Alarm signal

Alarm calls have been studied in many species, such as Belding's ground squirrels.

In the field of animal communication, an alarm signal is an antipredator adaptation referring to various signals emitted by social animals in response to danger. Many primates and birds have elaborate alarm calls for warning conspecifics of approaching predators. For example, the characteristic alarm call of the blackbird is a familiar sound in many gardens. Other animals, like fish and insects, may use other non-auditory signals, such as chemical messages. While visual signs have been suggested as alarm signals, they are easier to pinpoint by predators and less likely to be received by conspecifics, so have tended to be treated as a signal to the predator instead. An animal who signals an alarm is called an alarmer.

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.

Evidently, alarm signals promote survival by allowing the receivers of the alarm to escape from the source of peril, but this ecological safety system may come at a cost to the signaller. When an animal calls attention to itself by signalling, it may be more likely to be eaten by a predator than if it had kept quiet. This intuition has been verified by experimental data on ground squirrel predation rates and the connection between this and the noisy chirping or whistling alarm calls. However, there is also some evidence that alarm calls can increase individual fitness as well.[1]

Alarm calls are often high-frequency sounds because these sounds are harder to localize, despite common belief.[2][3]

Selective advantage

This cost/benefit tradeoff of alarm calling behaviour has sparked many interest debates among evolutionary biologists seeking to explain the occurrence of such apparently "self-sacrificing" behaviour. The central question is this: "If the ultimate purpose of any animal behaviour is to maximize the chances that an organism's own genes are passed on, with maximum fruitfulness, to future generations, why would an individual deliberately risk destroying itself (their entire genome) for the sake of saving others (other genomes)?".

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 our understanding of natural selection becomes more complicated than simply "survival of the fittest gene".

Other researchers, generally those who support the selfish gene theory, question the authenticity of this "altruistic" behaviour. For instance, it has been observed that vervets sometimes emit calls in the presence of a predator, and sometimes do not. Studies show that these vervets may call more often when they are surrounded by their own offspring and by other relatives who share many of their genes.[4] Other researchers have shown that some forms of alarm calling, for example, "aerial predator whistles" produced by Belding's ground squirrels, do not increase the chances that a caller will get eaten by a predator; the alarm call is advantageous to both caller and recipient by frightening and warding off the predator.

Another theory suggests that alarm signals function to attract further predators, which fight over the prey organism, giving it a better chance of escape.[5] Others still suggest they are a deterrent to predators, communicating the animals alertness to the predator. One such case is the eastern swamphen (Porphyrio porphyrio), which gives conspicuous visual tail flicks (see also aposematism, handicap principle and stotting).[6]

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, we can apply these findings to our understanding of altruism in human behaviour.

Monkeys with alarm calls

Vervet monkey in Dar es Salaam.

Vervet monkeys are the typical example of both animal alarm calls and of semantic capacity in non-human animals. They have three distinct calls for leopards, snakes, and eagles, and research shows that each call elicits different responses. When vervets are on the ground they respond to the eagle alarm call by looking up and running to cover, to leopard alarm calls primarily by looking up and running into a tree, and to the snake alarm call primarily by looking down. When in trees vervets responded to the eagle alarm call by looking up and down and running out of trees, to the leopard alarm call by running higher in the tree and looking both up and down, and to the snake alarm call by looking primarily down.[7][8]

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.[9] Similarly, the cotton-top tamarin is able to use a limited vocal range of alarm calls to distinguish between aerial and land predators.[10] 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.[11][12]

That these three species use vocalizations to warn others of danger has been called by some proof of proto-language in primates. However, there is some evidence that this behavior does not refer to the predators themselves but to threat, distinguishing calls from words.[13]

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.[14]

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,[15][16] whose work on this topic has been widely cited and debated.[17][18] 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 behavior of the alarm signaler to help make a good decision about the best escape route for themselves, without there having been any naming of predators.

False alarm calls

Deceptive vocalizations are given by male barn swallows.

Deceptive alarm calls are used by male swallows (Hirundo rustica).[19] Males give these false alarm calls when females leave the nest area during the mating season, and are thus able to disrupt extra-pair copulations. As this is likely to be costly to females, it can be seen as an example of sexual conflict.[20]

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.[21]

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.[22]

Alarm pheromones

Alarm signals need not be communicated only by auditory means. For example, many animals may use chemosensory alarm signals, communicated by chemicals known as pheromones. Minnows and catfish release alarm pheromones (Schreckstoff) when injured, which cause nearby fish to hide in dense schools near the bottom.[23] Animals are not the only organism to communicate threats to conspecifics either; some plants are able to perform a similar trick. Lima beans release volatile chemical signals that are received by nearby plants of the same species when infested with spider mites. This 'message' allows the recipients to prepare themselves by activating defense genes, making them less vulnerable to attack, and also attracting another mite species that is a predator of spider mites (indirect defence). Although it is conceivable that other plants are only intercepting a message primarily functioning to attract "bodyguards", some plants spread this signal on to others themselves, suggesting an indirect benefit from increased inclusive fitness.[24]

False chemical alarm signals are also employed. The aphid Myzus persicae is repelled by the wild potato Solanum berthaultii which releases a chemical from its leaves that acts as an allomone to disrupt aphid attacks.[25]

See also

References

  1. ↑ "The Predator Deterrence Function of Primate Alarm Calls". Ethology. 105: 477–490. doi:10.1046/j.1439-0310.1999.00396.x.
  2. ↑ http://mue.music.miami.edu/thesis/robert_hartman/robert_hartman_thesis.pdf
  3. ↑ http://primatology.net/2011/03/09/the-semantics-of-vervet-monkey-alarm-calls-part-i/
  4. ↑ "Vervet Monkey Alarm Calls: Manipulation through Shared Information?". 94: 150–166. JSTOR 4534456. Retrieved 22 May 2013.
  5. ↑ Chivers, D. P.; Brown, G. E.; Smith, R. J. F. (1996). "The Evolution of Chemical Alarm Signals: Attracting Predators Benefits Alarm Signal Senders". The American Naturalist. 148 (4): 649–659. doi:10.1086/285945.
  6. ↑ Woodland, D. J.; Jaafar, Z.; Knight, M. (1980). "The "Pursuit Deterrent" Function of Alarm Signals". The American Naturalist. 115 (5): 748–753. doi:10.1086/283596.
  7. ↑ Seyfarth, R. M.; Cheney, D. L.; Marler, P. (1980). "Monkey responses to three different alarm calls: evidence of predator classification and semantic communication". Science. 210 (4471): 801–803. PMID 7433999. doi:10.1126/science.7433999.
  8. ↑ Cheney, D. L.; Seyfarth, R. M. (1981). "Selective Forces Affecting the Predator Alarm Calls of Vervet Monkeys". Behaviour. 76 (1): 25–61. JSTOR 4534091. doi:10.1163/156853981x00022.
  9. ↑ http://www.plosone.org/article/info:doi/10.1371/journal.pone.0007808
  10. ↑ Neyman, P. F. 1978. Aspects of the ecology and social organization of free-ranging cotton-top tamarins (Saguinus oedipus) and the conservation status of the species. In: The Biology and Conservation of the Callitrichidae (Ed. by D. G. Kleiman), pp. 39e71. Washington, D.C.: Smithsonian Institution Press.
  11. ↑ Fichtel, C; Perry, S; Groslouis, J (2005). "Alarm calls of white-faced capuchin monkeys: an acoustic analysis". Animal Behaviour. 70: 165–176. doi:10.1016/j.anbehav.2004.09.020.
  12. ↑ Sproul et al. Cottontop tamarin, Saguinus oedipus, alarm calls contain sufficient information for recognition of individual identity. Animal Behaviour (2006) vol. 72 (6) pp. 1379-1385
  13. ↑ Zuberbühler, K (2000). "Referential labelling in Diana monkeys". Animal Behaviour. 59 (5): 917–927. PMID 10860519. doi:10.1006/anbe.1999.1317.
  14. ↑ Hammerschmidt, Kurt; Todt, Dietmar (1995). "Individual Differences in Vocalisations of Young Barbary Macaques (Macaca Sylvanus): A Multi-Parametric Analysis To Identify Critical Cues in Acoustic Signalling". Behaviour. 132 (5): 381–99. doi:10.1163/156853995x00621.
  15. ↑ Rendall, D.; Owren, M. J.; Ryan, M. J. (2009). "What do animal signals mean?". Animal Behaviour. 78: 233–240. doi:10.1016/j.anbehav.2009.06.007.
  16. ↑ Owren, M. J.; Rendall, D.; Ryan, M. J. (2010). "Redefining animal signaling: Influence versus information in communication". Biology and Philosophy. 25: 755–780. doi:10.1007/s10539-010-9224-4.
  17. ↑ Nordell, S. & Valone, T. (2014). Animal Behaviour: Concepts, Methods and Applications. Oxford University Press.
  18. ↑ Stegmann, U. E. editor (2013). Animal Communication Theory: Information and Influence. Cambridge University Press, Cambridge, UK. 452 pages
  19. ↑ Møller, A. P. (1990). "Deceptive use of alarm calls by male swallows Hirundo rustica: A new paternity guard". Behavioral Ecology. 1: 1–6. doi:10.1093/beheco/1.1.1.
  20. ↑ Arnqvist, G. & Rowe, L. (2005) Sexual conflict. Princeton University Press, Princeton ISBN 0-691-12217-2
  21. ↑ Wickler, W. (1968) Mimicry in Plants and Animals (Translated from the German) McGraw-Hill, New York. ISBN 0-07-070100-8. p. 108.
  22. ↑ Seyfarth, R; Cheney, D (1990). "The assessment by vervet monkeys of their own and another species' alarm calls". Animal Behaviour. 40 (4): 754–764. doi:10.1016/S0003-3472(05)80704-3.
  23. ↑ Campbell, N. & Reece, J. 2004. Biology 7th Edition - Benjamin Cummings ISBN 0-8053-7146-X
  24. ↑ Kobayashi, Y.; Yamamura, N. (2007). "Evolution of signal emission by uninfested plants to help nearby infested relatives". Evolutionary Ecology. 21 (3): 281–294. doi:10.1007/s10682-007-9165-9.
  25. ↑ Gibson, R.W.; Pickett, J.A. (1983). "Wild potato repels aphids by release of aphid alarm pheromone". Nature. 302 (5909): 608–609. doi:10.1038/302608a0.

Department of Systematics and Ecology, University of Kansas

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