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Notes to the database:
This database presents a representative selection of calls (N = 2707) that formed the basis of a quantitative analysis of the vocal repertoire of Barbary macaques (Hammerschmidt & Fischer 1998). In correspondence with our analysis, we noted for each call the sex of the caller, the individual identity (if known), the age, and a description of the context. For the purposes of the analysis, we established the a number of ‘main contexts’, each having a varying number of ‘subcontexts’. Finally, the 'cluster membership' is given. The cluster membership refers to a model of seven clusters that we chose according to the outcome of the analysis presented in the paper mentioned above.
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These are the main categories in the database:
Filename
- refers to the name of the soundfile
Cluster membership
We chose a 7-cluster model. Clusters are labelled with letters A through G.
Sex
Males
Females
Age
Infant: up to the age of 1 year
Young juvenile: age 1 – 2 years
Older juvenile: age 2 – 3.5 years
Subadult: females of 3.5 – 5 years of age; males 3.5 – 7 years of age
Adult: females age 5 years and older, males age 7 years and older.
Individual Identity
each individual is assigned an individual code
Main Context
Below, we give a brief description of the main contexts and the subcategories that we established.
During copulation, only females call whereas males remain silent. It was shown that the use of the mating call during copulation increases the likelihood of a ‘successful’ copulation, i.e. of an ejaculation in the male (Todt et al. 1995). There is also experimental evidence that these calls incite male-male competition (Semple 1998). During the mating season, females sometimes utter a call that sound like a mating call although the female is all by herself and not engaged in any close contact or interaction with a male (see Todt et al. 1995). Females also vocalize when they attempt to initiate a copulation, or when a male tries to do so. Sometimes, individuals vocalize after the male had dismounted from the female. Occasionally, third parties interfere during a copulation. Calls in this category were recorded from animals who attempted to do so. Accordingly, we established the following contexts:
Copulation
Copulation advertisement
After copulation
Disrupts copulation
Barbary macaques exhibit a system of extensive alloparental care involving both females and males (Paul 1984). Since males are typically interested in younger infants (Small 1990), the context ‘with male alloparent’ was observed for infants only, whereas the context ‘with mother’ or ‘with female alloparent’ occurs for infants and yearlings alike. We grouped calls accordingly into the following categories:
With female
With juvenile
With male
With Mother
Caregivers of all age classes and both sexes often emit typical vocalizations when they are staring at an infant, both when the infant is handled by other group members or when they are engaged in so-called ‘triadic interactions’ or ‘agonistic buffering’ (Deag & Crook 1971; Deag 1980; Taub 1980a; Taub 1980b; Taub 1984), i.e. when two animals are sitting together, often embracing each other, handling the infant, lip-smacking and teeth-chattering. We distinguished among situations in which the infant appears to be relaxed and ones in which it appears to be distressed and tries to get away from the caregiver.
Holds infant
Holds distressed infant
We recorded a large number of calls from females whose infants were in care of a male alloparent and who tried to retrieve their infant but were unsuccessful. In three instances, we were able to record calls from females whose dead infants had been removed. These subjects emitted very long sequences of calls many of which revealed a more tonal quality than the majority of Barbary macaque vocalizations. We distinguished between the two situations accordingly:
infant with other group member out of sight
dead infant removed by park staff
Young Barbary macaques are extremely vocal at dusk during sleeping cluster formation (“dusk calling”, see Hammerschmidt et al. 1994). Barbary macaque infants and juveniles regularly emitt long-lasting cry sequences when they are trying to establish body contact with their mothers in the sleeping tree. We grouped calls according to the following criteria: infant or juvenile is alone in the tree and away from any other animal while the group is settling down for the night; infant or juvenile attempts to establish body contact with a sleeping cluster. Frequently in such instances, the youngsters are rejected from adult animals, including their mothers; the young one has managed to establish body contact with other animals of that sleeping cluster.
alone in tree during sleeping cluster formation
close to sleeping cluster
in body contact with sleeping cluster
We recorded vocalizations from subjects at different age classes. Typically, females engage more in solitary play than males do and they stop participating in group playing at a younger age than males. A detailed analysis of the vocalizations emitted during group playing is currently in preparation (S. Kipper & D. Todt, unpublished data). We distinguished among situations in which the subject plays alone or with others.
plays alone
plays with others
On cold mornings, animals frequently establish large huddles after descent from their sleeping trees. These huddles are generally larger than sleeping clusters and, in contrast to sleeping clusters, often consist of adult subjects of both sexes. When younger animals are trying to achieve body contact with such a huddle or when they leave the huddle or the huddle breaks apart, they were observed to utter short series of calls. We noted whether the calling animal attempted to make or brake contact with animals in the huddle, or whether it was in contact while calling.
brakes contact
during contact
makes contact
In response to disturbances in their surroundings, we recorded calls from animals of all age-classes, except from infants. From our observations, however, it seemed as if subadult males and adult females were the most likely to call. A separate analysis showed that although these ‘alarm calls’ represent one single call type, a detailed analysis revealed that call can be discriminated in relation to the stimulus that elicited the calling. Playback experiments also showed that the subjects themselves were able to discriminate calls given in response to two different stimuli. For a more detailed description, see Fischer et al. 1995; Fischer 1998. We distinguished among calls with regard to the eliciting stimulus.
to dog
to observer
to park staff
to snake
In this context, we grouped calls that were given in quiet, apparently relaxed situations. Calls that were given when the caller looks at an infant were not recorded from animals who were actually in care of that infant, but from third parties.
sits alone
looks at infant
travels
We were able to record vocalizations given in agonistic situations from all age- and sexclasses. Calls were given both when subjects were actively challenging other group members and when they were threatened by other subjects. When subjects observe group fights in a distance, they sometimes utter low-frequency . In group fights and also during male-male fights which were observed most often during the mating season, subjects typically utter noisy screams. Challenging and threatening situations are more difficult to disentangle since animals often switch rapidly between approach and retreat. Hence, a variety of calls could be recorded during such situations. Lastly, we recorded a smaller number of calls after agonistic situations. As mentioned above, males rarely get harassed by other males since outside the mating season males most often try to avoid direct confrontation. We distinguished among the following categories.
challenges juvenile or female
challenges male
challenges observer
involved in group fight
post conflict
threatened by female
threatened by juvenile
threatened by male
watches fight in a distance
Acknowledgements
We thank Ellen Merz and Gilbert de Turckheim for permission to conduct this study at Rocamadour. Dietmar Todt provided essential resources and advice throughout our studies
References
Deag, J.M. 1980. Interactions between males and unweaned Barbary macaques: Testing the agonistic buffering hypothesis. Behaviour 75, 54-81.
Deag, J.M. & Crook, J.H. 1971. Social behaviour and agonistic buffering in the wild Barbary macaque, Macaca sylvanus. Folia primatologica 15, 183-200.
Fischer, J. 1998. Barbary macaques categorize shrill barks into two call types. Animal Behaviour, 55, 799-807.
Fischer, J. Hammerschmidt, K. & Todt D. 1995. Factors affecting acoustic variation in Barbary macaque (Macaca sylvanus) disturbance calls. Ethology, 101, 51-66.
Hammerschmidt, K., Ansorge, V., Fischer, J. & Todt, D. 1994. Dusk calling in Barbary macaques (Macaca sylvanus): Demand for Social Shelter. American Journal of Primatology 32, 277-289.
Hammerschmidt, K. & Fischer, J. 1998. The vocal repertoire of Barbary macaques: A quantitative analysis of a graded signal system. Ethology 104, 203-216.
Paul, A. 1984. Zur Sozialstruktur Und Sozialisation Semi-Freilebender Berberaffen (Macaca Sylvanus L.1758). Dissertation. Göttingen: Univ. Göttingen.
Semple, S. 1998. The function of Barbary macaque copulation calls. Proceedings of the Royal Society of London Series B-Biological Sciences 265, 287-291.
Small, M.F. 1990. Alloparental behaviour in Barbary macaques, Macaca sylvanus. Animal Behaviour 39, 297-306.
Taub, D.M. 1980a. Female choice and mating strategies among wild Barbary macaques (Macaca sylvanus). In: The Macaques: Studies in Ecology, Behaviour and Evolution (Ed. by D.M.Taub) New York, Van Nostrand Reinhold.
Taub, D.M. 1980b. Testing the 'agonistic buffering' hypothesis. 1. The dynamics of participation in the triadic interaction. Behavioral Ecology and Sociobiology 6, 187-197.
Taub, D.M. 1984. Male caretaking behaviour among wild Barbary macaques. In: Primate Paternalism (Ed. by D.M.Taub) New York, Van Nostrand Reinhold.
Todt, D., Hammerschmidt, K., Ansorge, V. & Fischer, J. 1995. The vocal behaviour of Barbary macaques: Call features and their performance in infants and adults. In: Current Topics in Primate Vocal Communication (Ed. by E. Zimmermann, J.D. Newman & U. Juergens), pp. 141-160. New York, Plenum Press.