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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.<ref name="recognition">{{cite journal | last1 = Hammerschmidt | first1 = Kurt | last2 = Todt | first2 = Dietmar | year = 1995 | title = Individual Differences in Vocalisations of Young Barbary Macaques (Macaca Sylvanus): A Multi-Parametric Analysis To Identify Critical Cues in Acoustic Signalling | url = | journal = Behaviour | volume = 132 | issue = 5| pages = 381–99 | doi=10.1163/156853995x00621}}</ref>
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.<ref name="recognition">{{cite journal | last1 = Hammerschmidt | first1 = Kurt | last2 = Todt | first2 = Dietmar | year = 1995 | title = Individual Differences in Vocalisations of Young Barbary Macaques (Macaca Sylvanus): A Multi-Parametric Analysis To Identify Critical Cues in Acoustic Signalling | url = | journal = Behaviour | volume = 132 | issue = 5| pages = 381–99 | doi=10.1163/156853995x00621}}</ref>


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,<ref>{{cite journal | last1 = Rendall | first1 = D. | last2 = Owren | first2 = M. J. | last3 = Ryan | first3 = M. J. | year = 2009 | title = What do animal signals mean? | url = | journal = Animal Behaviour | volume = 78 | issue = 2| pages = 233–240 | doi=10.1016/j.anbehav.2009.06.007}}</ref><ref>{{cite journal | last1 = Owren | first1 = M. J. | last2 = Rendall | first2 = D. | last3 = Ryan | first3 = M. J. | year = 2010 | title = Redefining animal signaling: Influence versus information in communication | url = | journal = Biology and Philosophy | volume = 25 | issue = 5| pages = 755–780 | doi=10.1007/s10539-010-9224-4}}</ref> whose work on this topic has been widely cited and debated.<ref>Nordell, S. & Valone, T. (2014). Animal Behaviour: Concepts, Methods and Applications. Oxford University Press.</ref><ref>Stegmann, U. E. editor (2013). Animal Communication Theory: Information and Influence. Cambridge University Press, Cambridge, UK. 452 pages</ref> 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.
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,<ref>{{cite journal | last1 = Rendall | first1 = D. | last2 = Owren | first2 = M. J. | last3 = Ryan | first3 = M. J. | year = 2009 | title = What do animal signals mean? | url = | journal = Animal Behaviour | volume = 78 | issue = 2| pages = 233–240 | doi=10.1016/j.anbehav.2009.06.007}}</ref><ref>{{cite journal | last1 = Owren | first1 = M. J. | last2 = Rendall | first2 = D. | last3 = Ryan | first3 = M. J. | year = 2010 | title = Redefining animal signaling: Influence versus information in communication | url = | journal = Biology and Philosophy | volume = 25 | issue = 5| pages = 755–780 | doi=10.1007/s10539-010-9224-4}}</ref> whose work on this topic has been widely cited and debated.<ref>Nordell, S. & Valone, T. (2014). Animal Behaviour: Concepts, Methods and Applications. Oxford University Press.</ref><ref>Stegmann, U. E. editor (2013). Animal Communication Theory: Information and Influence. Cambridge University Press, Cambridge, UK. 452 pages</ref> 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==
[[File:Chimpanzee (150887903).jpeg|alt=|thumb|200x200px|Chimpanzees produce alarm calls in response to threats.]]
[[Chimpanzee]]<nowiki/>s emit alarm calls in response to predators, such as leopards and snakes.<ref>{{Cite journal|last=Zuberbühler|first=Klaus|date=2000-01|title=Causal knowledge of predators' behaviour in wild Diana monkeys|url=http://dx.doi.org/10.1006/anbe.1999.1296|journal=Animal Behaviour|volume=59|issue=1|pages=209–220|doi=10.1006/anbe.1999.1296|issn=0003-3472}}</ref><ref>{{Cite journal|last=Crockford|first=Catherine|last2=Boesch|first2=Christophe|date=2003-07|title=Context-specific calls in wild chimpanzees, Pan troglodytes verus: analysis of barks|url=http://dx.doi.org/10.1006/anbe.2003.2166|journal=Animal Behaviour|volume=66|issue=1|pages=115–125|doi=10.1006/anbe.2003.2166|issn=0003-3472}}</ref> They produce three types of alarm calls: acoustically-variable 'hoos', 'barks', and 'SOS screams'.<ref name=":0">{{Cite journal|last=Schel|first=Anne Marijke|last2=Townsend|first2=Simon W.|last3=Machanda|first3=Zarin|last4=Zuberbühler|first4=Klaus|last5=Slocombe|first5=Katie E.|date=2013-10-16|title=Chimpanzee Alarm Call Production Meets Key Criteria for Intentionality|url=http://dx.doi.org/10.1371/journal.pone.0076674|journal=PLoS ONE|volume=8|issue=10|pages=e76674|doi=10.1371/journal.pone.0076674|issn=1932-6203}}</ref> Alarm signalling is impacted by receiver knowledge and caller age, can be coupled with receiver monitoring, and is important to the understanding of the evolution of [[hominoid]] communication.

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.<ref name=":1">{{Cite journal|last=Crockford|first=Catherine|last2=Wittig|first2=Roman M.|last3=Mundry|first3=Roger|last4=Zuberbühler|first4=Klaus|date=2012-01|title=Wild Chimpanzees Inform Ignorant Group Members of Danger|url=http://dx.doi.org/10.1016/j.cub.2011.11.053|journal=Current Biology|volume=22|issue=2|pages=142–146|doi=10.1016/j.cub.2011.11.053|issn=0960-9822}}</ref> Interestingly, 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.<ref name=":2">{{Cite journal|last=Crockford|first=Catherine|last2=Wittig|first2=Roman M.|last3=Zuberbühler|first3=Klaus|date=2017-11|title=Vocalizing in chimpanzees is influenced by social-cognitive processes|url=http://dx.doi.org/10.1126/sciadv.1701742|journal=Science Advances|volume=3|issue=11|pages=e1701742|doi=10.1126/sciadv.1701742|issn=2375-2548}}</ref> In a recent 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. In response to the non-snake-related calls from receivers, the callers increased their vocal and nonvocal signalling and coupled it with increased receiver monitoring.

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.<ref name=":3">{{Cite journal|last=Dezecache|first=Guillaume|last2=Crockford|first2=Catherine|last3=Zuberbühler|first3=Klaus|date=2019-07-06|title=The development of communication in alarm contexts in wild chimpanzees|url=https://doi.org/10.1007/s00265-019-2716-6|journal=Behavioral Ecology and Sociobiology|language=en|volume=73|issue=8|pages=104|doi=10.1007/s00265-019-2716-6|issn=1432-0762|pmc=PMC6612320|pmid=31346303}}</ref> There are several hypotheses for this lack of alarm calling in infants. Firstly, there may be 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.<ref name=":3" /> 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.<ref name=":3" /> Additionally, infants may lack the physical ability to produce these calls or lack the necessary experience to classify unfamiliar objects as dangerous and warranting an alarm signal.<ref name=":3" /> Therefore, alarm calling may require advanced levels of development, perception, categorization, and social cognition.<ref name=":3" />

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.<ref name=":1" /><ref name=":2" />

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 gaze alternations, from a threat to a nearby conspecific and back to the threat, or as the use of two gaze alternations.<ref name=":3" /> 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.<ref name=":3" /> 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.<ref name=":3" /> It has also been proposed to be a communicative behaviour or simply the result of shifts in attention between different environmental elements.<ref name=":0" /><ref name=":3" />

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.<ref name=":4">{{Cite journal|last=Crockford|first=Catherine|last2=Gruber|first2=Thibaud|last3=Zuberbühler|first3=Klaus|date=2018-05|title=Chimpanzee quiet hoo variants differ according to context|url=http://dx.doi.org/10.1098/rsos.172066|journal=Royal Society Open Science|volume=5|issue=5|pages=172066|doi=10.1098/rsos.172066|issn=2054-5703}}</ref> Context-dependent communication is beneficial because it facilities cooperative activities and social cohesion between signallers and receivers.<ref name=":4" /> Furthermore, alarm calls in chimpanzees 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.<ref name=":0" /><ref name=":2" /> Filling a gap in information and incorporating social cues and intentionality into communication are all components of human language.<ref name=":0" /><ref name=":2" /> These shared elements between chimpanzee and human communication suggest that our last common ancestor with chimpanzees also possessed these linguistic abilities.<ref name=":0" /><ref name=":2" /><ref>{{Citation|last=Hauser|first=Marc D.|title=The faculty of language: what is it, who has it, and how did it evolve?|url=http://dx.doi.org/10.1017/cbo9780511817755.002|work=The Evolution of Human Language|pages=14–42|publisher=Cambridge University Press|isbn=978-0-511-81775-5|access-date=2020-01-31|last2=Chomsky|first2=Noam|last3=Fitch|first3=W. Tecumseh}}</ref>


==False alarm calls==
==False alarm calls==

Revision as of 01:03, 31 January 2020

For devices giving alarm signals, see Alarm device.
"Alarm call" and "Alarmer" redirect here. For the Björk song, see Alarm Call. For the shoulder dystocia mnemonic, see ALARMER.
Alarm calls have been studied in many species, such as Belding's ground squirrels.

In animal communication, an alarm signal is an antipredator adaptation in the form of 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 alarm call of the blackbird is a familiar sound in many gardens. Other animals, like fish and insects, may use non-auditory signals, such as chemical messages. Visual signs such as the white tail flashes of many deer have been suggested as alarm signals; they are less likely to be received by conspecifics, so have tended to be treated as a signal to the predator instead.

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

Main article: Signalling theory
Alarm call by a rock squirrel

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.[5] 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.[citation needed]

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 animals 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]

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.[8][9]

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.[10] Similarly, the cotton-top tamarin is able to use a limited vocal range of alarm calls to distinguish between aerial and land predators.[11] 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.[12][13]

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.[14]

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.[15]

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,[16][17] whose work on this topic has been widely cited and debated.[18][19] 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

Chimpanzees produce alarm calls in response to threats.

Chimpanzees emit alarm calls in response to predators, such as leopards and snakes.[20][21] They produce three types of alarm calls: acoustically-variable 'hoos', 'barks', and 'SOS screams'.[22] Alarm signalling is impacted by receiver knowledge and caller age, can be coupled with receiver monitoring, and is important to the understanding of the evolution of hominoid communication.

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.[23] Interestingly, 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.[24] In a recent 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. In response to the non-snake-related calls from receivers, the callers increased their vocal and nonvocal signalling and coupled it with increased receiver monitoring.

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.[25] There are several hypotheses for this lack of alarm calling in infants. Firstly, there may be 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.[25] 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.[25] Additionally, infants may lack the physical ability to produce these calls or lack the necessary experience to classify unfamiliar objects as dangerous and warranting an alarm signal.[25] Therefore, alarm calling may require advanced levels of development, perception, categorization, and social cognition.[25]

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.[23][24]

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 gaze alternations, from a threat to a nearby conspecific and back to the threat, or as the use of two gaze alternations.[25] 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.[25] 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.[25] It has also been proposed to be a communicative behaviour or simply the result of shifts in attention between different environmental elements.[22][25]

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.[26] Context-dependent communication is beneficial because it facilities cooperative activities and social cohesion between signallers and receivers.[26] Furthermore, alarm calls in chimpanzees 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.[22][24] Filling a gap in information and incorporating social cues and intentionality into communication are all components of human language.[22][24] These shared elements between chimpanzee and human communication suggest that our last common ancestor with chimpanzees also possessed these linguistic abilities.[22][24][27]

False alarm calls

Deceptive vocalizations are given by male barn swallows.

Deceptive alarm calls are used by male swallows (Hirundo rustica).[28] 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.[29]

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.[30]

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.[31]

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.[32] 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.[33]

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.[34]

See also

References

  1. ^ Biology Letters. Titi monkey call sequences vary with predator location and type
  2. ^ Zuberbühler, Klaus; Jenny, David; Bshary, Redouan (1999). "The Predator Deterrence Function of Primate Alarm Calls" (PDF). Ethology. 105 (6): 477–490. doi:10.1046/j.1439-0310.1999.00396.x.
  3. ^ "Archived copy" (PDF). Archived from the original (PDF) on 2011-08-22. Retrieved 2011-03-20.{{cite web}}: CS1 maint: archived copy as title (link)
  4. ^ "The Semantics of Vervet Monkey Alarm Calls: Part I". 2011-03-09.
  5. ^ Cheney, Dorothy L.; Seyfarth, Robert M. (1985). "Vervet Monkey Alarm Calls: Manipulation through Shared Information?". Behaviour. 94 (1/2): 150–166. doi:10.1163/156853985X00316. JSTOR 4534456.
  6. ^ Chivers, D. P.; Brown, G. E.; Smith, R. J. F. (1996). "The Evolution of Chemical Alarm Signals: Attracting Predators Benefits Alarm Signal Senders" (PDF). The American Naturalist. 148 (4): 649–659. doi:10.1086/285945.
  7. ^ 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.
  8. ^ 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. Bibcode:1980Sci...210..801S. doi:10.1126/science.7433999. PMID 7433999.
  9. ^ Cheney, D. L.; Seyfarth, R. M. (1981). "Selective Forces Affecting the Predator Alarm Calls of Vervet Monkeys". Behaviour. 76 (1): 25–61. doi:10.1163/156853981x00022. JSTOR 4534091.
  10. ^ Ouattara, Karim; Lemasson, Alban; Zuberbühler, Klaus (2009). "Campbell's Monkeys Use Affixation to Alter Call Meaning". PLOS ONE. 4 (11): e7808. Bibcode:2009PLoSO...4.7808O. doi:10.1371/journal.pone.0007808. PMC 2771905. PMID 19915663.{{cite journal}}: CS1 maint: unflagged free DOI (link)
  11. ^ 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.
  12. ^ 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.
  13. ^ 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
  14. ^ Zuberbühler, K (2000). "Referential labelling in Diana monkeys". Animal Behaviour. 59 (5): 917–927. doi:10.1006/anbe.1999.1317. PMID 10860519.
  15. ^ 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.
  16. ^ Rendall, D.; Owren, M. J.; Ryan, M. J. (2009). "What do animal signals mean?". Animal Behaviour. 78 (2): 233–240. doi:10.1016/j.anbehav.2009.06.007.
  17. ^ Owren, M. J.; Rendall, D.; Ryan, M. J. (2010). "Redefining animal signaling: Influence versus information in communication". Biology and Philosophy. 25 (5): 755–780. doi:10.1007/s10539-010-9224-4.
  18. ^ Nordell, S. & Valone, T. (2014). Animal Behaviour: Concepts, Methods and Applications. Oxford University Press.
  19. ^ Stegmann, U. E. editor (2013). Animal Communication Theory: Information and Influence. Cambridge University Press, Cambridge, UK. 452 pages
  20. ^ Zuberbühler, Klaus (2000-01). "Causal knowledge of predators' behaviour in wild Diana monkeys". Animal Behaviour. 59 (1): 209–220. doi:10.1006/anbe.1999.1296. ISSN 0003-3472. {{cite journal}}: Check date values in: |date= (help)
  21. ^ Crockford, Catherine; Boesch, Christophe (2003-07). "Context-specific calls in wild chimpanzees, Pan troglodytes verus: analysis of barks". Animal Behaviour. 66 (1): 115–125. doi:10.1006/anbe.2003.2166. ISSN 0003-3472. {{cite journal}}: Check date values in: |date= (help)
  22. ^ a b c d e Schel, Anne Marijke; Townsend, Simon W.; Machanda, Zarin; Zuberbühler, Klaus; Slocombe, Katie E. (2013-10-16). "Chimpanzee Alarm Call Production Meets Key Criteria for Intentionality". PLoS ONE. 8 (10): e76674. doi:10.1371/journal.pone.0076674. ISSN 1932-6203.{{cite journal}}: CS1 maint: unflagged free DOI (link)
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External links

Department of Systematics and Ecology, University of Kansas

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