Sarath W. Kotagama

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

Positive Relationships Between Association Strength and Phenotypic Similarity Characterize the Assembly of Mixed-Species Bird Flocks Worldwide

Competition theory predicts that local communities should consist of species that are more dissimilar than expected by chance. We find a strikingly different pattern in a multicontinent data set (55 presence-absence matrices from 24 locations) on the composition of mixed-species bird flocks, which are important subunits of local bird communities the world over. By using null models and randomization tests followed by meta-analysis, we find the association strengths of species in flocks to be strongly related to similarity in body size and foraging behavior and higher for congeneric compared with noncongeneric species pairs. Given the local spatial scales of our individual analyses, differences in the habitat preferences of species are unlikely to have caused these association patterns; the patterns observed are most likely the outcome of species interactions. Extending group-living and social-information-use theory to a heterospecific context, we discuss potential behavioral mechanisms that lead to positive interactions among similar species in flocks, as well as ways in which competition costs are reduced. Our findings highlight the need to consider positive interactions along with competition when seeking to explain community assembly.

Two sympatric species of passerine birds imitate the same raptor calls in alarm contexts.

While some avian mimics appear to select sounds randomly, other species preferentially imitate sounds such as predator calls that are associated with danger. Previous work has shown that the Greater Racket-tailed Drongo (Dicrurus paradiseus) incorporates predator calls and heterospecific alarm calls into its own species-typical alarm vocalizations. Here, we show that another passerine species, the Sri Lanka Magpie (Urocissa ornata), which inhabits the same Sri Lankan rainforest, imitates three of the same predator calls that drongos do. For two of these call types, there is evidence that magpies also use them in alarm contexts. Our results support the hypothesis that imitated predator calls can serve as signals of alarm to multiple species.

The frequency of vocal mimicry associated with danger varies due to proximity to nest and nesting stage in a passerine bird

Several species of birds vocally imitate sounds associated with danger. Two anecdotal studies suggest that such ‘danger mimicry’ increases during nesting, but such a relationship has not been quantitatively demonstrated. Sri Lanka drongos (Dicrurus paradiseus lophorhinus) are known to imitate predators and other species’ mobbing and alarm calls in alarm contexts. Here we investigated whether drongos vary their production of danger mimicry in different nesting stages (building, incubation, nests with hatchlings, fledglings still outside of mixed-species flocks), and when foraging away from young in mixed-species flocks. We recorded drongos over two breeding seasons at 14 different nesting trees, used year-after-year. We found that of all the types of danger mimicry, imitation of predators was the most common and exclusive to drongos that had young offspring. Such predator mimicry was observed at a higher rate during the hatchling and fledgling stages compared to incubation or flocks. Danger mimicry did not, however, increase during this stage in isolation: drongo species-specific alarm calls also increased, and the close connection between these two types of calls did not appear to change. Although it is possible that the association between danger mimicry and species-specific alarm calls could help young birds learn sounds associated with danger, the performance of this behaviour does not seem exclusive enough to interactions between adult drongos and their offspring to meet functional definitions of teaching.

The effect of land-use on the diversity and mass-abundance relationships of understory avian insectivores in Sri Lanka and southern India

Understory avian insectivores are especially sensitive to deforestation, although regional differences in how these species respond to human disturbance may be linked to varying land-use histories. South Asia experienced widespread conversion of forest to agriculture in the nineteenth century, providing a comparison to tropical areas deforested more recently. In Sri Lanka and the Western Ghats of India, we compared understory insectivores to other guilds, and to insectivores with different vertical strata preferences, both inside mixed-species flocks and for the whole bird community. Overall species richness did not change across the land-use gradient, although there was substantial turnover in species composition between land-use types. We found that the proportion of species represented by insectivores was ~1.14 times higher in forest compared to agriculture, and the proportion of insectivores represented by understory species was ~1.32 times higher in forests. Mass-abundance relationships were very different when analyzed on mixed-species flocks compared to the total community, perhaps indicating reduced competition in these mutualisms. We show that South Asia fits the worldwide pattern of understory insectivores declining with increased land-use intensity, and conclude that these species can be used globally as indicator and/or umbrella species for conservation across different disturbance time scales.

Horizontal and vertical species turnover in tropical birds in habitats with differing land use.

Large tracts of tropical rainforests are being converted into intensive agricultural lands. Such anthropogenic disturbances are known to reduce species turnover across horizontal distances. But it is not known if they can also reduce species turnover across vertical distances (elevation), which have steeper climatic differences. We measured turnover in birds across horizontal and vertical sampling transects in three land-use types of Sri Lanka: protected forest, reserve buffer and intensive-agriculture, from 90 to 2100 m a.s.l. Bird turnover rates across horizontal distances were similar across all habitats, and much less than vertical turnover rates. Vertical turnover rates were not similar across habitats. Forest had higher turnover rates than the other two habitats for all bird species. Buffer and intensive-agriculture had similar turnover rates, even though buffer habitats were situated at the forest edge. Therefore, our results demonstrate the crucial importance of conserving primary forest across the full elevational range available.