of a distinct estrus call vocalization. She also displays frequent rubbing of cheeks, head, flank, and back on various substrates as well as rolling and spraying urine to advertise her estrus status. Another defining queen behavior displayed during courtship is the lordosis posture—crouching with perineal area lifted vertically and back legs treading. Estrus behaviors occur regardless of a male cat’s presence. In the proestrus phase, the queen displays courtship behaviors but acts aggressively if the male (tom) cat tries to mount. She only becomes immobile and allows mounting and biting of the neck as she enters the estrus phase and estrogen levels rise. Felids are induced ovulators, meaning physical stimulation of the vagina is needed to trigger release of an egg from the ovary (Hart and Hart 2014b).
Tomcats are drawn to the estrus female via olfactory cues. His courtship behavior is a sequence of genital sniffing, displaying the gaping or flehmen response, and then using teeth to grasp the nape of the queen’s neck before mounting. The tom also treads his back legs during mounting and intercourse. The female reacts strongly at the time of intromission, likely a result of hyperstimulation or discomfort from the tomcat penile spines. She produces a loud shriek accompanied by dilated pupils and darting away from the male (Houpt 2018).
Natural mating behavior consists of frequent copulations (approximately every 10 minutes for several hours) of less than 1 minute. In a free‐ranging cat environment, this mating strategy provides an opportunity for multiple males to mate with a queen during estrus. Numerous toms surround the queen and try to displace each other during or between copulation. Thus, multiple paternity within litters is common. Estrus ends about two days post‐ovulation, but an unbred cat will cycle approximately every three weeks throughout the breeding season, which is January through October in the Northern Hemisphere (Houpt 2018).
2.7 Sociality
2.7.1 Intraspecific
Domestication involves morphological changes as well as alterations to physiology, emotional systems, and social behavior (Coppinger and Smith 1983). Considering the domestic cat’s ancestral species as well as all other living members of the Felis genus are solitary and territorial individuals, the degree of intra‐ and interspecific social behavior displayed by the cat is extensive. Nevertheless, domestic cats tend to be solitary hunters in settings with sparse food sources. To allow cats to take full advantage of concentrated food niches provided by humans, feline social signaling and the ability to deescalate territorial tensions had to evolve (Bradshaw 2016). The social framework of feline colonies is matrilineal, meaning related females often form stable groups and are intolerant of unrelated females. In addition, females within a group often cooperatively raise kittens through group nursing and sharing of captured prey (Macdonald et al. 2000). Moreover, the basis of all affiliative cat social behavior seems to be maternal care. Many friendly behaviors observed between cats, such as allogrooming and allorubbing, as well as the purr and kneading behavior, are first seen between kittens and their dam (Bradshaw et al. 2012). Physical contact provided through mutual grooming and rubbing probably maintains group cohesion through both olfactory and tactile cues (Bradshaw 2016). The vertically raised tail is one of the most notable body postures for signaling friendly intentions from a distance.
Proximity, particularly with non‐group members, inevitably leads to some degree of conflict, prompting cats to communicate the intent to avoid conflict or willingness to fight by means of ritualized interactions (Bradshaw 2016). Threat behaviors convey a cat’s desire to maintain personal space or access to a resource. A confident cat conveying a threat may stare at and/or move without hesitation toward the intended receiver. The body posture is usually upright and ear pinnae turned to the side. The tail may be low but curved up at the level of the hocks with tail lashing occurring with higher‐arousal encounters. Firm head bunting and urine spraying can be threatening behaviors that incorporate tactile and/or olfactory signals without overt aggression. A confident cat rarely vocalizes during a threat. The cat receiving these signals may respond by moving away, typically with a lowered tail, body posture, and ear pinnae. Body posture becomes increasingly crouched, with the tail tucked under the body, ears flattened, and pupils dilated with elevated distress or fear. A roll onto the back is often observed in extreme situations as well. Although a roll may be seen during some friendly interactions, in this situation the cat is maneuvering to use the claws defensively if necessary (Leyhausen 1979). Vocalizations like hissing and yowling increase with the level of distress and threat perception as well. Although scratching and biting are possible, avoidance and appeasement behaviors are much more common, thereby reducing the risk of injury that may incur during a physical altercation (Dantas et al. 2011). The aggressor is not always confident, and a mix of fear and threat behaviors is often seen in both parties during an agonistic encounter. Aggression may become more prevalent in heavily populated and physically constrained colonies (Knowles et al. 2004).
2.7.2 Interspecific
Millions of cats interact with humans in various housed or free‐roaming environments, and the field of feline social cognition is uncovering what most cat owners already know—cats are capable of forming strong bonds with humans (Turner 2017). Geering (1986) found that food may be an initial attractant for a cat, but affiliative behaviors such as stroking are needed to maintain contact with a person. More recent studies have suggested that once a cat has bonded with a human, the cat shows a preference for contact from this person compared to others and feels more comfortable in exploring surroundings with familiar people present (Edwards et al. 2007). Cats tend to look for cues from a familiar person in the presence of a frightening stimulus (Merola et al. 2015) but, unlike dogs, may not necessarily seek human guidance when solving spatial tasks (Miklósi et al. 2005). Cats can both recognize the voice of familiar humans (Saito and Shinozuka 2013) and alter vocalizations directed toward humans. Cats emit the meow (Yeon et al. 2011) and modified purrs (McComb et al. 2009) almost exclusively toward humans when seeking resources such as food and affection, although similar sounds may be heard in the queen‐kitten context. Most affiliative behaviors displayed by cats toward humans—kneading with the forepaws, purring, allorubbing, and allogrooming—mirror conspecific behavior. Human scratching and stroking are likely perceived as reciprocation and maintenance of the bond. Human personality and perception of the cat as a social partner have been shown to influence the characteristic of both the specific human‐cat interaction and a cat’s general behaviors in the home (Wedl et al. 2011; Kotrschal et al. 2014). Arguably the most important factor in the degree of sociability of any cat is early life experiences with humans or any other social partner.
2.7.3 Aggression
Aggressive behavior is often a normal reaction to a stressful social situation, such as the presence of an unfamiliar cat or unwanted attention from a human. Cats tend to avoid conflict as much as possible, but physical constraints or limited resources may force a cat into proximity of the concerning human or cat, triggering aggression. Learning can shape emotions and behavioral responses. Negative emotional associations between individuals result from aversive experiences, and aggressive behavior may be reinforced as the most effective tactic to end unwanted interactions.
2.7.3.1 Play Aggression
Play often involves predatory‐like stalking, chasing, pouncing, scratching, and biting. This is considered a normal behavior in kittens and some adults, but aggressive play may be abnormal or warrant intervention if the biting and scratching are intense or the target of play attacks is vulnerable (e.g., child or senior person, geriatric cat). Kittens and cats who were hand raised, separated from littermates at a young age, or singleton kittens are more likely to display more frequent and intense play aggression (Mendl 1988).
Play is a normal aspect of development, and suppression of play may lead to long‐term problems. Aversive punishment is likely to create lasting associations of fear and frustration. The most appropriate methods for reducing play aggression are providing ample opportunities for play with appropriate toys, eliminating use of human hands or feet for play, and managing the environment to reduce exposure to inappropriate targets. For example, a family may keep a basket of toys handy and toss one of the objects in the opposite direction for a cat that tends to stalk and bite the ankles