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        <rdf:li rdf:resource="http://hdl.handle.net/10023/3428" />
        <rdf:li rdf:resource="http://hdl.handle.net/10023/3361" />
        <rdf:li rdf:resource="http://hdl.handle.net/10023/3301" />
        <rdf:li rdf:resource="http://hdl.handle.net/10023/3271" />
        <rdf:li rdf:resource="http://hdl.handle.net/10023/3268" />
        <rdf:li rdf:resource="http://hdl.handle.net/10023/3267" />
        <rdf:li rdf:resource="http://hdl.handle.net/10023/3266" />
        <rdf:li rdf:resource="http://hdl.handle.net/10023/3186" />
        <rdf:li rdf:resource="http://hdl.handle.net/10023/3082" />
        <rdf:li rdf:resource="http://hdl.handle.net/10023/3021" />
        <rdf:li rdf:resource="http://hdl.handle.net/10023/3013" />
        <rdf:li rdf:resource="http://hdl.handle.net/10023/3012" />
        <rdf:li rdf:resource="http://hdl.handle.net/10023/2983" />
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        <rdf:li rdf:resource="http://hdl.handle.net/10023/2555" />
        <rdf:li rdf:resource="http://hdl.handle.net/10023/2446" />
        <rdf:li rdf:resource="http://hdl.handle.net/10023/2003" />
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    <dc:date>2013-04-18T10:01:52Z</dc:date>
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  <item rdf:about="http://hdl.handle.net/10023/3428">
    <title>You are what you eat : Within-subject increases in fruit and vegetable consumption confer beneficial skin-color changes</title>
    <link>http://hdl.handle.net/10023/3428</link>
    <description>Abstract: Background: Fruit and vegetable consumption and ingestion of carotenoids have been found to be associated with human skin-color (yellowness) in a recent cross-sectional study. This carotenoid-based coloration contributes beneficially to the appearance of health in humans and is held to be a sexually selected cue of condition in other species. Methodology and Principal Findings: Here we investigate the effects of fruit and vegetable consumption on skin-color longitudinally to determine the magnitude and duration of diet change required to change skin-color perceptibly. Diet and skin-color were recorded at baseline and after three and six weeks, in a group of 35 individuals who were without makeup, self-tanning agents and/or recent intensive UV exposure. Six-week changes in fruit and vegetable consumption were significantly correlated with changes in skin redness and yellowness over this period, and diet-linked skin reflectance changes were significantly associated with the spectral absorption of carotenoids and not melanin. We also used psychophysical methods to investigate the minimum color change required to confer perceptibly healthier and more attractive skin-coloration. Modest dietary changes are required to enhance apparent health (2.91 portions per day) and attractiveness (3.30 portions). Conclusions: Increased fruit and vegetable consumption confers measurable and perceptibly beneficial effects on Caucasian skin appearance within six weeks. This effect could potentially be used as a motivational tool in dietary intervention.
Description: R Whitehead was funded by an ESRC Studentship.</description>
    <dc:date>2012-03-07T00:00:00Z</dc:date>
    <dc:creator>Whitehead, Ross</dc:creator>
    <dc:creator>Re, Daniel</dc:creator>
    <dc:creator>Xiao, Dengke</dc:creator>
    <dc:creator>Ozakinci, Gozde</dc:creator>
    <dc:creator>Perrett, David Ian</dc:creator>
    <dc:description>Background: Fruit and vegetable consumption and ingestion of carotenoids have been found to be associated with human skin-color (yellowness) in a recent cross-sectional study. This carotenoid-based coloration contributes beneficially to the appearance of health in humans and is held to be a sexually selected cue of condition in other species. Methodology and Principal Findings: Here we investigate the effects of fruit and vegetable consumption on skin-color longitudinally to determine the magnitude and duration of diet change required to change skin-color perceptibly. Diet and skin-color were recorded at baseline and after three and six weeks, in a group of 35 individuals who were without makeup, self-tanning agents and/or recent intensive UV exposure. Six-week changes in fruit and vegetable consumption were significantly correlated with changes in skin redness and yellowness over this period, and diet-linked skin reflectance changes were significantly associated with the spectral absorption of carotenoids and not melanin. We also used psychophysical methods to investigate the minimum color change required to confer perceptibly healthier and more attractive skin-coloration. Modest dietary changes are required to enhance apparent health (2.91 portions per day) and attractiveness (3.30 portions). Conclusions: Increased fruit and vegetable consumption confers measurable and perceptibly beneficial effects on Caucasian skin appearance within six weeks. This effect could potentially be used as a motivational tool in dietary intervention.</dc:description>
  </item>
  <item rdf:about="http://hdl.handle.net/10023/3361">
    <title>Vocal copying of individually distinctive signature whistles in bottlenose dolphins</title>
    <link>http://hdl.handle.net/10023/3361</link>
    <description>Abstract: Vocal learning is relatively common in birds but less so in mammals. Sexual selection and individual or group recognition have been identified as major forces in its evolution. While important in the development of vocal displays, vocal learning also allows signal copying in social interactions. Such copying can function in addressing or labelling selected conspecifics. Most examples of addressing in non-humans come from bird song, where matching occurs in an aggressive context. However, in other animals, addressing with learned signals is very much an affiliative signal. We studied the function of vocal copying in a mammal that shows vocal learning as well as complex cognitive and social behaviour, the bottlenose dolphin (Tursiops truncatus). Copying occurred almost exclusively between close associates such as mother–calf pairs and male alliances during separation and was not followed by aggression. All copies were clearly recognizable as such because copiers consistently modified some acoustic parameters of a signal when copying it. We found no evidence for the use of copying in aggression or deception. This use of vocal copying is similar to its use in human language, where the maintenance of social bonds appears to be more important than the immediate defence of resources.</description>
    <dc:date>2013-04-22T00:00:00Z</dc:date>
    <dc:creator>King, Stephanie Laura</dc:creator>
    <dc:creator>Sayigh, Laela</dc:creator>
    <dc:creator>Wells, Randall</dc:creator>
    <dc:creator>Fellner, Wendi</dc:creator>
    <dc:creator>Janik, Vincent M.</dc:creator>
    <dc:description>Vocal learning is relatively common in birds but less so in mammals. Sexual selection and individual or group recognition have been identified as major forces in its evolution. While important in the development of vocal displays, vocal learning also allows signal copying in social interactions. Such copying can function in addressing or labelling selected conspecifics. Most examples of addressing in non-humans come from bird song, where matching occurs in an aggressive context. However, in other animals, addressing with learned signals is very much an affiliative signal. We studied the function of vocal copying in a mammal that shows vocal learning as well as complex cognitive and social behaviour, the bottlenose dolphin (Tursiops truncatus). Copying occurred almost exclusively between close associates such as mother–calf pairs and male alliances during separation and was not followed by aggression. All copies were clearly recognizable as such because copiers consistently modified some acoustic parameters of a signal when copying it. We found no evidence for the use of copying in aggression or deception. This use of vocal copying is similar to its use in human language, where the maintenance of social bonds appears to be more important than the immediate defence of resources.</dc:description>
  </item>
  <item rdf:about="http://hdl.handle.net/10023/3301">
    <title>Estimated tissue and blood N2 levels and risk of in vivo bubble formation in deep-, intermediate- and shallow diving toothed whales during exposure to naval sonar.</title>
    <link>http://hdl.handle.net/10023/3301</link>
    <description>Abstract: Naval sonar has been accused of causing whale stranding by a mechanism which increases formation of tissue N2 gas bubbles. Increased tissue and blood N2 levels, and thereby increased risk of decompression sickness (DCS), is thought to result from changes in behavior or physiological responses during diving. Previous theoretical studies have used hypothetical sonar-induced changes in both behavior and physiology to model blood and tissue N2 tension PN2, but this is the first attempt to estimate the changes during actual behavioral responses to sonar. We used an existing mathematical model to estimate blood and tissue N2 tension PN2 from dive data recorded from sperm, killer, long-finned pilot, Blainville’s beaked, and Cuvier’s beaked whales before and during exposure to Low- (1–2 kHz) and Mid- (2–7 kHz) frequency active sonar. Our objectives were: (1) to determine if differences in dive behavior affects risk of bubble formation, and if (2) behavioral- or (3) physiological responses to sonar are plausible risk factors. Our results suggest that all species have natural high N2 levels, with deep diving generally resulting in higher end-dive PN2 as compared with shallow diving. Sonar exposure caused some changes in dive behavior in both killer whales, pilot whales and beaked whales, but this did not lead to any increased risk of DCS. However, in three of eight exposure session with sperm whales, the animal changed to shallower diving, and in all these cases this seem to result in an increased risk of DCS, although risk was still within the normal risk range of this species. When a hypothetical removal of the normal dive response (bradycardia and peripheral vasoconstriction), was added to the behavioral response during model simulations, this led to an increased variance in the estimated end-dive N2 levels, but no consistent change of risk. In conclusion, we cannot rule out the possibility that a combination of behavioral and physiological responses to sonar have the potential to alter the blood and tissue end-dive N2 tension to levels which could cause DCS and formation of in vivo bubbles, but the actually observed behavioral responses of cetaceans to sonar in our study, do not imply any significantly increased risk of DCS.</description>
    <dc:date>2012-05-10T00:00:00Z</dc:date>
    <dc:creator>Kvadsheim, Petter H</dc:creator>
    <dc:creator>Miller, Patrick</dc:creator>
    <dc:creator>Tyack, Peter Lloyd</dc:creator>
    <dc:creator>Sivle, Lise D</dc:creator>
    <dc:creator>Lam, Frans Peter</dc:creator>
    <dc:creator>Fahlman, Andreas</dc:creator>
    <dc:description>Naval sonar has been accused of causing whale stranding by a mechanism which increases formation of tissue N2 gas bubbles. Increased tissue and blood N2 levels, and thereby increased risk of decompression sickness (DCS), is thought to result from changes in behavior or physiological responses during diving. Previous theoretical studies have used hypothetical sonar-induced changes in both behavior and physiology to model blood and tissue N2 tension PN2, but this is the first attempt to estimate the changes during actual behavioral responses to sonar. We used an existing mathematical model to estimate blood and tissue N2 tension PN2 from dive data recorded from sperm, killer, long-finned pilot, Blainville’s beaked, and Cuvier’s beaked whales before and during exposure to Low- (1–2 kHz) and Mid- (2–7 kHz) frequency active sonar. Our objectives were: (1) to determine if differences in dive behavior affects risk of bubble formation, and if (2) behavioral- or (3) physiological responses to sonar are plausible risk factors. Our results suggest that all species have natural high N2 levels, with deep diving generally resulting in higher end-dive PN2 as compared with shallow diving. Sonar exposure caused some changes in dive behavior in both killer whales, pilot whales and beaked whales, but this did not lead to any increased risk of DCS. However, in three of eight exposure session with sperm whales, the animal changed to shallower diving, and in all these cases this seem to result in an increased risk of DCS, although risk was still within the normal risk range of this species. When a hypothetical removal of the normal dive response (bradycardia and peripheral vasoconstriction), was added to the behavioral response during model simulations, this led to an increased variance in the estimated end-dive N2 levels, but no consistent change of risk. In conclusion, we cannot rule out the possibility that a combination of behavioral and physiological responses to sonar have the potential to alter the blood and tissue end-dive N2 tension to levels which could cause DCS and formation of in vivo bubbles, but the actually observed behavioral responses of cetaceans to sonar in our study, do not imply any significantly increased risk of DCS.</dc:description>
  </item>
  <item rdf:about="http://hdl.handle.net/10023/3271">
    <title>African perceptions of female attractiveness</title>
    <link>http://hdl.handle.net/10023/3271</link>
    <description>Abstract: Little is known about mate choice preferences outside Western, educated, industrialised, rich and democratic societies, even though these Western populations may be particularly unrepresentative of human populations. To our knowledge, this is the first study to test which facial cues contribute to African perceptions of African female attractiveness and also the first study to test the combined role of facial adiposity, skin colour (lightness, yellowness and redness), skin homogeneity and youthfulness in the facial attractiveness preferences of any population. Results show that youthfulness, skin colour, skin homogeneity and facial adiposity significantly and independently predict attractiveness in female African faces. Younger, thinner women with a lighter, yellower skin colour and a more homogenous skin tone are considered more attractive. These findings provide a more global perspective on human mate choice and point to a universal role for these four facial cues in female facial attractiveness.</description>
    <dc:date>2012-10-29T00:00:00Z</dc:date>
    <dc:creator>Coetzee, Vinet</dc:creator>
    <dc:creator>Faerber, Stella J</dc:creator>
    <dc:creator>Greeff, Jaco M</dc:creator>
    <dc:creator>Lefevre, Carmen Emilia</dc:creator>
    <dc:creator>Re, Daniel</dc:creator>
    <dc:creator>Perrett, David Ian</dc:creator>
    <dc:description>Little is known about mate choice preferences outside Western, educated, industrialised, rich and democratic societies, even though these Western populations may be particularly unrepresentative of human populations. To our knowledge, this is the first study to test which facial cues contribute to African perceptions of African female attractiveness and also the first study to test the combined role of facial adiposity, skin colour (lightness, yellowness and redness), skin homogeneity and youthfulness in the facial attractiveness preferences of any population. Results show that youthfulness, skin colour, skin homogeneity and facial adiposity significantly and independently predict attractiveness in female African faces. Younger, thinner women with a lighter, yellower skin colour and a more homogenous skin tone are considered more attractive. These findings provide a more global perspective on human mate choice and point to a universal role for these four facial cues in female facial attractiveness.</dc:description>
  </item>
  <item rdf:about="http://hdl.handle.net/10023/3268">
    <title>Oxygenated-blood colour change thresholds for perceived facial redness, health, and attractiveness</title>
    <link>http://hdl.handle.net/10023/3268</link>
    <description>Abstract: Blood oxygenation level is associated with cardiovascular fitness, and raising oxygenated blood colouration in human faces increases perceived health. The current study used a two-alternative forced choice (2AFC) psychophysics design to quantify the oxygenated blood colour (redness) change threshold required to affect perception of facial colour, health and attractiveness. Detection thresholds for colour judgments were lower than those for health and attractiveness, which did not differ. The results suggest redness preferences do not reflect a sensory bias, rather preferences may be based on accurate indications of health status. Furthermore, results suggest perceived health and attractiveness may be perceptually equivalent when they are assessed based on facial redness. Appearance-based motivation for lifestyle change can be effective; thus future studies could assess the degree to which cardiovascular fitness increases face redness and could quantify changes in aerobic exercise needed to increase facial attractiveness.</description>
    <dc:date>2011-03-23T00:00:00Z</dc:date>
    <dc:creator>Re, Daniel E.</dc:creator>
    <dc:creator>Whitehead, Ross D.</dc:creator>
    <dc:creator>Xiao, Dengke</dc:creator>
    <dc:creator>Perrett, David I.</dc:creator>
    <dc:description>Blood oxygenation level is associated with cardiovascular fitness, and raising oxygenated blood colouration in human faces increases perceived health. The current study used a two-alternative forced choice (2AFC) psychophysics design to quantify the oxygenated blood colour (redness) change threshold required to affect perception of facial colour, health and attractiveness. Detection thresholds for colour judgments were lower than those for health and attractiveness, which did not differ. The results suggest redness preferences do not reflect a sensory bias, rather preferences may be based on accurate indications of health status. Furthermore, results suggest perceived health and attractiveness may be perceptually equivalent when they are assessed based on facial redness. Appearance-based motivation for lifestyle change can be effective; thus future studies could assess the degree to which cardiovascular fitness increases face redness and could quantify changes in aerobic exercise needed to increase facial attractiveness.</dc:description>
  </item>
  <item rdf:about="http://hdl.handle.net/10023/3267">
    <title>Repeated elicitation of the acoustic startle reflex leads to sensitisation in subsequent avoidance behaviour and induces fear conditioning</title>
    <link>http://hdl.handle.net/10023/3267</link>
    <description>Abstract: Background: Autonomous reflexes enable animals to respond quickly to potential threats, prevent injury and mediate fight or flight responses. Intense acoustic stimuli with sudden onsets elicit a startle reflex while stimuli of similar intensity but with longer rise times only cause a cardiac defence response. In laboratory settings, habituation appears to affect all of these reflexes so that the response amplitude generally decreases with repeated exposure to the stimulus. The startle reflex has become a model system for the study of the neural basis of simple learning processes and emotional processing and is often used as a diagnostic tool in medical applications. However, previous studies did not allow animals to avoid the stimulus and the evolutionary function and long-term behavioural consequences of repeated startling remain speculative. In this study we investigate the follow-up behaviour associated with the startle reflex in wild-captured animals using an experimental setup that allows individuals to exhibit avoidance behaviour. Results: We present evidence that repeated elicitation of the acoustic startle reflex leads to rapid and pronounced sensitisation of sustained spatial avoidance behaviour in grey seals (Halichoerus grypus). Animals developed rapid flight responses, left the exposure pool and showed clear signs of fear conditioning. Once sensitised, seals even avoided a known food source that was close to the sound source. In contrast, animals exposed to non-startling (long rise time) stimuli of the same maximum sound pressure habituated and flight responses waned or were absent from the beginning. The startle threshold of grey seals expressed in units of sensation levels was comparable to thresholds reported for other mammals (93 dB). Conclusions: Our results demonstrate that the acoustic startle reflex plays a crucial role in mediating flight responses and strongly influences the motivational state of an animal beyond a short-term muscular response by mediating long-term avoidance. The reflex is therefore not only a measure of emotional state but also influences emotional processing. The biological function of the startle reflex is most likely associated with mediating rapid flight responses. The data indicate that repeated startling by anthropogenic noise sources might have severe effects on long-term behaviour. Future, studies are needed to investigate whether such effects can be associated with reduced individual fitness or even longevity of individuals.</description>
    <dc:date>2011-01-01T00:00:00Z</dc:date>
    <dc:creator>Gotz, Thomas</dc:creator>
    <dc:creator>Janik, Vincent M.</dc:creator>
    <dc:description>Background: Autonomous reflexes enable animals to respond quickly to potential threats, prevent injury and mediate fight or flight responses. Intense acoustic stimuli with sudden onsets elicit a startle reflex while stimuli of similar intensity but with longer rise times only cause a cardiac defence response. In laboratory settings, habituation appears to affect all of these reflexes so that the response amplitude generally decreases with repeated exposure to the stimulus. The startle reflex has become a model system for the study of the neural basis of simple learning processes and emotional processing and is often used as a diagnostic tool in medical applications. However, previous studies did not allow animals to avoid the stimulus and the evolutionary function and long-term behavioural consequences of repeated startling remain speculative. In this study we investigate the follow-up behaviour associated with the startle reflex in wild-captured animals using an experimental setup that allows individuals to exhibit avoidance behaviour. Results: We present evidence that repeated elicitation of the acoustic startle reflex leads to rapid and pronounced sensitisation of sustained spatial avoidance behaviour in grey seals (Halichoerus grypus). Animals developed rapid flight responses, left the exposure pool and showed clear signs of fear conditioning. Once sensitised, seals even avoided a known food source that was close to the sound source. In contrast, animals exposed to non-startling (long rise time) stimuli of the same maximum sound pressure habituated and flight responses waned or were absent from the beginning. The startle threshold of grey seals expressed in units of sensation levels was comparable to thresholds reported for other mammals (93 dB). Conclusions: Our results demonstrate that the acoustic startle reflex plays a crucial role in mediating flight responses and strongly influences the motivational state of an animal beyond a short-term muscular response by mediating long-term avoidance. The reflex is therefore not only a measure of emotional state but also influences emotional processing. The biological function of the startle reflex is most likely associated with mediating rapid flight responses. The data indicate that repeated startling by anthropogenic noise sources might have severe effects on long-term behaviour. Future, studies are needed to investigate whether such effects can be associated with reduced individual fitness or even longevity of individuals.</dc:description>
  </item>
  <item rdf:about="http://hdl.handle.net/10023/3266">
    <title>Redness enhances perceived aggression, dominance and attractiveness in men's faces</title>
    <link>http://hdl.handle.net/10023/3266</link>
    <description>Abstract: In a range of non-human primate, bird and fish species, the intensity of red coloration in males is associated with social dominance, testosterone levels and mate selection. In humans too, skin redness is associated with health, but it is not known whether - as in non-human species - it is also associated with dominance and links to attractiveness have not been thoroughly investigated. Here we allow female participants to manipulate the CIELab a* value (red-green axis) of skin to maximize the perceived aggression, dominance and attractiveness of photographs of men's faces, and make two findings. First, participants increased a* (increasing redness) to enhance each attribute, suggesting that facial redness is perceived as conveying similar information about a male's qualities in humans as it does in non-human species. Second, there were significant differences between trial types: the highest levels of red were associated with aggression, an intermediate level with dominance, and the least with attractiveness. These differences may reflect a trade-off between the benefits of selecting a healthy, dominant partner and the negative consequences of aggression.</description>
    <dc:date>2012-01-01T00:00:00Z</dc:date>
    <dc:creator>Stephen, Ian D.</dc:creator>
    <dc:creator>Oldham, Francesca H.</dc:creator>
    <dc:creator>Perrett, David I.</dc:creator>
    <dc:creator>Barton, Robert A.</dc:creator>
    <dc:description>In a range of non-human primate, bird and fish species, the intensity of red coloration in males is associated with social dominance, testosterone levels and mate selection. In humans too, skin redness is associated with health, but it is not known whether - as in non-human species - it is also associated with dominance and links to attractiveness have not been thoroughly investigated. Here we allow female participants to manipulate the CIELab a* value (red-green axis) of skin to maximize the perceived aggression, dominance and attractiveness of photographs of men's faces, and make two findings. First, participants increased a* (increasing redness) to enhance each attribute, suggesting that facial redness is perceived as conveying similar information about a male's qualities in humans as it does in non-human species. Second, there were significant differences between trial types: the highest levels of red were associated with aggression, an intermediate level with dominance, and the least with attractiveness. These differences may reflect a trade-off between the benefits of selecting a healthy, dominant partner and the negative consequences of aggression.</dc:description>
  </item>
  <item rdf:about="http://hdl.handle.net/10023/3186">
    <title>Spontaneous emergence, imitation and spread of alternative foraging techniques among groups of vervet monkeys</title>
    <link>http://hdl.handle.net/10023/3186</link>
    <description>Abstract: Animal social learning has become a subject of broad interest, but demonstrations of bodily imitation in animals remain rare. Based on Voelkl and Huber's study of imitation by marmosets, we tested four groups of semi-captive vervet monkeys presented with food in modified film canisters (“aethipops’). One individual was trained to take the tops off canisters in each group and demonstrated five openings to them. In three groups these models used their mouth to remove the lid, but in one of the groups the model also spontaneously pulled ropes on a canister to open it. In the last group the model preferred to remove the lid with her hands. Following these spontaneous differentiations of foraging techniques in the models, we observed the techniques used by the other group members to open the canisters. We found that mouth opening was the most common technique overall, but the rope and hands methods were used significantly more in groups they were demonstrated in than in groups where they were not. Our results show bodily matching that is conventionally described as imitation. We discuss the relevance of these findings to discoveries about mirror neurons, and implications of the identity of the model for social transmission.</description>
    <dc:date>2012-10-10T00:00:00Z</dc:date>
    <dc:creator>van de Waal, Erica</dc:creator>
    <dc:creator>Whiten, Andrew</dc:creator>
    <dc:description>Animal social learning has become a subject of broad interest, but demonstrations of bodily imitation in animals remain rare. Based on Voelkl and Huber's study of imitation by marmosets, we tested four groups of semi-captive vervet monkeys presented with food in modified film canisters (“aethipops’). One individual was trained to take the tops off canisters in each group and demonstrated five openings to them. In three groups these models used their mouth to remove the lid, but in one of the groups the model also spontaneously pulled ropes on a canister to open it. In the last group the model preferred to remove the lid with her hands. Following these spontaneous differentiations of foraging techniques in the models, we observed the techniques used by the other group members to open the canisters. We found that mouth opening was the most common technique overall, but the rope and hands methods were used significantly more in groups they were demonstrated in than in groups where they were not. Our results show bodily matching that is conventionally described as imitation. We discuss the relevance of these findings to discoveries about mirror neurons, and implications of the identity of the model for social transmission.</dc:description>
  </item>
  <item rdf:about="http://hdl.handle.net/10023/3082">
    <title>Identification of learning mechanisms in a wild meerkat population</title>
    <link>http://hdl.handle.net/10023/3082</link>
    <description>Abstract: Vigorous debates as to the evolutionary origins of culture remain unresolved due to an absence of methods for identifying learning mechanisms in natural populations. While laboratory experiments on captive animals have revealed evidence for a number of mechanisms, these may not necessarily reflect the processes typically operating in nature. We developed a novel method that allows social and asocial learning mechanisms to be determined in animal groups from the patterns of interaction with, and solving of, a task. We deployed it to analyse learning in groups of wild meerkats (Suricata suricatta) presented with a novel foraging apparatus. We identify nine separate learning processes underlying the meerkats' foraging behaviour, in each case precisely quantifying their strength and duration, including local enhancement, emulation, and a hitherto unrecognized form of social learning, which we term `observational perseverance'. Our analysis suggests a key factor underlying the stability of behavioural traditions is a high ratio of specific to generalized social learning effects. The approach has widespread potential as an ecologically valid tool to investigate learning mechanisms in natural groups of animals, including humans.</description>
    <dc:date>2012-08-08T00:00:00Z</dc:date>
    <dc:creator>Hoppitt, William John Edward</dc:creator>
    <dc:creator>Samson, Jamie</dc:creator>
    <dc:creator>Laland, Kevin N.</dc:creator>
    <dc:creator>Thornton, Alex</dc:creator>
    <dc:description>Vigorous debates as to the evolutionary origins of culture remain unresolved due to an absence of methods for identifying learning mechanisms in natural populations. While laboratory experiments on captive animals have revealed evidence for a number of mechanisms, these may not necessarily reflect the processes typically operating in nature. We developed a novel method that allows social and asocial learning mechanisms to be determined in animal groups from the patterns of interaction with, and solving of, a task. We deployed it to analyse learning in groups of wild meerkats (Suricata suricatta) presented with a novel foraging apparatus. We identify nine separate learning processes underlying the meerkats' foraging behaviour, in each case precisely quantifying their strength and duration, including local enhancement, emulation, and a hitherto unrecognized form of social learning, which we term `observational perseverance'. Our analysis suggests a key factor underlying the stability of behavioural traditions is a high ratio of specific to generalized social learning effects. The approach has widespread potential as an ecologically valid tool to investigate learning mechanisms in natural groups of animals, including humans.</dc:description>
  </item>
  <item rdf:about="http://hdl.handle.net/10023/3021">
    <title>Social learning of vocal structure in a nonhuman primate?</title>
    <link>http://hdl.handle.net/10023/3021</link>
    <description>Abstract: Background: Non-human primate communication is thought to be fundamentally different from human speech, mainly due to vast differences in vocal control. The lack of these abilities in non-human primates is especially striking if compared to some marine mammals and bird species, which has generated somewhat of an evolutionary conundrum. What are the biological roots and underlying evolutionary pressures of the human ability to voluntarily control sound production and learn the vocal utterances of others? One hypothesis is that this capacity has evolved gradually in humans from an ancestral stage that resembled the vocal behavior of modern primates. Support for this has come from studies that have documented limited vocal flexibility and convergence in different primate species, typically in calls used during social interactions. The mechanisms underlying these patterns, however, are currently unknown. Specifically, it has been difficult to rule out explanations based on genetic relatedness, suggesting that such vocal flexibility may not be the result of social learning. Results: To address this point, we compared the degree of acoustic similarity of contact calls in free-ranging Campbell's monkeys as a function of their social bonds and genetic relatedness. We calculated three different indices to compare the similarities between the calls' frequency contours, the duration of grooming interactions and the microsatellite-based genetic relatedness between partners. We found a significantly positive relation between bond strength and acoustic similarity that was independent of genetic relatedness. Conclusion: Genetic factors determine the general species-specific call repertoire of a primate species, while social factors can influence the fine structure of some the call types. The finding is in line with the more general hypothesis that human speech has evolved gradually from earlier primate-like vocal communication.</description>
    <dc:date>2011-12-16T00:00:00Z</dc:date>
    <dc:creator>Lemasson, Alban</dc:creator>
    <dc:creator>Ouattara, Karim</dc:creator>
    <dc:creator>Petit, Eric J.</dc:creator>
    <dc:creator>Zuberbuhler, Klaus</dc:creator>
    <dc:description>Background: Non-human primate communication is thought to be fundamentally different from human speech, mainly due to vast differences in vocal control. The lack of these abilities in non-human primates is especially striking if compared to some marine mammals and bird species, which has generated somewhat of an evolutionary conundrum. What are the biological roots and underlying evolutionary pressures of the human ability to voluntarily control sound production and learn the vocal utterances of others? One hypothesis is that this capacity has evolved gradually in humans from an ancestral stage that resembled the vocal behavior of modern primates. Support for this has come from studies that have documented limited vocal flexibility and convergence in different primate species, typically in calls used during social interactions. The mechanisms underlying these patterns, however, are currently unknown. Specifically, it has been difficult to rule out explanations based on genetic relatedness, suggesting that such vocal flexibility may not be the result of social learning. Results: To address this point, we compared the degree of acoustic similarity of contact calls in free-ranging Campbell's monkeys as a function of their social bonds and genetic relatedness. We calculated three different indices to compare the similarities between the calls' frequency contours, the duration of grooming interactions and the microsatellite-based genetic relatedness between partners. We found a significantly positive relation between bond strength and acoustic similarity that was independent of genetic relatedness. Conclusion: Genetic factors determine the general species-specific call repertoire of a primate species, while social factors can influence the fine structure of some the call types. The finding is in line with the more general hypothesis that human speech has evolved gradually from earlier primate-like vocal communication.</dc:description>
  </item>
  <item rdf:about="http://hdl.handle.net/10023/3013">
    <title>Community-specific evaluation of tool affordances in wild chimpanzees</title>
    <link>http://hdl.handle.net/10023/3013</link>
    <description>Abstract: The notion of animal culture, defined as socially transmitted community-specific behaviour patterns, remains controversial, notably because the definition relies on surface behaviours without addressing underlying cognitive processes. In contrast, human cultures are the product of socially acquired ideas that shape how individuals interact with their environment. We conducted field experiments with two culturally distinct chimpanzee communities in Uganda, which revealed significant differences in how individuals considered the affording parts of an experimentally provided tool to extract honey from a standardised cavity. Firstly, individuals of the two communities found different functional parts of the tool salient, suggesting that they experienced a cultural bias in their cognition. Secondly, when the alternative function was made more salient, chimpanzees were unable to learn it, suggesting that prior cultural background can interfere with new learning. Culture appears to shape how chimpanzees see the world, suggesting that a cognitive component underlies the observed behavioural patterns.</description>
    <dc:date>2011-11-03T00:00:00Z</dc:date>
    <dc:creator>Gruber, Thibaud</dc:creator>
    <dc:creator>Muller, Martin N.</dc:creator>
    <dc:creator>Reynolds, Vernon</dc:creator>
    <dc:creator>Wrangham, Richard</dc:creator>
    <dc:creator>Zuberbuehler, Klaus</dc:creator>
    <dc:description>The notion of animal culture, defined as socially transmitted community-specific behaviour patterns, remains controversial, notably because the definition relies on surface behaviours without addressing underlying cognitive processes. In contrast, human cultures are the product of socially acquired ideas that shape how individuals interact with their environment. We conducted field experiments with two culturally distinct chimpanzee communities in Uganda, which revealed significant differences in how individuals considered the affording parts of an experimentally provided tool to extract honey from a standardised cavity. Firstly, individuals of the two communities found different functional parts of the tool salient, suggesting that they experienced a cultural bias in their cognition. Secondly, when the alternative function was made more salient, chimpanzees were unable to learn it, suggesting that prior cultural background can interfere with new learning. Culture appears to shape how chimpanzees see the world, suggesting that a cognitive component underlies the observed behavioural patterns.</dc:description>
  </item>
  <item rdf:about="http://hdl.handle.net/10023/3012">
    <title>Communication during sex among female bonobos : effects of dominance, solicitation and audience</title>
    <link>http://hdl.handle.net/10023/3012</link>
    <description>Abstract: Bonobo females frequently form close bonds, which give them social power over other group members. One potential mechanism to facilitate female bonding is the performance of sexual interactions. Using naturalistic observations and experiments, we found various patterns that determined female-female sexual interactions. First, while low-ranked females interacted with all females, sexual interactions between high-ranked females were rare. Second, during genital contacts, females sometimes produced 'copulation calls', which were significantly affected by the rank of the caller and partner, as well as the solicitation direction. Third, there was a significant effect of the alpha female as a bystander, while variables relating to physical experience had no effects. Overall, results highlight the importance of sexual interactions for bonobo female social relations. Copulation calls are an important tool during this process, suggesting that they have become ritualised, beyond their reproductive function, to serve as broader social signals in flexible and potentially strategic ways.</description>
    <dc:date>2012-03-01T00:00:00Z</dc:date>
    <dc:creator>Clay, Zanna</dc:creator>
    <dc:creator>Zuberbuehler, Klaus</dc:creator>
    <dc:description>Bonobo females frequently form close bonds, which give them social power over other group members. One potential mechanism to facilitate female bonding is the performance of sexual interactions. Using naturalistic observations and experiments, we found various patterns that determined female-female sexual interactions. First, while low-ranked females interacted with all females, sexual interactions between high-ranked females were rare. Second, during genital contacts, females sometimes produced 'copulation calls', which were significantly affected by the rank of the caller and partner, as well as the solicitation direction. Third, there was a significant effect of the alpha female as a bystander, while variables relating to physical experience had no effects. Overall, results highlight the importance of sexual interactions for bonobo female social relations. Copulation calls are an important tool during this process, suggesting that they have become ritualised, beyond their reproductive function, to serve as broader social signals in flexible and potentially strategic ways.</dc:description>
  </item>
  <item rdf:about="http://hdl.handle.net/10023/2983">
    <title>Production and perception rules underlying visual patterns : effects of symmetry and hierarchy</title>
    <link>http://hdl.handle.net/10023/2983</link>
    <description>Abstract: Formal language theory has been extended to two-dimensional patterns, but little is known about two-dimensional pattern perception. We first examined spontaneous two-dimensional visual pattern production by humans, gathered using a novel touch screen approach. Both spontaneous creative production and subsequent aesthetic ratings show that humans prefer ordered, symmetrical patterns over random patterns. We then further explored pattern-parsing abilities in different human groups, and compared them with pigeons. We generated visual plane patterns based on rules varying in complexity. All human groups tested, including children and individuals diagnosed with autism spectrum disorder (ASD), were able to detect violations of all production rules tested. Our ASD participants detected pattern violations with the same speed and accuracy as matched controls. Children's ability to detect violations of a relatively complex rotational rule correlated with age, whereas their ability to detect violations of a simple translational rule did not. By contrast, even with extensive training, pigeons were unable to detect orientation-based structural violations, suggesting that, unlike humans, they did not learn the underlying structural rules. Visual two-dimensional patterns offer a promising new formally-grounded way to investigate pattern production and perception in general, widely applicable across species and age groups.</description>
    <dc:date>2012-07-19T00:00:00Z</dc:date>
    <dc:creator>Westphal, Gesche</dc:creator>
    <dc:creator>Huber, Ludwig</dc:creator>
    <dc:creator>Gomez, Juan-Carlos</dc:creator>
    <dc:creator>Fitch, William Tecumseh Sherman</dc:creator>
    <dc:description>Formal language theory has been extended to two-dimensional patterns, but little is known about two-dimensional pattern perception. We first examined spontaneous two-dimensional visual pattern production by humans, gathered using a novel touch screen approach. Both spontaneous creative production and subsequent aesthetic ratings show that humans prefer ordered, symmetrical patterns over random patterns. We then further explored pattern-parsing abilities in different human groups, and compared them with pigeons. We generated visual plane patterns based on rules varying in complexity. All human groups tested, including children and individuals diagnosed with autism spectrum disorder (ASD), were able to detect violations of all production rules tested. Our ASD participants detected pattern violations with the same speed and accuracy as matched controls. Children's ability to detect violations of a relatively complex rotational rule correlated with age, whereas their ability to detect violations of a simple translational rule did not. By contrast, even with extensive training, pigeons were unable to detect orientation-based structural violations, suggesting that, unlike humans, they did not learn the underlying structural rules. Visual two-dimensional patterns offer a promising new formally-grounded way to investigate pattern production and perception in general, widely applicable across species and age groups.</dc:description>
  </item>
  <item rdf:about="http://hdl.handle.net/10023/2560">
    <title>Assessing public engagement with science in a university primate research centre in a national zoo</title>
    <link>http://hdl.handle.net/10023/2560</link>
    <description>Abstract: Recent years have seen increasing encouragement by research institutions and funding bodies for scientists to actively engage with the public, who ultimately finance their work. Animal behaviour as a discipline possesses several features, including its inherent accessibility and appeal to the public, that may help it occupy a particularly successful niche within these developments. It has also established a repertoire of quantitative behavioural methodologies that can be used to document the public's responses to engagement initiatives. This kind of assessment is becoming increasingly important considering the enormous effort now being put into public engagement projects, whose effects are more often assumed than demonstrated. Here we report our first attempts to quantify relevant aspects of the behaviour of a sample of the hundreds of thousands of visitors who pass through the ‘Living Links to Human Evolution Research Centre’ in Edinburgh Zoo. This University research centre actively encourages the public to view ongoing primate research and associated science engagement activities. Focal follows of visitors and scan sampling showed substantial ‘dwell times’ in the Centre by common zoo standards and the addition of new engagement elements in a second year was accompanied by significantly increased overall dwell times, tripling for the most committed two thirds of visitors. Larger groups of visitors were found to spend more time in the Centre than smaller ones. Viewing live, active science was the most effective activity, shown to be enhanced by novel presentations of carefully constructed explanatory materials. The findings emphasise the importance and potential of zoos as public engagement centres for the biological sciences.</description>
    <dc:date>2012-04-04T00:00:00Z</dc:date>
    <dc:creator>Bowler, Mark Timothy</dc:creator>
    <dc:creator>Buchanan-Smith, Hannah M</dc:creator>
    <dc:creator>Whiten, Andrew</dc:creator>
    <dc:description>Recent years have seen increasing encouragement by research institutions and funding bodies for scientists to actively engage with the public, who ultimately finance their work. Animal behaviour as a discipline possesses several features, including its inherent accessibility and appeal to the public, that may help it occupy a particularly successful niche within these developments. It has also established a repertoire of quantitative behavioural methodologies that can be used to document the public's responses to engagement initiatives. This kind of assessment is becoming increasingly important considering the enormous effort now being put into public engagement projects, whose effects are more often assumed than demonstrated. Here we report our first attempts to quantify relevant aspects of the behaviour of a sample of the hundreds of thousands of visitors who pass through the ‘Living Links to Human Evolution Research Centre’ in Edinburgh Zoo. This University research centre actively encourages the public to view ongoing primate research and associated science engagement activities. Focal follows of visitors and scan sampling showed substantial ‘dwell times’ in the Centre by common zoo standards and the addition of new engagement elements in a second year was accompanied by significantly increased overall dwell times, tripling for the most committed two thirds of visitors. Larger groups of visitors were found to spend more time in the Centre than smaller ones. Viewing live, active science was the most effective activity, shown to be enhanced by novel presentations of carefully constructed explanatory materials. The findings emphasise the importance and potential of zoos as public engagement centres for the biological sciences.</dc:description>
  </item>
  <item rdf:about="http://hdl.handle.net/10023/2555">
    <title>Evidence for weak or linear conformity but not for hyper-conformity in an everyday social learning context</title>
    <link>http://hdl.handle.net/10023/2555</link>
    <description>Abstract: Conformity is thought to be an important force in cultural evolution because it has the potential to stabilize cooperation in large groups, potentiate group selection and thus explain uniquely human behaviors. However, the effects of such conformity on cultural and biological evolution will depend much on the way individuals are influenced by the frequency of alternative behavioral options witnessed. Theoretical modeling has suggested that only what we refer to as ‘hyper-conformity’, an exaggerated tendency to perform the most frequent behavior witnessed in other individuals, is able to increase within-group homogeneity and between-group diversity, for instance. Empirically however, few experiments have addressed how the frequency of behavior witnessed affects behavior. Accordingly we performed an experiment to test for the presence of conformity in a natural situation with humans. Visitors to a Zoo exhibit were invited to write or draw answers to questions on A5 cards and potentially win a small prize. We manipulated the proportion of existing writings versus drawings visible to visitors and measured the proportion of written cards submitted. We found a strong and significant effect of the proportion of text displayed on the proportion of text in the answers, thus demonstrating social learning. We show that this effect is approximately linear, with potentially a small, weak-conformist component but no hyper-conformist one. The present experiment therefore provides evidence for linear conformity in humans in a very natural context.</description>
    <dc:date>2012-02-20T00:00:00Z</dc:date>
    <dc:creator>Claidiere, Nicolas</dc:creator>
    <dc:creator>Bowler, Mark Timothy</dc:creator>
    <dc:creator>Whiten, Andrew</dc:creator>
    <dc:description>Conformity is thought to be an important force in cultural evolution because it has the potential to stabilize cooperation in large groups, potentiate group selection and thus explain uniquely human behaviors. However, the effects of such conformity on cultural and biological evolution will depend much on the way individuals are influenced by the frequency of alternative behavioral options witnessed. Theoretical modeling has suggested that only what we refer to as ‘hyper-conformity’, an exaggerated tendency to perform the most frequent behavior witnessed in other individuals, is able to increase within-group homogeneity and between-group diversity, for instance. Empirically however, few experiments have addressed how the frequency of behavior witnessed affects behavior. Accordingly we performed an experiment to test for the presence of conformity in a natural situation with humans. Visitors to a Zoo exhibit were invited to write or draw answers to questions on A5 cards and potentially win a small prize. We manipulated the proportion of existing writings versus drawings visible to visitors and measured the proportion of written cards submitted. We found a strong and significant effect of the proportion of text displayed on the proportion of text in the answers, thus demonstrating social learning. We show that this effect is approximately linear, with potentially a small, weak-conformist component but no hyper-conformist one. The present experiment therefore provides evidence for linear conformity in humans in a very natural context.</dc:description>
  </item>
  <item rdf:about="http://hdl.handle.net/10023/2446">
    <title>Female chimpanzees use copulation calls flexibly to prevent social competition</title>
    <link>http://hdl.handle.net/10023/2446</link>
    <description>Abstract: The adaptive function of copulation calls in female primates has been debated for years. One influential idea is that copulation calls are a sexually selected trait, which enables females to advertise their receptive state to males. Male-male competition ensues and females benefit by getting better mating partners and higher quality offspring. We analysed the copulation calling behaviour of wild female chimpanzees (Pan troglodytes schweinfurthii) at Budongo Forest, Uganda, but found no support for the male-male competition hypothesis. Hormone analysis showed that the calling behaviour of copulating females was unrelated to their fertile period and likelihood of conception. Instead, females called significantly more while with high-ranking males, but suppressed their calls if high-ranking females were nearby. Copulation calling may therefore be one potential strategy employed by female chimpanzees to advertise receptivity to high-ranked males, confuse paternity and secure future support from these socially important individuals. Competition between females can be dangerously high in wild chimpanzees, and our results indicate that females use their copulation calls strategically to minimise the risks associated with such competition.</description>
    <dc:date>2008-06-18T00:00:00Z</dc:date>
    <dc:creator>Townsend, Simon W.</dc:creator>
    <dc:creator>Deschner, Tobias</dc:creator>
    <dc:creator>Zuberbuehler, Klaus</dc:creator>
    <dc:description>The adaptive function of copulation calls in female primates has been debated for years. One influential idea is that copulation calls are a sexually selected trait, which enables females to advertise their receptive state to males. Male-male competition ensues and females benefit by getting better mating partners and higher quality offspring. We analysed the copulation calling behaviour of wild female chimpanzees (Pan troglodytes schweinfurthii) at Budongo Forest, Uganda, but found no support for the male-male competition hypothesis. Hormone analysis showed that the calling behaviour of copulating females was unrelated to their fertile period and likelihood of conception. Instead, females called significantly more while with high-ranking males, but suppressed their calls if high-ranking females were nearby. Copulation calling may therefore be one potential strategy employed by female chimpanzees to advertise receptivity to high-ranked males, confuse paternity and secure future support from these socially important individuals. Competition between females can be dangerously high in wild chimpanzees, and our results indicate that females use their copulation calls strategically to minimise the risks associated with such competition.</dc:description>
  </item>
  <item rdf:about="http://hdl.handle.net/10023/2003">
    <title>Implied motion activation in cortical area MT can be explained by visual low-level features</title>
    <link>http://hdl.handle.net/10023/2003</link>
    <description>Abstract: To investigate form-related activity inmotion-sensitive cortical areas, we recorded cell responses to animate implied motion in macaque middle temporal (MT) and medial superior temporal (MST) cortex and investigated these areas using fMRI in humans. In the single-cell studies, we compared responses with static images of human or monkey figures walking or running left or right with responses to the same human and monkey figures standing or sitting still. We also investigated whether the view of the animate figure (facing left or right) that elicited the highest response was correlated with the preferred direction for moving random dot patterns. First, figures were presented inside the cell's receptive field. Subsequently, figures were presented at the fovea while a dynamic noise pattern was presented at the cell's receptive field location. The results show that MT neurons did not discriminate between figures on the basis of the implied motion content. Instead, response preferences for implied motion correlated with preferences for low-level visual features such as orientation and size. No correlation was found between the preferred view of figures implying motion and the preferred direction for moving random dot patterns. Similar findings were obtained in a smaller population of MST cortical neurons. Testing human MT+ responses with fMRI further corroborated the notion that low-level stimulus features might explain implied motion activation in human MT+. Together, these results suggest that prior human imaging studies demonstrating animate implied motion processing in area MT+ can be best explained by sensitivity for low-level features rather than sensitivity for the motion implied by animate figures.</description>
    <dc:date>2011-06-01T00:00:00Z</dc:date>
    <dc:creator>Lorteije, Jeannette A. M.</dc:creator>
    <dc:creator>Barraclough, Nick E.</dc:creator>
    <dc:creator>Jellema, Tjeerd</dc:creator>
    <dc:creator>Raemaekers, Mathijs</dc:creator>
    <dc:creator>Duijnhouwer, Jacob</dc:creator>
    <dc:creator>Xiao, Dengke</dc:creator>
    <dc:creator>Oram, Mike W.</dc:creator>
    <dc:creator>Lankheet, Martin J. M.</dc:creator>
    <dc:creator>Perrett, David I.</dc:creator>
    <dc:creator>van Wezel, Richard J. A.</dc:creator>
    <dc:description>To investigate form-related activity inmotion-sensitive cortical areas, we recorded cell responses to animate implied motion in macaque middle temporal (MT) and medial superior temporal (MST) cortex and investigated these areas using fMRI in humans. In the single-cell studies, we compared responses with static images of human or monkey figures walking or running left or right with responses to the same human and monkey figures standing or sitting still. We also investigated whether the view of the animate figure (facing left or right) that elicited the highest response was correlated with the preferred direction for moving random dot patterns. First, figures were presented inside the cell's receptive field. Subsequently, figures were presented at the fovea while a dynamic noise pattern was presented at the cell's receptive field location. The results show that MT neurons did not discriminate between figures on the basis of the implied motion content. Instead, response preferences for implied motion correlated with preferences for low-level visual features such as orientation and size. No correlation was found between the preferred view of figures implying motion and the preferred direction for moving random dot patterns. Similar findings were obtained in a smaller population of MST cortical neurons. Testing human MT+ responses with fMRI further corroborated the notion that low-level stimulus features might explain implied motion activation in human MT+. Together, these results suggest that prior human imaging studies demonstrating animate implied motion processing in area MT+ can be best explained by sensitivity for low-level features rather than sensitivity for the motion implied by animate figures.</dc:description>
  </item>
  <item rdf:about="http://hdl.handle.net/10023/1587">
    <title>The syntax and meaning of wild gibbon songs</title>
    <link>http://hdl.handle.net/10023/1587</link>
    <description>Abstract: Spoken language is a result of the human capacity to assemble simple vocal units into more complex utterances, the basic carriers of semantic information. Not much is known about the evolutionary origins of this behaviour. The vocal abilities of non-human primates are relatively unimpressive in comparison, with gibbon songs being a rare exception. These apes assemble a repertoire of call notes into elaborate songs, which function to repel conspecific intruders, advertise pair bonds, and attract mates. We conducted a series of field experiments with white-handed gibbons at Khao Yai National Park, Thailand, which showed that this ape species uses songs also to protect themselves against predation. We compared the acoustic structure of predatory-induced songs with regular songs that were given as part of their daily routine. Predator-induced songs were identical to normal songs in the call note repertoire, but we found consistent differences in how the notes were assembled into songs. The responses of out-of-sight receivers demonstrated that these syntactic differences were meaningful to conspecifics. Our study provides the first evidence of referential signalling in a free-ranging ape species, based on a communication system that utilises combinatorial rules.</description>
    <dc:date>2006-12-20T00:00:00Z</dc:date>
    <dc:creator>Clarke, Esther</dc:creator>
    <dc:creator>Reichard, Ulrich H.</dc:creator>
    <dc:creator>Zuberbuehler, Klaus</dc:creator>
    <dc:description>Spoken language is a result of the human capacity to assemble simple vocal units into more complex utterances, the basic carriers of semantic information. Not much is known about the evolutionary origins of this behaviour. The vocal abilities of non-human primates are relatively unimpressive in comparison, with gibbon songs being a rare exception. These apes assemble a repertoire of call notes into elaborate songs, which function to repel conspecific intruders, advertise pair bonds, and attract mates. We conducted a series of field experiments with white-handed gibbons at Khao Yai National Park, Thailand, which showed that this ape species uses songs also to protect themselves against predation. We compared the acoustic structure of predatory-induced songs with regular songs that were given as part of their daily routine. Predator-induced songs were identical to normal songs in the call note repertoire, but we found consistent differences in how the notes were assembled into songs. The responses of out-of-sight receivers demonstrated that these syntactic differences were meaningful to conspecifics. Our study provides the first evidence of referential signalling in a free-ranging ape species, based on a communication system that utilises combinatorial rules.</dc:description>
  </item>
</rdf:RDF>

