ALARM CALLING IN SRI LANKAN MIXED-SPECIES BIRD FLOCKS

Abstract Vocal alarm calls are important to the vigilance and likely the organization of mixed-species flocks, but community-wide studies of alarm calling in flocks are lacking. We investigated which species alarm-call, and the characteristics of their calls, in a large flock system of a Sri Lankan rainforest. We recorded naturally elicited alarm calls during several attacks by Accipiter hawks and while following flocks for 10 h. We then artificially elicited alarms by throwing a stick to the side of the flock, in a total of 70 trials at 30 flock sites. The Orange-billed Babbler (Turdoides rufescens) was the most frequent caller to both the artificial and natural stimuli, followed by the Greater Racket-tailed Drongo (Dicrurus paradiseus). Several other species also called, and multiple species often called to the same stimulus (in 23 trials, and in all of the hawk attacks). The species differed in their rapidity of response and in their sensitivity to different natural stimuli. Calls of the gregarious babbler usually provided a first, unreliable warning of an incoming threat, whereas later calls of other species emphasized the seriousness of the threat. We suggest that birds in mixed-species flocks may be particularly aware of aerial predators for two reasons: (1) a “numbers effect,” whereby nongregarious species are more aware of predators when surrounded by large numbers of other species; and (2) an “information effect,” whereby species differ in the information available in their alarm calls, leading to an accumulation of information in a mixed-species flock. Llamadas de Alarma en Bandadas Mixtas de Aves en Sri Lanka

[1]  E. Goodale,et al.  The composition and spatial organisation of mixed- species flocks in a Sri Lankan rainforest , 2004 .

[2]  M. Moynihan,et al.  The organization and probable evolution of some mixed species flocks of Neotropical birds , 1962 .

[3]  P. Marler,et al.  Characteristics of Some Animal Calls , 1955, Nature.

[4]  R. Kannan A Guide to the Birds of India, Pakistan, Nepal, Bangladesh, Bhutan, Sri Lanka, and the Maldives Richard Grimmett Carol Inskipp Tim Inskipp , 1999 .

[5]  E. Curio,et al.  The hearing of an avian predator and its avian prey , 1986, Behavioral Ecology and Sociobiology.

[6]  P. Gaddis Mixed Flocks, Accipiters, and Antipredator Behavior , 1980 .

[7]  H. L. Bell A Bird Community of Lowland Rainforest in New Guinea. 5. Mixed-Species Feeding Flocks , 1983 .

[8]  M. Elgar,et al.  PREDATOR VIGILANCE AND GROUP SIZE IN MAMMALS AND BIRDS: A CRITICAL REVIEW OF THE EMPIRICAL EVIDENCE , 1989, Biological reviews of the Cambridge Philosophical Society.

[9]  S. Haftorn CONTEXTS AND POSSIBLE FUNCTIONS OF ALARM CALLING IN THE WILLOW TIT, PARUS MONTANUS; THE PRINCIPLE OF ‘BETTER SAFE THAN SORRY’ , 2000 .

[10]  T. Caro,et al.  Pursuit-deterrence revisited. , 1995, Trends in ecology & evolution.

[11]  J. Dick,et al.  Compact handbook of the birds of India and Pakistan , 1987 .

[12]  K. Sanderson,et al.  Alarm Vocalisations in Australian Birds: Convergent Characteristics and Phylogenetic Differences , 1994 .

[13]  J. Terborgh,et al.  Mixed flocks and polyspecific associations: Costs and benefits of mixed groups to birds and monkeys , 1990, American journal of primatology.

[14]  J. Diamond Flocks of Brown and Black New Guinean Birds: A Bicoloured Mixed-species Foraging Association , 1987 .

[15]  J. M. Smith,et al.  The Evolution of Alarm Calls , 1965, The American Naturalist.

[16]  K. Sullivan,et al.  The advantages of social foraging in downy woodpeckers , 1984, Animal Behaviour.

[17]  A. J. Gaston Social behaviour within groups of jungle babblers (Turdoides striatus) , 1977, Animal Behaviour.

[18]  O. Hogstad Alarm calling by willow tits, Parus montanus, as mate investment , 1995, Animal Behaviour.

[19]  M. Ficken Acoustic characteristics of alarm calls associated with predation risk in chickadees , 1990, Animal Behaviour.

[20]  J. Dick,et al.  Handbook of the birds of India and Pakistan together with those of Bangladesh, Nepal, Bhutan and Sri Lanka , 1978 .

[21]  D. H. Morse,et al.  Ecological Aspects of Some Mixed-Species Foraging Flocks of Birds , 1970 .

[22]  J. Thiollay Frequency of Mixed Species Flocking in Tropical Forest Birds and Correlates of Predation Risk: An Intertropical Comparison , 1999 .

[23]  A. Zahavi,et al.  Cooperative Breeding in Birds: Arabian Babblers: the quest for social status in a cooperative breeder , 1990 .

[24]  P. Sherman Nepotism and the evolution of alarm calls. , 1977, Science.

[25]  K. Sullivan Selective alarm calling by downy woodpeckers in mixed-species flocks , 1985 .

[26]  S. L. Lima,et al.  Collective detection of predatory attack by social foragers: fraught with ambiguity? , 1995, Animal Behaviour.

[27]  The role of alarm responses in the formation of mixed-species flocks of heathland birds , 1981, Behavioral Ecology and Sociobiology.

[28]  G. Powell Sociobiology and Adaptive Significance of Interspecific Foraging Flocks in the Neotropics , 1985 .

[29]  M. Ficken,et al.  Chickadee alarm calls: Does mate investment pay dividends? , 1979, Animal Behaviour.

[30]  H. Pulliam,et al.  On the advantages of flocking. , 1973, Journal of theoretical biology.

[31]  Michael H. Kutner Applied Linear Statistical Models , 1974 .

[32]  C. Munn Birds that ‘cry wolf’ , 1986, Nature.

[33]  R. L. Hutto The Composition and Social-Organization of Mixed-Species Flocks in a Tropical Deciduous Forest in Western Mexico , 1994 .

[34]  K. Sanderson,et al.  ACOUSTIC DISCRIMINATION OF PASSERINE ANTI-PREDATOR SIGNALS BY AUSTRALIAN RAPTORS , 1998 .

[35]  E. Curio,et al.  The adaptive significance of avian mobbing: I. Teleonomic hypotheses and predictions. , 1978 .

[36]  J. Podos Correlated evolution of morphology and vocal signal structure in Darwin's finches , 2001, Nature.

[37]  D. H. Morse Feeding Behavior and Predator Avoidance in Heterospecific Groups , 1977 .