Animal communication

Animal communication is any behaviour on the part of one animal that has an effect on the current or future behaviour of another animal. The study of animal communication, called zoosemiotics (distinguishable from anthroposemiotics, the study of human communication) has played an important part in the development 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.

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.



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. Nonetheless, we do not feel comfortable talking about it as communication. Our discomfort suggests that we should modify the definition of communication in some way, either by saying that communication should generally be to the adaptive advantage of the communicator, or by saying that it involves something more than the inevitable consequence of the animal going about its ordinary life.

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.

A more controversial example of prey to predator communication is stotting, a highly noticeable form of running shown by some antelopes such as Thomson's gazelle in the presence of a predator; it has been argued that this demonstrates to the predator that the particular prey individual is fit and healthy and therefore not worth pursuing.

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 behaviour of domestic animals, or give commands to them, fit the definition of interspecific 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.

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

Forms of communication
Most of the following forms of communication can also be used for interspecific 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 is brightly colored, mainly red, yellow, white or shining, 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).

Another important forms of communication is bird song, usually performed mainly by males, though in some species the sexes sing in alternation (this is called duetting and serves mainly purposes of strengthening pair-bonding and repelling competitors). Bird song is just the best known case of vocal communication; other instances 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 if they are a part of the hive and grants their safe entry.

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 weed 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.
 * 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.

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 not just peripheral but also 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. It was the fundamental insight of sociobiology 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. In the case of communication, an important discussion by John R. 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.

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 subtlely different ways to convey many meanings including: 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.
 * 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)

Communication and understanding
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. Richard 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.

Recently, research has demonstrated a further speculated concept in animal communication - that at least one species other than humans uses symbolic, personal, names. At present this has been identified only in dolphins, who use whistling communication to convey information including the equivalent of personal names. The names are specific to individuals, who will respond even when voice, speaker, inflection and other cues are removed from the sound. .

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 Iraneaus Eibl-Eibesfeldt have argued that facial gestures such as smiling, grimacing, and the eye-brow 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.

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

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. The first good evidence of syntax in non-humans, reported in 2006, is from the putty-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":
 * 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 meaningless phonetic elements, or 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.


 * 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."