Vocalizations in the Grey Butcherbird Cracticus Torquatus with Emphasis on Structure in Male Breeding Song: Implications for the Function and Evolution of Song from a Study of a Southern Hemisphere Species.

Abstract

An undue focus on Temperate Zone oscines (songbirds or passerines) has led to a geographical bias in interpretation of song function and evolution. This bias led initially to relatively simplistic theories of the function of bird song with vocalizations divided into 'songs' and 'calls'. Songs were complex, learned vocalizations, given by males in the breeding season, functioning in territory defence and mate attraction and stimulation. Calls, on the other hand, were simple innate vocalizations serving more immediate needs such as begging for food and raising an alarm. Female song, where it occurred, was considered an aberration. Further studies suggested that complex songs were associated with mate attraction functions while simpler songs were associated with territory defence. However it became apparent that the distinction between songs and calls was not nearly so clear-cut and the supposed connection between complexity and function in song was questioned. Moreover it was realised that female song could not be dismissed as a mere aberration. Another problem was the function of the dawn chorus, where research had failed to find a consistent, all-encompassing explanation. Since most studies had been done on Northern Hemisphere songbirds, it was becoming clear that the geographical focus needed to be broadened. The life histories of Northern Hemisphere Temperate Zone songbirds are very different from those in many other regions. In contrast to the situation in this zone, maintenance of year-round territory, territory defence by both male and female, life-long social monogamy and extensive female vocalization are widespread in tropical, subtropical and Southern Hemisphere regions. Recently it has been suggested that more intensive studies of vocalizations in these regions might help clarify some of these issues and consequently an endemic Australian passerine was chosen for the current study. The study focused on the vocalizations of the grey butcherbird Cracticus torquatus Artamidae, which displays the life history features described above. The main study population was located in the Brisbane suburbs of Rainworth and Bardon. Additional data were gathered from other Brisbane suburbs and bushland sites within the city and at Lake Broadwater near Dalby, Queensland. Vocalizations were initially recorded electronically and analysed using Canary sound editing program. Vocalization data were supplemented using an aural recording method, which was independently checked for reliability. Behavioural data including posture during vocalizations and interactions with other birds were also gathered. Initial investigations revealed the existence of two main categories of vocalizations - those given by the family group during the day, all year round and those given at dawn by males during the breeding season. For the focal study populations, group vocalizations were studied throughout the year over several years, however the song given at dawn by males during the breeding season proved to be quite complex and three birds from three territories in the main study area were chosen as case studies. Recordings were made of the vocalizations of the three case study birds over three breeding seasons. Starting and finishing times (with respect to civil twilight) were recorded in order to determine both changes in song bout duration and starting time throughout the breeding season. Additional birds from the same area, from the other Brisbane suburbs and from the bushland sites were studied to check the validity of conclusions drawn from the case studies. The results of the investigations revealed a vocalization structure that contrasted strongly with the simple picture of bird song drawn from study of Northern Hemisphere Temperate Zone passerines. The vocalizations given during the day, often referred to as the 'song' of this species, were very different from the early dawn song given by the male during the breeding season. This latter appeared to be song sensu stricto according to the paradigms developed for Northern Hemisphere birds. The daytime vocalizations, however, fitted neither the classic definition of 'song' nor the classic definition of 'call'. This relatively long-term study revealed different starting time patterns and periodicity for day vocalizations and male breeding season song. Day vocalizations commenced at a fairly constant time with respect to civil twilight throughout the year but breeding season song started progressively earlier from the beginning through to the middle of the season then progressively later till the end of the season. Relative finishing time of breeding song however remained constant so that the duration of breeding season song gradually increased then decreased paralleling the change in starting times. A consequence of the two distinct classes of vocalizations was that there were essentially two distinct 'dawn choruses'. One, consisting of group vocals, was sung all year round; the other was given by males singing 'breeding season' song. Since there was no reason the expect that a single function would necessarily be ascribed to both choruses, this raised the possibility that some of the confusion surrounding the function and nature of the 'dawn chorus' originated from a failure to recognize the existence of two such choruses. Variation in time and space showed further differences between the two vocalization classes. The breeding song of each male was distinctly different from that of his neighbours and there was a marked change in the repertoire of any individual from one year to the next. In marked contrast, daytime vocalization repertoires of neighbouring groups were virtually indistinguishable and changed little from year to year. These findings, together with information from recent literature, suggested that the two song classes had a different ontogeny, function and possibly evolution. It was proposed that territory declaration was the function of dawn singing by grey butcherbird family groups but that the function most consistent with adult male dawn song was attraction of females for extra-pair copulations. It was suggested that chorusing itself was to some extent an accidental by-product of the advantage to the individual or group of singing at dawn although a recently proposed function, the social dynamics function, could not be ruled out. Further differences from the Northern Hemisphere situation were detected in subsong. First, subsong was given by birds in their first year and also by adult males. The finding of subsong in adult males was not without precedent as it has been documented previously for a small number of passerines, especially those that change repertoire from year to year. Subsong in young (first year) birds, however, was unusual in that birds practised in small groups rather than in the complete isolation usually associated with subsong. Moreover they did not practise adult male song but instead practised group daytime vocalizations. It was suggested that it was important for birds to learn to sing in company for the important task of group territory defence. Further investigation of the literature and observations during the present study revealed similar vocalization classes and behaviours in other members of the Artamidae and other endemic Australasian taxa. These literature investigations also revealed that the possession two song vocalization classes was quite widespread although they tended to be restricted (but not exclusive) to males rather than found in males and females. These findings led to further research into the significance of Australia in the evolution of songbirds, the role of co-operative breeding in Australian passerines, and finally to an hypothesis for a possible origin of male bird song. It is suggested that male song arose in a social environment where the male and female were in frequent vocal communication. If the tendency to seek extra-pair copulations (EPCs) and female choice had already been incorporated into the suite of passerine behaviours, it would be necessary to avoid the mate during such activities and the male would need to advertise with a s