display". Advertisement not associated with sex may actually avoid future physical aggression. Post-aggression victory signals may decrease the probability of future aggressive events. "Victory displays" have been postulated to function in any situation involving allocation of resources, including time. If diffusion of aggressive situations saves time and injury, this should be a favored strategy since both time and the calories necessary to heal injuries can be better spent.
It has been under-appreciated that passively aggressive cats can spray. These cats may be acting normally for their temperament, or may be pathologically anxious because of the complex social environment in which they find themselves. Cats who are exhibiting a passive, rather than an active physical aggression, are usually less confident than other cats. They can use spraying as a) a passive threat, b) a response to a preceding physical threat from an animal with whom they do not feel they can adequately contest, c) a response to an olfactory cue from another animal who may be higher or lower on the hierarchy, but who still is not a viable target for a physical contest, or d) as an anxious or fearful response to uncertain circumstances. The latter has been under-appreciated as a form of passive aggression. Animals that are less than confident may, when a social system changes by the addition or absence of another animal, learn to deform that social system further by spraying. This is a form of advertisement that would get them information that they otherwise do not have available. Depending on the response to their spraying, they are able to ascertain information that they cannot get in a direct manner.
Cats who are exhibiting spraying related to social status have also been under-appreciated. This type of status-related spraying could also involve marking with urine or feces, but it could be a purely visual rather than a visual and olfactory display. The postures that go along with spraying are stereotypic, do not have to be accompanied by urine spraying, and would be recognized by another cat as associated with assertion of status and/or the claim to territory. In many cases, cats of high status will be defining a "mine" versus a "not-mine" situation. The objects they may be claiming could be locations, inanimate objects, or people. In this case, the spraying has less to do with the rest of the underlying social interactions; it acts to preempt status for them. In status-related spraying, the presence or absence of other cats in the social system is relatively less important than the overall statement of assertion of position from the individual doing the spraying.
The mechanism for avoiding conflict in households is usually a hierarchical system based on deference . Many of the individuals who have examined feline social systems in wild cats, such as mountain lions, have focused on females as family units. In such situations, usually the females in the group are related and the males are unrelated to the females. One male will control access to breeding of many females and very few younger males ever have any reproductive success. In evolutionary systems involving high variance in reproductive success, contest for position is common. Selection over evolutionary time may have reinforced social systems that facilitated such competition. Such social systems would be maintained in the absence of sexual overtones. This is exactly what happens in domestic cat situations involving neutered animals. Even within the free-ranging, wild felid social hierarchies, females have an independent hierarchy that is largely based on age, but is also influenced by other social characteristics and physical characteristics such as strength. Bernstein and Strack have noted that most individuals who possess more than one house cat are able to rank their cats according to some deference-based or conflict-based hierarchy and to articulate reasons pertaining to specific behaviors for why they think one cat is more highly ranked than the other. However, this deference-based structure serves to avoid conflict. In such households cats have been shown to time- and space-share. If given adequate opportunity (e.g., 3-D space, hiding holes) to share space and time, overt aggression is rare, as is any covert aggression involving spraying.
If we are to fully address elimination disorders and their social ramifications, it is important that we pay attention to these newer findings.
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