Alarm signal
![](http://upload.wikimedia.org/wikipedia/commons/thumb/f/f1/Spermophilus_beldingi.jpg/220px-Spermophilus_beldingi.jpg)
In
Different calls may be used for predators on the ground or from the air. Often, the animals can tell which member of the group is making the call, so that they can disregard those of little reliability.[1]
Evidently, alarm signals promote survival by allowing the receivers of the alarm to escape from the source of peril; this can evolve by kin selection, assuming the receivers are related to the signaller. However, alarm calls can increase individual fitness, for example by informing the predator it has been detected.[2]
Alarm calls are often high-frequency sounds because these sounds are harder to localize.[3][4]
Selective advantage
This cost/benefit tradeoff of alarm calling behaviour has sparked many interest debates among
Altruism
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Some scientists have used the evidence of alarm-calling behaviour to challenge the theory that "evolution works only/primarily at the level of the gene and of the gene's 'interest' in passing itself along to future generations." If alarm-calling is truly an example of altruism, then human understanding of natural selection becomes more complicated than simply "survival of the fittest gene".
Other researchers, generally those who support the
Predator-directed signaling
Another theory suggests that alarm signals function to attract further predators, which fight over the prey organism, giving it a better chance of escape.[6] Others still suggest they are a deterrent to predators, communicating the prey's alertness to the predator. One such case is the western swamphen (Porphyrio porphyrio), which gives conspicuous visual tail flicks (see also aposematism, handicap principle and stotting).[7]
Further research
Considerable research effort continues to be directed toward the purpose and ramifications of alarm-calling behaviour, because, to the extent that this research has the ability to comment on the occurrence or non-occurrence of altruistic behaviour, these findings can be applied to the understanding of altruism in human behaviour.
Monkeys with alarm calls
![](http://upload.wikimedia.org/wikipedia/commons/thumb/a/a0/Chlorocebus_pygerythrus.jpg/200px-Chlorocebus_pygerythrus.jpg)
Vervet monkeys
Vervet monkeys (Chlorocebus Pygerythus) are some of the most studied monkeys when it comes to vocalization and alarm calls within the nonhuman primates. They are most known for making alarm calls in the presence of their most common predators (leopards, eagles, and snakes). Alarm calls of the vervet monkey are considered arbitrary in relation to the predator that they signify, in the sense that while the calls may be distinct to the threat that the monkeys are perceiving, the calls do not mimic the actual sounds of the predator – it is like yelling "Danger!" when seeing an angry dog rather than making barking sounds. This type of alarm calls is seen as the earliest example of symbolic communication (the relationship between signifier and signified is arbitrary and purely conventional) in nonhuman primates.[8]
However, there is much debate on whether the vervet monkeys alarm calls are actual "words" in the sense of purposely manipulating sounds to communicate specific meaning or are unintentional sounds t hat are made when interacting with an outside stimulus. Like small children who cannot communicate words effectively make random noises when being played with or are stimulated by something in their immediate environment. As children grow and begin learning how to communicate the noises, they make are very broad in relation to their environment. They begin to recognize the things in their environment but there more things than known words or noises so a certain sound may reference multiple things. As children get older, they can become more specific about the noises and words made in relation to the things in their environment. It is thought that as Vervet monkeys get older they are able to learn and break the broad categories into more specific sub categories to a specific context.[9]
In an experiment conducted by Dr. Tabitha Price, they used custom software to gather the acoustic sounds of male and female Vervet monkeys from East Africa and male Vervet monkey from South Africa. The point of the experiment was to gather the acoustic sounds of these monkeys when stimulated by the presence of snakes (mainly Python), raptors, terrestrial animals (mostly Leopards), and aggression. Then to determine if the calls could be distinguished with a known context.
The experiment determined that while the Vervet monkeys were able to categorize different predators and members of different social groups, however their ability to communicate specific threats is not proven. The chirps and barks that Vervet monkeys make as an eagle swoops in are the same chirps and barks that are made in moments of high arousal. Similarly, the barks made for leopards are the same that are made during aggressive interactions. The environment that they exist in is too complex for their ability to communicate about everything in their environment specifically.[10]
In an experiment conducted by Dr. Julia Fischer, a drone was flown over Vervet monkeys and recorded the sounds produced. The Vervet monkeys made alarm calls that were almost identical to the eagle calls of East African Vervets. When a sound recording of the drone was played back a few days later to a monkey that was alone and away from the main group it looked up and scanned the sky. Dr. Fischer concluded that Vervet monkeys can be exposed to a new threat once and understand what it means.
It is still debated whether or not Vervet monkeys are actually aware of what the alarm calls mean. One side of the argument is that the monkeys give alarm calls because they are simply excited. The other side of the argument is that the alarm calls create mental representation of predators in the listeners minds. The common middle ground argument is that they give alarm calls because they want others to elicit a certain response, not necessarily because they want the group to think that there is a specific threat near.[9]
Ultimately there is not enough evidence to support whether or not the calls are simply identifying a threat or calling for specific action due to the threat.
Campbell's mona monkeys
Campbell's mona monkeys also generate alarm calls, but in a different way than vervet monkeys. Instead of having discrete calls for each predator, Campbell monkeys have two distinct types of calls which contain different calls which consist in an acoustic continuum of affixes which change meaning. It has been suggested that this is a homology to human morphology.[11] Similarly, the cotton-top tamarin is able to use a limited vocal range of alarm calls to distinguish between aerial and land predators.[12] Both the Campbell monkey and the cotton-top tamarin have demonstrated abilities similar to vervet monkeys' ability to distinguish likely direction of predation and appropriate responses.[13][14]
That these three
Barbary macaque
Another species that exhibits alarm calls is the Barbary macaque. Barbary macaque mothers are able to recognize their own offspring's calls and behave accordingly.[16]
Diana monkeys
Diana monkeys also produce alarm signals. Adult males respond to each other's calls, showing that calling can be contagious.[17] Their calls differ based on signaller sex, threat type, habitat, and caller ontogenetic or lifetime predator experience.
Diana monkeys emit different alarm calls as a result of their sex. Male alarm calls are primarily used for resource defence, male–male competition, and communication between groups of conspecifics.[18] Female alarm calls are mainly used for communication within groups of conspecifics to avoid predation.[19]
Alarm calls are also predator-specific. In
In Taï National Park and Tiwai Island, Sierra Leone, specific acoustic markers in the alarm calls of Diana monkeys convey both threat type and caller familiarity information to a receiver. In Taï National Park, males respond to eagle alarm signals based on predator type and caller familiarity. When the caller is unfamiliar to the receiver, the response call is a 'standard' eagle alarm call, characterized by a lack of frequency transition at the onset of the call.[17] When the caller is familiar, the response call is an atypical eagle alarm call, characterized by a frequency transition at onset, and the response is faster than to that of an unfamiliar caller.[17] On Tiwai Island, males respond in the opposite way to eagle alarm signals.[17] When the caller is familiar, the response call is a 'standard' eagle alarm call, without a frequency transition at onset. When the caller is unfamiliar, the response call is an atypical eagle alarm call, with a frequency transition at onset.[17]
The differences in alarm call responses are due to differences in habitat. In Taï National Park, there is a low predation risk from eagles, high primate abundance, strong intergroup competition, and a tendency for group encounters to result in high levels of aggression.[17] Therefore, even familiar males are a threat to whom males respond with aggression and an atypical eagle alarm call.[17] Only unfamiliar males, who are likely to be solitary and non-threatening, do not receive an aggressive response and receive only a typical alarm call.[17] On Tiwai Island, there is a high predation risk from eagles, low primate abundance, a tendency for group encounters to result in peaceful retreats, low resource competition, and frequent sharing of foraging areas.[17] Therefore, there is a lack of aggression towards familiar conspecifics to whom receivers respond with a 'standard' eagle call.[17] There is only aggression towards unfamiliar conspecifics, to whom receivers respond with an atypical call.[17] Simply put, a response with a typical eagle alarm call prioritizes the risk of predation, while a response with an atypical alarm call prioritizes social aggression.[17]
Diana monkeys also display a predisposition for flexibility in acoustic variation of alarm call assembly related to caller
Sexual selection for predator-specific alarm signals
In guenons, selection is responsible for the evolution of predator-specific alarm calls from loud calls. Loud calls travel long distances, greater than that of the home range, and can be used as beneficial alarm calls to warn conspecifics or showcase their awareness of and deter a predator.[24][25][26] A spectrogram of a subadult male call shows that the call is a composition of elements from a female alarm call and male loud call, suggesting the transition from the latter to the former during puberty and suggesting that alarm calls gave rise to loud calls through sexual selection.[26] Evidence of sexual selection in loud calls includes structural adaptations for long-range communication, co-incidence of loud calls and sexual maturity, and sexual dimorphism in loud calls.[26]
Controversy over the semantic properties of alarm calls
Not all scholars of animal communication accept the interpretation of alarm signals in monkeys as having semantic properties or transmitting "information". Prominent spokespersons for this opposing view are Michael Owren and Drew Rendall,[27][28] whose work on this topic has been widely cited and debated.[29][30] The alternative to the semantic interpretation of monkey alarm signals as suggested in the cited works is that animal communication is primarily a matter of influence rather than information, and that vocal alarm signals are essentially emotional expressions influencing the animals that hear them. In this view monkeys do not designate predators by naming them, but may react with different degrees of vocal alarm depending on the nature of the predator and its nearness on detection, as well as by producing different types of vocalization under the influence of the monkey's state and movement during the different types of escape required by different predators. Other monkeys may learn to use these emotional cues along with the escape behaviour of the alarm signaller to help make a good decision about the best escape route for themselves, without there having been any naming of predators.
Chimpanzees with alarm calls
Receiver knowledge
Alarm signalling varies depending on the receiver's knowledge of a certain threat. Chimpanzees are significantly more likely to produce an alarm call when conspecifics are unaware of a potential threat or were not nearby when a previous alarm call was emitted.[33] When judging if conspecifics are unaware of potential dangers, chimpanzees do not solely look for behavioural cues, but also assess receiver mental states and use this information to target signalling and monitoring.[34] In a recent[when?] experiment, caller chimpanzees were shown a fake snake as a predator and were played pre-recorded calls from receivers. Some receivers emitted calls that were snake-related, and therefore represented receivers with knowledge of the predator, while other receivers emitted calls that were not snake-related, and therefore represented receivers without knowledge of the predator.[34] In response to the non-snake-related calls from receivers, the signallers increased their vocal and nonvocal signalling and coupled it with increased receiver monitoring.[34]
Caller age
Chimpanzee age impacts the frequency of alarm signalling. Chimpanzees over 80 months of age are more likely to produce an alarm call than those less than 80 months of age.[35] There are several hypotheses for this lack of alarm calling in infants zero to four years of age.[35] The first hypothesis is a lack of motivation to produce alarm calls because of mothers in close proximity that minimize the infant's perception of a threat or that respond to a threat before the infant can.[35] Infants may also be more likely to use distress calls to catch their mother's attention in order for her to produce an alarm call.[35] Infants might also lack the physical ability to produce alarm calls or lack the necessary experience to classify unfamiliar objects as dangerous and worthy of an alarm signal.[35] Therefore, alarm calling may require advanced levels of development, perception, categorization, and social cognition.[35]
Other factors
Other factors, such as signaller arousal, receiver identity, or increased risk of predation from calling, do not have a significant effect on the frequency of alarm call production.[33][34]
Receiver monitoring
However, while alarm signals can be coupled with receiver monitoring, there is a lack of consensus on the definition, starting age, and purpose of monitoring. It is either defined as the use of three subsequent gaze alternations, from a threat to a nearby conspecific and back to the threat, or as the use of two gaze alternations.[35] Moreover, while some studies only report gaze alternation as starting in late juveniles, other studies report gaze alternation in infants as early as five months of age.[35] In infants and juveniles, it is potentially a means of social referencing or social learning through which younger chimpanzees check the reactions of more experienced conspecifics in order to learn about new situations, such as potential threats.[35] It has also been proposed to be a communicative behaviour or simply the result of shifts in attention between different environmental elements.[32][35]
Evolution of hominoid communication
The evolution of hominoid communication is evident through chimpanzee 'hoo' vocalizations and alarm calls. Researchers propose that communication evolved as natural selection diversified 'hoo' vocalizations into context-dependent 'hoos' for travel, rest, and threats.[36] Context-dependent communication is beneficial and likely maintained by selection as it facilities cooperative activities and social cohesion between signallers and receivers that can increase the likelihood of survival.[36] Alarm calls in chimpanzees also point to the evolution of hominoid language. Callers assess conspecifics' knowledge of threats, fill their need for information, and, in doing so, use social cues and intentionality to inform communication.[32][34] Filling a gap in information and incorporating social cues and intentionality into communication are all components of human language.[32][34] These shared elements between chimpanzee and human communication suggest an evolutionary basis, most likely that the last common human ancestor with chimpanzees also possessed these linguistic abilities.[32][34][37]
False alarm calls
![](http://upload.wikimedia.org/wikipedia/commons/thumb/a/a6/Barn_Swallow-Mindaugas_Urbonas.jpg/200px-Barn_Swallow-Mindaugas_Urbonas.jpg)
Counterfeit alarm calls are also used by thrushes to avoid intraspecific competition. By sounding a bogus alarm call normally used to warn of aerial predators, they can frighten other birds away, allowing them to eat undisturbed.[40]
Vervets seem to be able to understand the referent of alarm calls instead of merely the acoustic properties, and if another species' specific alarm call (terrestrial or aerial predator, for instance) is used incorrectly with too high of a regularity, the vervet will learn to ignore the analogous vervet call as well.[41]
Alarm pheromones
Alarm signals need not be communicated only by
Deceptive chemical alarm signals are also employed. For example, the wild potato, Solanum berthaultii, emits the aphid alarm-pheromone, (E)-β-farnesene, from its leaves, which functions as a repellent against the green peach aphid, Myzus persicae.[46]
See also
- Group selection
- Kin selection
- Mobbing call
References
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External links
- Chickadees' alarm-call carry information about size, threat of predator
- The Trek of the Pika "A story complete with sounds of pika and marmot calls" 2002-10-30
- Characteristics of arctic ground squirrel alarm calls Oecologia Volume 7, Number 2 / June, 1971
- Why do Yellow-bellied Marmots Call? Daniel T. Blumstein & Kenneth B. Armitage
Department of Systematics and Ecology, University of Kansas