Wednesday, June 1, 2011

Domestic Cats (Featuring the cats at CAPS!)

Social Life

The Signalling repertoire of the cat and its wild relatives

The scent signals of domestic cats are products of their acute sense of smell, which could have developed primarily to detect food, and from their origin as territorial animals which needed to communicate with neighbors that they may rarely encounter face to face. Communication is said to happen when one animal responds to the signals sent by another.  In many cases there is no reason to believe that both animals agree about the message being transferred; signallers often attempt to manipulate the behavior of recipients to their own advantage, while recipients attempt to mind-read these descriptions. The influence of domestication on signalling adds another layer to the explanation of why signals take the form they do. For the cat, the ancestral species Felis silvestris libyca is believed to be exclusively territorial, and so its signalling vocabulary must have changed as it evolved to live at high densities and become sociable. When individual animals live close togehter, and benefit by cooperating, they must have the ability to resolve conflicts without resorting to physical violence, especially when both parties are as well armed as a cat.

Communication between domestic cats

Olfactory communication

The ancestral species of the domestic cat was likely exclusively territorial, as are most of the smaller species in the Felidae. Since widely spaced animals rarely encounter one another face to face, they tend to communicate by scent marks, which allow a delay of several hours or days between the deposition of the signal and its reception. For well-armed carnivores, there is also the advantage that potentially dangerous encounters with rivals can be avoided by the use of olfactory signals, both those deposited on the layers of medium and those carried directly from the body surface by air current, The potential disadvantage of relying on scent signals is lack of control, both in direction of the message, which depends on the wind, and of who receives it, since a scent-mark cannot be turned off at will, both lead to potential revealing of the information that the scent contains. Despite these problems, members of the Carnivora rely extensively on scent for communication.

Many domestic cats live at a densities several times higher than their wild counterparts, and it is thus possible that their scent communication has been changed during the course of domestication. Cats who live in groups can potentially not only exchange information through scents,  but also exchange the scents themselves to product colony or group specific odors. Comparisons with other species suggest that the domestic cat should have a complex and adaptable repertoire of scent signals, so it is no surprise that  several sources of scent have been found, though their functions in comminication are unknown.


Cats can adopt 2 distinctly different postures for urination, indicating that at least one has some use in signalling. Kittens, juveniles and adult females usually squat to urinate, then usually cover the urine with soil or litter. Although this can be interpreted as an attempt to hide urine and the information that its odor contains. Such deposits are sniffed by other cats if encountered. The duration of sniffing tends to increase with the unfamiliarity of the depositor, suggesting that the sniffer is responding to and gathering information from the odor. This may only be a common occurrence where cats are living in high densities; trying to hide it may be effective in widely spaced territories.

Deliberate scent marking with urine is done by spraying, in which the cat backs up to a vertical surface, and urinates backwards, usually while quivering its tail. While mature males are the most frequent sprayers, adult females do spray. In closed or high density colonies there may be some suppression of spraying in females and younger males, resulting in most spray-marks being produced by a small number of 'dominant' males. Spraying by tom cats is increased by the closeness of oestrus females, resulting in an annual peak in the UK in February/March.

The odor of sprayed urine is pungent, so it may carry other secretions, possibly from the perputial or anal glands. The anal gland secretion, which is voided by very frightened cats, has a strong odor, but this is not, to the human nose, similar to that of sprayed urine. The odor of sprayed urine increases after deposition and is likely largely due to the microbial and oxidative degradation of the two unusual amino acids. The male can excrete large amounts of felinine, whereas females produce less, thus less pungent sprayed urine. This excretion may have a significant effect on the sulphur containing amino acid requirements of the male; it is possible that the amount of felinine in urine, and hence the strength of the odor, is a reflection on the success of the male in obtaining high quality food, and is thus an 'honest' signal advertising his fitness as a mate to females and as a competitor to other males.

The territorial function of urine spraying, if any, is unclear. Spray marks are rarely seen to act as a deterrent on their own, but this is the case for most territorial scent marks. Even those which mark edges of territories, which those of tom cats do not. Maybe since the odor of scent marks changes with age, they could be assisting cats to space themselves out while hunting, so they can avoid areas which have been disturbed recently. This is unlikely to be a reliable strategy, because cats that did not spray urine could put themselves at an advantage because other cats would waste time and effort hunting in places where prey was still wary due to recent presence of a predator.

All cats, but especially adult males, investigate spray marks intensely,. especially if they are produced by oestrus females, which suggests that they do contain relevant information. Initial inspection is normally by sniffing, often followed by flehmen, in which the upper lip is raised and the mouth held partially open; this can persist for 30 seconds or more. During flehmen the may make more physical contact with the source of the odor, and move the tongue to and fro behind the incisors. Both airborne and fluid borne molecules of the scent are carried to an accessory olfactory organ of unknown function in the cat, Since flehmen is only done in response to odors from other cats, it is assumed that it is used to gather and possibly store social information.

Many species within the Carnivora use faeces, often with glandular secretions added, to convey information but the evidence that domestic cats do this is only circumstantial. Near to the core of the home range, feces are usually buried but they may be left exposed at another site. Cats usually sniff the places where they have just buried feces but tend not to do so after leaving them exposed. This suggests that one of the functions of burying feces is to minimise the chance that the olfactory information they contain will be detected by another cat, although hygiene could be a more likely explanation.

Though it has a role in conditioning the front claws, scratching must result in the deposition of scent from the glands on the paws (interdigital glands). The same scratching site is often used repeatedly, resulting in a clear visual marker which draws attention to the olfactory information. The scratching sites are distributed along regularly used routes, rather than at the edge of the territory./homerange.

Skin glands
Domestic cats have several skin glands in addition to the interdigital glands including the submandibular gland beneath the chin, the perioral glands at the corners of the mouth, temporal glands on each side of the forehead, a gland at the base of the tail which can over secrete in intact males, giving rise to the condition stud-tail, and caudal glands, which are diffusely distributed along the tail. The pinnae external ears also produce a waxy secretion. It is unclear if each of these glands produce a unique secretion, each with a well-defined function, or whether there  is overlap. The secretions of the glands of the head are rubbed onto prominent objects by a behavior pattern known as bunting. The exact form of this appears to depend upon the height of the object, so that high objects are usually marked with forehead and ears, objects at head height with a wipe of the head from the corner of the mouth to the ear, and lower objects with the underside of the chin and then the side of the throat. This variety suggests that similar odors are deposited from all parts of the head, either because there is redundance between the glandular secretions, or because they become thoroughly mixed on the coat through grooming.

Intact males tent to rubmark more often than do anoestrus females or juveniles, and sometimes spray urine on top of their own rub marks or vice versa. Other rubmarks, although performed on visually prominent objects, such as projecting twigs or corners of artificial structures, are not associated with another visual or obvious olfactory cue and are thus not obvious to the human. Carts appear to be able to locate them easily, suggesting that they are pungent to the cat nose, and often overmark them with their own cephalic secretions. The rubmarks of intact females contain information about the oestrus cycles, as indicated by the degree of interest shown by males, but aside from this there is little information on the function of this behavior. Some cats also rubmark repeatedly in the vicinity of humans, but this could be a transferred verison of cat human rubbing.

Cat cat rubbing is a visual and tactile display which must also result in the exhchange of scents between the pelages of the cat, though it is unclear whether this is relevant, ie. in the establishment of group odors shared by cats who are friendly towards one another. When cats sniff each other, they tend to concentrate on the head region, rather than the flanks and tail, where shared odors would accumulate, showing that even if group odors do exits, individual odors contain more valuable information.

Auditory Communication

Cat's vocalisations usually occur during one of four types of interaction: agonistic, sexual, mother-young, and cat-human. Most of the aggressvie and defensive sounds are strained-intensity calls, since under these circumstances the cat is likely to be tensing her whole body in preparation for a fight. Tension in the throat is thought to be the reason why cats drool during fights, or have to break off from vocalising to swallow repeatedly. The low pitch of the growl and the long duration of the yowl are thought to be designed to convey the size and strength of the cat, and the abruptness and volume of the pain shriek may be designed to shock or startle the attacker into loosening its grip. The female and male sexual calls are of high intensity, advertising fitness to potential partners and rivals of the same sex.

The calls produced by kittens less than 3 weeks old are defensive spit, purring, and a distress call which has aural characteristics similar to the adult miaw. The latter is vocalised when the kitten becomes isolated, cold, or trapped. The call elicited by cold is noticeably higher pitched than the other two, though this difference disappears as the kitten becomes capable of thermoregulation at about four weeks. Restraint induces a call which is similar in pitch to that caused by isolation, but is noticeably longer, and the isolation call is generally the loudest. It is likely that mother cat can distinguish between the calls, and respond accordingly.

Purring is a ubiquitous vocalisation among cats, but its function is not understood, and until recently, its means of production was not clear. It is produced during both inhalation and exhalation, with a brief pause at the transition between the phases, thus sounds as if it a continuous sound. It is generated by a sudden build up and release of pressure as the glottis is closed and then opens, resulting in a sudden separation of the vocal folds, which generate the sound. The laryngeal muscles which move the glottis are driven by a free running neural oscillator, generating a cycle of contraction and released every 30-40 milliseconds.

Although purring is though to indicate pleasure, it is produced in a wide veriety of situations, most of which involve contact between the cat and a person or another cat. Kittens can purr almost from birth, and do so mainly when they are suckling, which can encourage the mother to continue to nurse them. Adults may purr when in contact with a familiar partner, and during tactile stimulation with objects, when rolling or rubbing. All of these situations can be seen as pleasurable to the cat, but there is one serious exception: vets often experience cats who purr continuously when they are chronically ill or appear to be in severe pain. Purring may function as a manipulative contact and care soliciting  signal, possibly derived from its function in the newborn.

Apart from purring, the vocalisation that is commonest in cat human interactions is the miow. This is very rarely heard during cat cat interactions and may thus be a learned response, based on its effectiveness in getting human attention. It is very easy to train in food deprived cats. There are variations in frequency, duration and form, both within and between individuals, which makes it seem that the meow does not have an intraspecies specific meaning. It is likely that each cat learns by simple association that meowing results in feeding, access to desired locations and other resources provided by humans, and some cats learn to produce different meows for different purposes.

Visual communication
Wild type striped tabby domestic cats are cryptically marked, and have no obvious structure that has been specially adapted for signalling. Despite its relatively immobile flat face compared with the wolf, the cat has a varied vocabulary of visual signals, mainly used in regulating aggressive behavior. There is nothing to suggest that the changes to the pelage introduced post domestication, e.g. orange, white spotting, long hair, have had any noticeable effect on ability to signal, in contrast to the profound loss of visual signalling structures in some breeds of dog.

Many of the postures adopted in agonistic encounters can be interpreted as attempts by the cat to change its apparent size, thereby influencing the outcome of the interaction. An aggressive cat will piloerect and stand at full height, whereas a cat that wants to withdraw from a contest will crouch on the ground, flatten its ears and withdraw its head into its shoulders, indicating that it is not ready to launch a biting attack. The defensive-aggressive posture is presented when the aggressor is about to complete its attack and also to potential predators such as dogs. This is normally adopted side on to the opponent, to maximize its visual impact. Though more extreme, it is similar in form to the side step posture used by kittens in play, since this posture tends to disrupt bouts of social play, it is likely that one developed from the other.

Presumably all of these postures are interpreted by that cat's opponent, and used in deciding how to proceed in the encounter, but there is little evidence as to how each posture influences its outcome. Competitive encounters between animals of the same species tend to involve signals which are both obvious, and aimed at manipulating the behavior of the recipient, which should try to combat this by mind reading. The friendly displays of cats are easy to read, but the extent to which each posture is a form of bluffing, and how effective each is at deceiving its recipient, remain to be studied.

In the preliminary stages of friendly encounters, cats tend to avoid looking at each other.  Cats monitor the position of the other, but tend to look away before being looked at. In these conditions, mutual gaze may be being interpreted as a threat signal. In encounters with no friendly content, the amount of mutual gaze was not different from that predicted from the amount of time that each looked at the other, and so may not be being used a signal.

Rolling is a factor of female sexual proestrus behavior, where it is usually accompanied by purring, scratching and rhythmic opening and closing of the claws, and is interspersed with bouts of object rubbing. Male to male rolling appears to be a form of submissive or appeasement behavior since it is never directed by mature males towards immature males and is often followed by the mature male ignoring or tolerating the immature male's presence.

The cat's highly mobile tail, with its independently moveable tip, appears suitable for use as a signalling organ as well as helping a cat balance. It is tucked away between the hind legs in the submissive/defensive posture but this is unlikely to convey much information that is not already provided by the posture. Lashing of the tail from side to side is a factor of aggressive behavior, but its value as a signal is unknown.
The vertically held tail, tail up, TU, is associated with friendly behavior but its function as a signal has only recently been guessed. The vertical tail signal an intention to interact in a friendly way, it is needed because of the potentially deadly consequences of being approached by a cat with unknown intentions.

Tactile communication

Though simple physical contact, as when two cats rest together, may have social significance, the two most obvious forms of tactile communication are cat cat rubbing their heads, flanks or tails on one another (allorubbing), and one cat licking another (allogrooming).

Rubbing tends to take place between cats of unqeual size or status. It is unknown the social meaning of rubbing, whether the transfer of scent that must be inevitable has any significance.

While grooming of one member of a social group by another is significant in many species, it is only recently that it has been studied in the domestic cat. Allogrooming  is a form of redirected aggression or dominance behavior. Sometimes there is no effect of kinship on the choice of partners, which suggests that it is not to maintain bonds between kin. It is possible that allogrooming has other roles in free ranging breeding colonies.

There are multiple main groups of behavior suggested for the cat:
contact, including allogrooming, rubbing; aggressive, defensive, play, sexual, maternal behavior (associated with both contact and rubbing).

Cats' reactions to familiar and unfamiliar people are different due to paternity (genetics) and early socialisation. Growling at a person is inhibited by socialisation but unaffected by paternity, hissing showed stronger paternal effects, frequency of TU is highest in both friendly-father and socialized cats, purring is not affected by paternity, only enhanced by socialisation in the presence of a familair person.

Communication in wild cats: the effect of domestication on signalling behavior

Given the small number of generations since domestication, we can assume that the domestic cat's vocabulary of signals is mostly unchanged from its ancestor, the African wildcat F.s. libyca. However, domestication has noticeable increased the requirement for social communication, both intra and interspecific. It should thus be possible to study the effects of domestication on communication behavior through a comparison of signalling in the domestic cat with that of wild cats.

Three major lineages are believed to exist : the ocelot lineage, which includes the small South American cats, the domestic cat lineage, which includes the small Mediterranean cats, and the pantherine lineage, made up of large and small cats from several continents.

Spatial organization in wild cats

Both the function of a signal and the modality are very dependent on the distance between the emitter and the receiver. Communication is thus intertwined with spatial organization. For any predator feeding on sparesly distributed small prey, non overlapping hunting areas are likely. Field studies have shown this is true for most wild cats, including F. silvestris. There are three outstanding exceptions: The lion Panthero leo, the cheetah Acinonyx jubats, and the domestic cat, all of which have been found living communally. The domestic cat is, however, by no means an exclusive group living species, and has been often shown to be solitary when food is at low density and sparsely distributed. Group living is most often caused by an aritificial hoarding of food associated with human settlements. The change in niche caused by domestication may have caused a decrease in the adaptive value of solitary life, and a corresponding change in intraspecific communication.

Comunication in the wild Felidae; differences between lineages

Even in solitary species or individuals, signalling is needed for mating, parent-young interactions and maintenace of territorial boundaries. The wide range of signals shown by these largely solitary animals is shown by most species.  They have been shown to exhibit a night repertoire of signals despite being mainly solitary. However, the nocturnal and solitary behavior of these species slows the study of communication, thus much of the data, especially on small cats, has been collected on captive cats.

Olfactory communication

Olfactory signals are long lasting and would thus be expected to play an important part in communication between both social and solitary members of the Felidae.


Urine is emitted in spraying  more often in males than females. Sprayed urine may contain anal gland secretions, whereas squat urination is uinlikely to contain anything extra. Squat urinations differ in that the urine is usually raked into the soil with the hind feet, known as scuffing/scraping or raking. It has been suggested that this action may mix urine into the soil and help transfer the urine scent, and possibly also the scent from the glands on the feet to the environment, however, the communicative function of this behavior is unknown.


Wild cats had additionally been shown to scrape their hind feet without urination or defecation, The absence of urine, feces or anal gland secretions implies that scrapes are acting as visual signals as well as olfactory ones, though scraping may help to pass secretions from the glands in the feet on to the medium. Mountain lions Puma concolor mark home ranges, visually and/or chemically.

Scraping occurs in most species of small cats within the ocelot and panthera lineages, but in only one species within the domestic cat lineage, Pallas's cat. This species likely diverged from the rest of the domestic cat lineage at an early stage, in which case this behavior may suggest an evolutionary loss/change among an ancestral member of the domestic cat lineage. This behavior was based in the genus Felis, used only by cats in the same lineage as the domestic cat.

Feces deposition
The method of feces deposition varies according to species. However, it is hard to see if there is an evolutionry pattern to these differences or whether they are dependent on local conditions.  Canadian lynx Lynx canadensis and European wildcat  F.s. silvestris use two methods, depending on where the defecation took place; feces were localised and covered within territories, but left uncovered in prominent positions at points between territories which were used as mating meeting sites in the lynx. This suggests that the method of defecation may depend on local conditions rather than on phylogeny.

Skin Glands
As for the domestic cat, tree scratching functions to remove loose claw sheaths. but it is also used as part of the scent marking in most cats, often occurring in the same areas as other methods of scent marking. It may also leave a visual signal. This behavior occurs in a diverse range of felids (Pallas's cat, sand cat, fishing cat, jungle cat, domestic cat, serval, caracal, African golden cat, ocelot, Canadian lynx, tiger, lion, jaguar, leopard, cheetah) and appears to have altered little in its character or function during the course of felid evolution.

Object rubbing has been suggested to have three ways of acting: it acts as a method of scent marking by depositing gland secretions such as saliva on objects, object rubbing picks up the scent, as many species of small felids rub on objects previously sprayed with urine. Scents were seen being both picked up, e.g. urine and deposited, e.g. saliva, by a variety of species. Observations from many species suggest that object rubbing acts as a visual signal during reproductive and oestrous behaviors and in many species of small cats as it does in the domestic cat. These observations suggest that all three lineages of undomesticated cats used object rubbing similarly, as a signal of both visual and olfactory nature.

The function of scent marking was investigated in tigers Pantehra tigris. It plays a role in establishing and maintaining territories. Scent marking was concentrated at potential contact zones where major routes of travel approached territorial borders, where scent marks give invaders some information about the likelihood of encountering another animal, and thus also about its risk of injury by being in that area. This fits with the observation that scent marks rarely act as an immediate detterrent to invaders. Previous theories on the function of scent marking have involved the idea that this provided territorial information about the location of each individual cat, which may also be tree, although the benefit to the producer of the signal is unclear. The second function of scent marking is that it signals the onset of oestrus in the females. A change in the marking rate of the female was a good indicator of reproduction in a variety of small cat species.

Acoustic communication
Acoustic signals in felids have a wide range of messages, and are used across long distances as well as during close contact in group living felids as well as solitary ones. For example, calls can display territorial advertisment, defensive and offensive threat spit, hiss, growl, snarl, close range affiliation pristen, gurgle, puffing, mating signals, both for sexual advertisement in male and female sexual calls, and during copulation, infant signals of contact purr, meow, and distress meow, identification messages, call sequence duration in lions, and to encourage assembly of a group roaring of lions.  It is impossible to create a detailed manual of felid calls because detailed information on many species is rare. There are mentions of other sounds, but it is not usually possible to tell whether these are distinct sounds or just continuations of a scale of a previously recorded call, or a slight variation between species. The roar is found only in the Pantera lineage. Other differences include the close range friendly affiliation call, which differs in structure across the three lineages, there being three types; gurgle, prusten and puffing, all of which are thought to have the same function in different species. Threat and infant sounds appear to be relatively uniform. The less common calls include the wah wah and the chatter. It is unkown how widespread these two sounds are across lineages.

Visual communication

As in domestic cats, social rolling in wild felids is a factor of sexual behavior, although in captive cats it does also occur in general social situations. There is, however, no evidence of social rolling in wild cats being used in the submissive manner as for the domestic cat. With the exception of TU, which occurs with crouch as a defensive posture, no other tail position appears to act as a signal in wild cats. The one exception is the lion, which has shown a TU position with rubbing, though it was not described as a signal  There has been no study into the use of body and face signals in wild cats, excepting use of the visual signals in the lion.

Tactile communication

Tactile communication in free ranging solitary cats usually occurs a s a component of either mating or mother-young behavior. In social cats, especially the lion and domestic cat, tactile sginals are often used as general social signals and more specifically in a reproductive or parental context. Tactile signals appear to be used in a similar manner in these two social species, despite their different evolutionary lineages.

Social rubbing among small felids has not been documented in the wild. It may be derived from the mating ritual, due to its occurrence during reproductive behavior. Social rubbing among lions has been reported in more detail, occurring as an associated behavior between adults. It occurs particularly after members of the group have been separated, and also after friendly interactions. It indicates that the intention of the animals are peaceful, Males rarely rub on females or cubs, while females rub on both males and females, and cubs rubbed mostly on females. Rubbing acts as a pacifying gesture, producing more benefit for a subordinate animal than for a dominant. This system has also worked for the other group living cat, the domestic cat, Both lions (panthera lineage) and domestic cats (domestic cat lineage) appear to use social rubbing to placate. The signal implies that rubbing may have a similar function in mating behavior among solitary cats, i.e. indicating that the intentions of the animal are peaceful, both before and after copuation. This signal has diversified to be used in other social contexts among two sociable species, F.s. catis and O.leo, depsite their different lineages.

Social grooming in solitary cats occurs both as part of mating behavior, and in mother-young interactions, in which it it practical in maintaining cubs' cleanliness. In the social lion it occurs in these two situations, and in a non specific social situation, often when two are resting together. The function has not been known. Normally it is seen in social cats.

The effect of domestication on cat cat signals

During domestication F. silvestris must have adapted to living at higher densities, and then subsequently adopted group living. Since the signals used by solitary animals have different properties from those needed by group living cats, this move may have led to an evolutionary change in the signalling patterns used by this species. Signals must be formed originally from non signal movement, by ritualisation. Further ritualisation can then happen, where a signal diversifies, creating several functionally distinct signals, via the following stages:

1. Signal occurs in one context only.
2. Signal appears in two contexts, taking on a second function, but remains structurally unchanged.
3. The two signals become structurally distinct in the two contexts.

Domestication can provide an insight into the process of ritualisation of signals, because it is possible to compare the domestic cat with relatives that behave very similarly to its ancestors, thus it is possible to determine whether any diversitfication or ritualisation of signals has occured during domestication. Differences between signals used by the domestic cat and wild cats are differences which may have been caused by domestication, both by changing the situation in whcih intraspecific behavior is expressed, e.g. high local population densities, and by introducing a need for interspecific cat human communication.

1. The evolution fo a new signal from a non-signal behvior: Tail Up

 The action of TU as an intergrative part of urine spraying, is thought to occur in all species of cats. The tail is raises vertically during spraying and is immediately lowered.  In domestic cats, it additionally acts as a friendly signal,  differing from the raised tail that occurs during urine spraying, in both contexts being link to friendly behaviors, particularly social rubbing and occuring for prolonged periods, often remaining upright during movement.

It may have evolved in the domestic cat. There is no evidence in any of the wild species. All species carried out social and object rubbing without raising their tail. This is in contrast to domestic cats, where rubbing is almost exclusively carried out with the tail held vertical. The raised tail during spraying is, in contrast, seen in all species. There is no mention of TU occurring in any context other than urine spraying.

TU may have evolved as a friendly signal during domestication, resulting in increased sociality, which may have caused the necessity of an additional visual signal. It canot be ruled out that TU may have developed at an earlier stage, possibly among one of the wild forms of F. silvestris. There are a few behavioral studies on the undomesticated subspecies of F. silvestris, particualy the African subspecies, which may account for the absence of any mention of TU.

Social behavior in lions

Social rubbing in both mating and general social situations often occurs with tail raised, during head rubbing and anal sniffing contact, the animals raises the tail so that it either arches over their back or tips towards the other animal.  It occurs with friendly behaviors or rubbing and anal sniffing; it is applied in a different way from the raised tail during spraying. Its function in lions may be similar to that in domestic cats,  i.e. friendly signal. The occurrence of TU as a friendly signal is only evident in F.s. catus and P. leo, from different evolutionary lineages, but not in any other undomesticated species of fetid, implies that this signal may have evolved separately in the two species, possibly as a result of similar selective pressures acting only on the two most social species of cats.

Now new behaviors have been found as a result of domestication, despite many quantitative differences in the character of signals. Thus it would be especially interesting if they are found to have evolved as a result of domestication.

2. an established signal diversifies to develop a secondary function, i.e. occurs in a new context but does not change in structure

Social rolling  in wild felids is a sexual signal, occuring as part of the reproductive repertoire. In domestic cats it is still used in this manner, but is also used as a submissive gesture in groups of domestic cats. There is nothing to show that wild cats use social rolling for this function, though it is possible that its role in sexual behavior is a submissive one, in which it is only a small step to its general use as a submissive behavior in groups of domestic cats.

Social rubbing and grooming are also both sexual signals in wild cats. In the domestic cat, they are also used in a general social greeting. This change in context and function could be attributed to domestictation. The use of the behavior in a wide variety of affiliate contexts is likely a natural ability of all felids rather than a product of domestication.

Neotonised signals
Meow, knead and purr are all juvenile behaviors, with the exception of purr, which also occurs in adults. In cat human interactions, adult cats can use all three habitually. Wild cats cannot naturally revert to performing kitten behavior when adult. The domestic cat has evolved, culturally or genetically, the ability to use kitten behaviors towards humans when an adult (neoteny).

3.An established signal diversified in both structure and function to beome a different signal.

There are no sure examples of this in the cat cat relationship, but a change in signal structure does occur in cat human signals.

Rubbing in domestic cat occurs in a human directed situation, rubbing occurs at a higher frequency and at a higher intensity than in a cat cat situation. It is likely to have occurred partly due to the change in receiver psychology, also due to the change in meaning of the signal. It is likely that much of human directed cat behavior is exhibited as either a food or attention-getting signal. This is in contrast to the message given in the cat cat situation, where it acs as a subtle friendly signal, A food eliciting signal would be loud  and prominent, whereas a friendly, cooperative signal between members of a colony would be a subtle cue. The difference in the type of message that is being given by rubbing may cause a difference in frequency and intensity with which the signal is given. This ritualisation of an established cat cat signal in the cat human situation may have also occurred in other common cat human signals such as the meow.

Despite substantial publications on communication of the cat, sveral important issues remain to be resolved. Did everything that has been described as communication really involved transmission of information from one cat to another, and have all the signals produced by the domestic cat been identified? Most signals have been defined on the basis that they have patterns that are obvious to humans and which appear to evoke responses from another cat. However, more study and interpretation of a behavior pattern as a signal requires that it should be tested independently of the context within which it normally occurs.

It is possible that the domestic cat gives subtle signals which have not been identified as such. Cooperative signals may be difficult to detect experimentally, since they should be produced with the minimum amount of energy needed, and should keep the signaller as inconspicuous as possible, to minimise detection by a predator, i.e. the grunts emitted by vervet monkeys, though indistinguishable to humans, are produced in at least two distinct forms with different meanings. Since sociality in the domestic cat may be primitive, and may even have evoled as a consequence of cats' association with humans, but maybe we see the cat's social system as primitive because we have not yet identified all the signals by which relationships are established and maintained, and so do not yet fully comprehend those that we have identified.

It is still unknown, whether the traditional concept of dominants, which are so useful in interpreting the social behavior of other species, can be applied to the cat. Signals, roles of allogrooming and rubbing in redirecting and averting aggression need further study.

Our understanding of the role of scent signals in social behavior has lagged behing that of some other mammals, even though synthetic analogues of the 'facial pheremones' of the cat are now commercially available for controlling indoor urination and aggression. All of the scent marking done by cats is in need of reevaluation in terms of the benefits received by the depositor, and the recipient, as we have attempted to do for spray urination by males.

The cat gives us  many opportunities to examine the effects of domestication on signalling. This may have happened in two non-exclusive ways: either through hard wired changes in the structure and/or meaning of signals which are inherited,or an enhanced ability to learn to communicate in new ways, especially when signalling to humans. The TU signal is an example pf the former, appearing as a method for avoiding unnecessary conflict as cats adapted to living in high densities around human settlement. Neotenisation may have extended the use of some signals, especially vocalisation, from the juvenile to the adult. Vocal signals, most notably meow, since they vary a lot in form from one cat to another, may show an increased adaptability in performance, enabling the development of an interspecific as well as an intraspecific vocabulary.

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Singing Nettles Herbal Clinic by Amanda Dainow is licensed under a Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License.
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