Throughout the year, African pied starlings live in flocks of 15—25 individuals. Stable monogamous pairs re-use the same nest sites in successive breeding seasons September—January. Several nests can be found close to each other in burrows or holes. During the breeding season, up to seven helpers can feed the young with the parents, and helpers may feed young at three different nests during a single breeding season Craig, During the non-breeding season, pied starling groups may be nomadic and join other groups at communal roost sites Craig and Feare, This study was conducted in the Eastern Cape region in South Africa where the four species occur, often at the same sites.
Songs have been recorded since , mainly during the breeding season. Red-winged starling vocalizations were recorded primarily on Rhodes University campus in Grahamstown and in the vicinity — Pale-winged starling vocalizations were recorded at one site: Graaff-Reinet Pied starling vocalizations were recorded at three sites: Table Farm, Queenstown and Graaff-Reinet to and Most recordings were obtained in the morning 6—10 a. According to Fry et al.
Vocalizations were analyzed using homemade software for song analyses ANA, Richard, The amount of song recorded is summarized in Table 3.
Table 3. Here we focused our analyses on the temporal aspects of songs. Indeed, most studies on interspecific comparisons of vocalizations have focused on quantitative aspects, such as the repertoire size Catchpole, ; Kroodsma, ; MacComb and Semple, Whereas temporal aspects of vocal signals or vocal interactions have so far been little studied, they nonetheless could provide a wealth of information regarding the influence of social life on the evolution of vocal communication. We predicted that social life, in terms of the number of social partners or distance between partners for example, would affect the temporal structure of song.
Two categories of songs could be distinguished: discontinuous songs , corresponding to unitary notes or short motifs a fixed combination of acoustic elements produced at discrete intervals, and continuous songs in which long sequences are produced, with less than 0. For each species, we measured: 1- sequence duration, 2- intervals between two successive sequences or two successive discontinuous motifs, 3- the motif duration, 4- the number of motifs per sequence, 5- intervals between two successive motifs within a sequence.
The four species showed clear differences in the temporal organization of their song. Considering the proportion of continuous and discontinuous songs, a gradient was observed from O. Interestingly, this gradient corresponded to the increase in the complexity of social life Figure 11 : the more the species showed a complex and especially family type of social organization in terms of number of congeners and nest proximity , the more their songs were produced in a continuous manner. Song sequence duration and sequence interval duration for the 4 species of African sturnids.
Song overlap was never observed in O. On the contrary, in L. Both alternating and overlapping song interactions are also regularly observed in O. Whistles of a male and a female O. Chorus of L. The data presented here on one animal model reveal the interest of focusing on one question here the temporal features of song that may or may not lead to alternating vocal interactions and examining the different facets of the question. To the question: do European starlings show turn-taking in their vocal exchanges between males?
It has been proposed that warbling could play some stimulating role on the physiology of the listeners but also on the emitters too as found in budgerigars by Brockway and Adret-Hausberger and Jenkins Warbling is often associated with excitation behaviors such as visual displays and the production of high pitched trills, especially in the breeding season Verheyen, Fundamentally, male starlings show movements of the head, typical of observation, during the silent interval between successive whistles and an erect posture while they are more in an oblique posture, with or without wing displays and a low reactivity while warbling.
Excitation may lead to more overlap. Interestingly, the comparative study of African starlings reflects these findings: the more communal the species, the more song overlap and choruses appear during close-range interactions, and the more continuous the song. The more territorial and long distant interactive a species is, the more alternation there is, hence the more discontinuous the song structures are. Some species like the European starling and the pale-winged starling show both song styles, reflecting the different contexts of interaction. According to Hauser , the timing of calling in macaques may be altered in such a way that it is used by individuals to manipulate or facilitate social relationships.
One may speculate that the need for mutual intelligibility and information seeking but also the need for giving and receiving attention, a potential mediator of social bonding Fedurek et al. Humans too may produce choirs that are perceived as a communal display rather than an interaction between individuals. In the Dogons, as mentioned earlier, observing rules in language coincides with law and order in the society Calame-Griaule, It is true too that spacing of the vocalizations requires calmness, control and attention toward the others instead of being self-centered.
For France et al. This is reminiscent of the gradient observed in species of the starling family Sturnidae.
Other communal breeders and group living animals such as the Australian magpies also favor choruses and overlap of songs e. This recalls some human conversations where the dominant individual disregards the other's turn. For Takahashi et al. Indeed the temporal features of animal vocal interactions in many ways parallel human communication. The use of rule systems for vocal communication is not limited to human beings. This review makes two additional points: turn-taking is one characteristic feature of human conversations but choruses might well be of interest if the social evolution of language and the intercultural aspects are to be considered; more integrative studies such as those described here and in progress for starlings are needed in order to tackle the question of the evolution of rule-governed communication in language.
The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest. Adret-Hausberger, M. Social influences on the whistled songs of starlings. Temporal dynamics of dialects in the whistled songs of starlings.
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These two levels have been studied separately. We hypothesize that joint action exploits forms of ostension found at the second level, and that ostensive communication draws on forms of coordination found at the first level. Through our integrated study of the two levels we aim to redefine the relation between coordination, communication, and cultural transmission.
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At a second level, established through ostensive communication, common ground includes also general knowledge of enduring relevance. These two levels have been studied separately. We hypothesize that joint action exploits forms of ostension found at the second level, and that ostensive communication draws on forms of coordination found at the first level.
Through our integrated study of the two levels we aim to redefine the relation between coordination, communication, and cultural transmission. Our common program will close gaps between research that has focused on the processes and representations that enable joint action Call, Knoblich and research that has addressed intentional communication and its role in cultural transmission Gergely, Sperber.
It will integrate the study of a embodied cognitive mechanisms for interpersonal coordination, b shared intentions and shared task representations, c ostensive communication, d natural pedagogy, and e how all of the above provide the micro-mechanisms of cultural transmission.