Animal communication

Metacommunications: signals that modify the meaning of subsequent signals. The best known example is the play face and tail signals in dogs, which indicate that a subsequent aggressive signal is part of a play fight rather than a serious aggressive episode.

Animal communication is any behavior on the part of one animal that has an effect on the current or future behaviour of another animal. The study of animal communication, sometimes called Zoosemiotics (distinguishable from anthroposemiotics, the study of human communication) has played an important part in the methodology of ethology, sociobiology, and the study of animal cognition.

Animal communication, and indeed the understanding of the animal world in general, is a rapidly growing field, and even in the 21st century so far, many prior understandings related to diverse fields such as personal symbolic name use, animal emotions, animal culture and learning, and even sexual conduct, long thought to be well understood, have been revolutionized.

Validation

A lamb investigates a rabbit, an example of interspecific communication through body language and scent.

Forms of communication

The best known forms of communication involve the display of distinctive body parts, or distinctive bodily movements; often these occur in combination, so a distinctive movement acts to reveal or emphasise a distinctive body part. An example that was important in the history of ethology was the parent Herring Gull's presentation of its bill to a chick in the nest. Like many gulls, the Herring Gull has a brightly coloured bill, yellow with a red spot on the lower mandible near the tip. When it returns to the nest with food, the parent stands over its chick and taps the bill on the ground in front of it; this elicits a begging response from a hungry chick (pecking at the red spot), which stimulates the parent to regurgitate food in front of it. The complete signal therefore involves a distinctive morphological feature (body part), the red-spotted bill, and a distinctive movement (tapping towards the ground) which makes the red spot highly visible to the chick. Investigations by Niko Tinbergen and his colleagues showed that the red colour of the bill, and its high contrast, are crucial for eliciting the appropriate response from the chick (It is unresolved whether this actually is an inborn behavior in all its complexity, or simply a combination of generalized curiosity on part of the chick, and generalized parental/feeding instincts acting together to produce a simple learning process via reward. Gull chicks peck at everything that are brightly colored, mainly red, yellow, white or shining, and high-contrast objects, but the parent's bill is the only such object that will constantly yield food as a reward when pecked at. Accidental swallowing of pieces of brightly colored plastic or glass is a common cause of mortality amongst gull chicks). Some cephalopods, such as the octopus, have specialized skin cells that can change the apparent colour, opacity, and reflectiveness of their skin.^[1] In addition to being used for camouflage, rapid changes in skin colour are used while hunting and in courtship rituals.^[2] Many animals communicate through vocalisations. One form of animal communication is bird song, often performed by males, though in some species the sexes sing in alternation (this is called duetting). Whale song has been found to have different dialects based on location.^[3] Other instances of communication include the warning cries of many monkeys, the territorial calls of gibbons, and the mating calls of many species of frog.

Less obvious (except in a few cases) is olfactory communication. Many mammals, in particular, have glands that generate distinctive and long-lasting smells, and have corresponding behaviours that leave these smells in places where they have been. Often the scented substance is introduced into urine or feces. Sometimes it is distributed through sweat, though this does not leave a semi-permanent mark as scents deposited on the ground do. Some animals have glands on their bodies whose sole function appears to be to deposit scent marks: for example Mongolian gerbils have a scent gland on their stomachs, and a characteristic ventral rubbing action that deposits scent from it. Golden hamsters and cats have scent glands on their flanks, and deposit scent by rubbing their sides against objects; cats also have scent glands on their foreheads. Bees carry with them a pouch of material from the hive which they release as they reenter, the smell of which indicates that they are a part of the hive and grants their safe entry.

Most of these forms of communication can also be used for interspecies communication.

A rarer form of animal communication is electrocommunication. It is seen primarily in aquatic life, though some mammals, notably the platypus and echidnas are capable of electroreception and thus theoretically of electrocommunication.

Functions of communication

While there are as many kinds of communication as there are kinds of social behaviour, a number of functions have been studied in particular detail. They include:

• agonistic interaction: everything to do with contests and aggression between individuals. Many species have distinctive threat displays that are made during competition over food, mates or territory; much bird song functions in this way. Often there is a matched submission display, which the threatened individual will make if it is acknowledging the social dominance of the threatener; this has the effect of terminating the aggressive episode and allowing the dominant animal unrestricted access to the resource in dispute. Some species also have affiliative displays which are made to indicate that a dominant animal accepts the presence of another.
• courtship rituals: signals made by members of one sex to attract or maintain the attention of potential mate, or to cement a pair bond. These frequently involve the display of body parts, body postures (gazelles assume characteristic poses as a signal to initiate mating), or the emission of scents or calls, that are unique to the species, thus allowing the individuals to avoid mating with members of another species which would be infertile. Animals that form lasting pair bonds often have symmetrical displays that they make to each other: famous examples are the mutual presentation of reeds by Great Crested Grebes, studied by Julian Huxley, the triumph displays shown by many species of geese and penguins on their nest sites and the spectacular courtship displays by birds of paradise and manakins.
• ownership/territorial: signals used to claim or defend a territory, food, or a mate.
• food-related signals: many animals make "food calls" that attract a mate, or offspring, or members of a social group generally to a food source. When parents are feeding offspring, the offspring often have begging responses (particularly when there are many offspring in a clutch or litter - this is well known in altricial songbirds, for example). Perhaps the most elaborate food-related signal is the dance language of honeybees studied by Karl von Frisch.
• alarm calls: signals made in the presence of a threat from a predator, allowing all members of a social group (and often members of other species) to run for cover, become immobile, or gather into a group to reduce the risk of attack.
• metacommunications: signals that modify the meaning of subsequent signals. The best known example is the play face in dogs, which signals that a subsequent aggressive signal is part of a play fight rather than a serious aggressive episode.

Interpretation of animal communication

It is important to note that whilst many gestures and actions have common, stereotypical meanings, researchers regularly seem to find that animal communication is often more complex and subtle than previously believed, and that the same gesture may have multiple distinct meanings depending on context and other behaviors. So generalizations such as "X means Y" are often, but not always accurate. For example, even a simple domestic dog's tail wag may be used in subtly different ways to convey many meanings including:

• Excitement
• Anticipation
• Playfulness
• Contentment/enjoyment
• Relaxation or anxiety
• Questioning another animal or a human as to intentions
• Tentative role assessment on meeting another animal
• Reassurance ("I'm hoping to be friendly, are you?")
• Brief acknowledgement ("I hear you", or "I'm aware and responsive if you want my attention")
• Statement of interest ("I want that (food/toy/activity), if you're willing")
• Uncertainty/apprehension
• Submissive placation (if worried by a more dominant animal)

Combined with other body language, in a specific context, many gestures such as yawns, direction of vision, and so on all convey meaning. Thus statements that a particular action "means" something should always be interpreted to mean "often means" something. As with human beings, who may smile or hug or stand a particular way for multiple reasons, many animals reuse gestures too.

Intraspecies vs. interspecies communication

The sender and receiver of a communication may be of the same species or of different species. The majority of animal communication is intraspecific (between two or more individuals of the same species). However, there are some important instances of interspecific communication. Also, the possibility of interspecific communication, and the form it takes, is an important test of some theoretical models of animal communication.

A European starling (Sturnus vulgaris) singing

Intraspecies communication

The majority of animal communication occurs within a single species, and this is the context in which it has been most intensively studied.

Most of the forms and functions of communication described above are relevant to intra-species communication.

Interspecies communication

Many examples of communication take place between members of different species. Animals communicate to other animals with various signs: visual, sound, echolocation, body language, and smell.

Prey to predator

If a prey animal moves or makes a noise in such a way that a predator can detect and capture it, that fits the definition of "communication" given above. This type of communication is known as interceptive eavesdropping, where a predator intercepts the message being conveyed to conspecifics.

This Chihuahua is baring his teeth to signify an attack is imminent if the photographer comes closer to take his bone.

There are however some actions of prey species that are clearly communications to actual or potential predators. A good example is warning colouration: species such as wasps that are capable of harming potential predators are often brightly coloured, and this modifies the behaviour of the predator, who either instinctively or as the result of experience will avoid attacking such an animal. Some forms of mimicry fall in the same category: for example hoverflies are coloured in the same way as wasps, and although they are unable to sting, the strong avoidance of wasps by predators gives the hoverfly some protection. There are also behavioral changes that act in a similar way to warning colouration. For example, canines such as wolves and coyotes may adopt an aggressive posture, such as growling with their teeth bared, to indicate they will fight if necessary, and rattlesnakes use their well-known rattle to warn potential predators of their poisonous bite. Sometimes, a behavioral change and warning colouration will be combined, as in certain species of amphibians which have a brightly coloured belly, but on which the rest of their body is coloured to blend in with their surroundings. When confronted with a potential threat, they show their belly, indicating that they are poisonous in some way.

Another example of prey to predator communication, is referred to as a pursuit-deterrent signal. Pursuit-deterrent signals occur when prey indicates to a predator that pursuit would be unprofitable because the signaler is prepared to escape. Pursuit-deterrent signals provide a benefit to both the signaler and receiver; they prevent the sender from wasting time and energy fleeing, and they prevent the receiver from investing in a costly pursuit that is unlikely to result in capture. Such signals can advertise prey’s ability to escape, and reflect phenotypic condition (quality advertisement), or can advertise that the prey has detected the predator (perception advertisement). Pursuit-deterrent signals have been reported for a wide variety of taxa, including fish (Godin and Davis 1995), lizards (Cooper et al. 2004), ungulates (Caro 1995), rabbits (Holley 1993), primates (Zuberbuhler et al. 1997), rodents (Shelley and Blumstein 2005, Clark 2005), and birds (Alvarez 1993, Murphy 2006, 2007). The most familiar example of quality advertisement pursuit-deterrent signal is stotting, a pronounced combination of running while simultaneously hopping shown by some antelopes such as Thomson's gazelle in the presence of a predator. At least 11 hypothesis for stotting have been proposed. A leading theory today is that it alerts predators that the element of surprise has been lost. Predators like cheetahs rely on surprise attacks, proven by the fact that chases are rarely successful when they stot. Predators know not to waste energy on a chase that will likely be unsuccessful (optimal foraging behavior).

Predator to prey

Some predators communicate to prey in ways that change their behaviour and make them easier to catch, in effect deceiving them. A well-known example is the angler fish, which has a fleshy growth protruding from its forehead and dangling in front of its jaws; smaller fish try to take the lure, and in so doing are perfectly placed for the angler fish to eat them.

Symbiotic species

Interspecies communication also occurs in various kinds of mutualism and symbiosis. For example, in the cleaner fish/grouper system, groupers signal their availability for cleaning by adopting a particular posture at a cleaning station.

Human/animal communication

Various ways in which humans interpret the behavior of domestic animals, or give commands to them, fit the definition of interspecies communication. Depending on the context, they might be considered to be predator to prey communication, or to reflect forms of commensalism. The recent experiments on animal language are perhaps the most sophisticated attempt yet to establish human/animal communication, though their relation to natural animal communication is uncertain.

Other aspects of animal communication

Evolution of communication

The importance of communication is clear from the fact that animals have evolved elaborate body parts to facilitate it. They include some of the most striking structures in the animal kingdom, such as the peacock's tail. Birdsong appears to have brain structures entirely devoted to its production. But even the red spot on a herring gull's bill, and the modest but characteristic bowing behaviour that displays it, require evolutionary explanation.

There are two aspects to the required explanation:

• identifying a route by which an animal that lacked the relevant feature or behaviour could acquire it;
• identifying the selective pressure that makes it adaptive for animals to develop structures that facilitate communication, emit communications, and respond to them.

Significant contributions to the first of these problems were made by Konrad Lorenz and other early ethologists. By comparing related species within groups, they showed that movements and body parts that in the primitive forms had no communicative function could be "captured" in a context where communication would be functional for one or both partners, and could evolve into a more elaborate, specialised form. For example, Desmond Morris showed in a study of grass finches that a beak-wiping response occurred in a range of species, serving a preening function, but that in some species this had been elaborated into a courtship signal.

The second problem has been more controversial. The early ethologists assumed that communication occurred for the good of the species as a whole, but this would require a process of group selection which is believed to be mathematically impossible in the evolution of sexually reproducing animals. Altruism towards an unrelated group is not widely accepted in the scientific community, but rather can be seen as a sort of reciprocal altruism, expecting the same behavior from others, a benefit of living in a group. Sociobiologists argued that behaviours that benefited a whole group of animals might emerge as a result of selection pressures acting solely on the individual. A gene-centered view of evolution proposes that behaviors that enabled a gene to become wider established within a population would become positively selected for, even if their effect on individuals or the species as a whole was detrimental.^[4] In the case of communication, an important discussion by John Krebs and Richard Dawkins established hypotheses for the evolution of such apparently altruistic or mutualistic communications as alarm calls and courtship signals to emerge under individual selection. This led to the realisation that communication might not always be "honest" (indeed, there are some obvious examples where it is not, as in mimicry). The possibility of evolutionarily stable dishonest communication has been the subject of much controversy, with Amotz Zahavi in particular arguing that it cannot exist in the long term. Sociobiologists have also been concerned with the evolution of apparently excessive signalling structures such as the peacock's tail; it is widely thought that these can only emerge as a result of sexual selection, which can create a positive feedback process that leads to the rapid exaggeration of a characteristic that confers an advantage in a competitive mate-selection situation.

One theory to explain the evolution of traits like a peacock's tail is 'runaway selection'. This requires two traits-a trait that exists, like the bright tail, and a prexisting bias in the female to select for that trait. Females prefer the more elaborate tails, and thus those males are able to mate successfully. Exploiting the psychology of the female, a positive feedback loop is enacted and the tail becomes bigger and brighter. Eventually, the evolution will level off because the survival costs to the male do not allow for the trait to be elaborated any further. Two theories exist to explain runaway selection. The first is the good genes hypothesis. This theory states that an elaborate display is an honest signal of fitness and truly is a better mate. The second is the handicap hypothesis. This explains that the peacock's tail is a handicap, requiring energy to keep and makes it more visible to predators. Regardless, the individual is able to survive, even though its genes are not as good per se.

Cognitive aspects

Ethologists and sociobiologists have characteristically analysed animal communication in terms of more or less automatic responses to stimuli, without raising the question of whether the animals concerned understand the meaning of the signals they emit and receive. That is a key question in animal cognition. There are some signalling systems that seem to demand a more advanced understanding. A much discussed example is the use of alarm calls by vervet monkeys. Robert Seyfarth and Dorothy Cheney showed that these animals emit different alarm calls in the presence of different predators (leopards, eagles, and snakes), and the monkeys that hear the calls respond appropriately - but that this ability develops over time, and also takes into account the experience of the individual emitting the call. Metacommunication, discussed above, also seems to require a more sophisticated cognitive process.

A recently published paper^[5] demonstrated that bottlenose dolphins can recognize identity information from whistles even when otherwise stripped of the characteristics of the whistle; making dolphins the only animals other than humans that have been shown to transmit identity information independent of the caller’s voice or location. The paper concludes that:

The fact that signature whistle shape carries identity information independent from voice features presents the possibility to use these whistles as referential signals, either addressing individuals or referring to them, similar to the use of names in humans. Given the cognitive abilities of bottlenose dolphins, their vocal learning and copying skills, and their fission–fusion social structure, this possibility is an intriguing one that demands further investigation.

—V. M. Janik, et al. ^[5]

Animal communication and human behaviour

Another controversial issue is the extent to which humans have behaviours that resemble animal communication, or whether all such communication has disappeared as a result of our linguistic capacity. Some of our bodily features - eyebrows, beards and moustaches, deep adult male voices, perhaps female breasts - strongly resemble adaptations to producing signals. Ethologists such as Irenäus Eibl-Eibesfeldt have argued that facial gestures such as smiling, grimacing, and the eyebrow flash on greeting are universal human communicative signals that can be related to corresponding signals in other primates. Given the recency with which spoken language has emerged, it is likely that human body language does include some more or less involuntary responses that have a similar origin to the communication we see in other animals.

Humans also often seek to mimic animals' communicative signals in order to interact with the animals. For example, cats have a mild affiliative response involving closing their eyes; humans often close their eyes towards a pet cat to establish a tolerant relationship. Stroking, petting and rubbing pet animals are all actions that probably work through their natural patterns of interspecific communication.

Dogs have shown an ability to understand communication from a species other than their own. They were able to use human communicative gestures such as pointing and looking to find hidden food and toys^[6].

Animal communication and linguistics

For linguistics, the interest of animal communication systems lies in their similarities to and differences from human language:

1. Human languages are characterized for having a double articulation (in the characterization of French linguist André Martinet). It means that complex linguistic expressions can be broken down in meaningful elements (such as morphemes and words), which in turn are composed of smallest phonetic elements that affect meaning, called phonemes. Animal signals, however, do not exhibit this dual structure.
2. In general, animal utterances are responses to external stimuli, and do not refer to matters removed in time and space. Matters of relevance at a distance, such as distant food sources, tend to be indicated to other individuals by body language instead, for example wolf activity before a hunt, or the information conveyed in honeybee dance language. It is therefore unclear to what extent utterances are automatic responses and to what extent deliberate intent plays a part.
3. Human language is largely learned culturally, while animal communication systems are known largely by instinct.
4. In contrast to human language, animal communication systems are usually not able to express conceptual generalizations. (Cetaceans and some primates may be notable exceptions).
5. Human languages combine elements to produce new messages (a property known as creativity). One factor in this is that much human language growth is based upon conceptual ideas and hypothetical structures, both being far greater capabilities in humans than animals. This appears far less common in animal communication systems, although current research into animal culture is still an ongoing process with many new discoveries.

A recent and interesting area of development is the discovery that the use of syntax in language, and the ability to produce "sentences", is not limited to humans either.^[7] The first good evidence of syntax in non-humans, reported^[8] in 2006, is from the greater spot-nosed monkey (Cercopithecus Nictitans) of Nigeria. This is the first evidence that some animals can take discrete units of communication, and build them up into a sequence which then carries a different meaning from the individual "words":

The putty-nosed monkeys have two main alarm sounds. A sound known onomatopoeiacally as the 'pyow' warns against a lurking leopard, and a coughing sound that scientists call a 'hack' is used when an eagle is hovering nearby.

"Observationally and experimentally we have demonstrated that this sequence [of up to three 'pyows' followed by up to four 'hacks'] serves to elicit group movement... the 'pyow-hack' sequence means something like "let's go!" [a command telling others to move]... The implications are that primates at least may be able to ignore the usual relationship between an individual alarm call, and the meaning it might convey under certain circumstances... To our knowledge this is the first good evidence of a syntax-like natural communication system in a non-human species."

Similar results have also recently been reported in the Campbell's Mona Monkey^[9]

See also

• Animal behavior
• Biocommunication
• Biosemiotics
• Emotion in animals
• Forms of activity and interpersonal relations
• International Society for Biosemiotic Studies
• Sir Philip Sidney game
• Zoomusicology

References

External links

• The Elgin Center for Zoosemiotic Research
• Zoosemiotics: animal communication on the web
• The Animal Communication Project
• International Bioacoustics Council research on animal language.
• Animal Sounds different animal sounds to listen and download.
• The British Library Sound Archive contains over 150,000 recordings of animal sounds and natural atmospheres from all over the world.