Eben Goodale

Interspecific information transfer influences animal community structure

Acquiring information from the cues and signals of other species of the same trophic level is widespread among animals, and can help individuals exploit resources and avoid predators. But can such interspecific information transfer also influence the spatial structure of species within communities? Whereas some species use heterospecific information without changing their position, we review research that indicates that heterospecific information is a driving factor in the formation or maintenance of temporary or stable mixed-species groups. Heterospecific information can also influence the organization of such groups, including leadership. Further, animals sometimes select habitats using heterospecific information. We survey interspecific information transfer, and evaluate the morphological, ecological and behavioral factors that make some species information sources and others information seekers.

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.

Persistence of song types in Darwin's finches, Geospiza fortis, over four decades

Learned bird songs evolve via cultural evolution, with song patterns transmitted across generations by imitative learning. In Darwins finches of the Galápagos Islands, males learn songs from their fathers, and song types can be maintained across multiple generations. However, little is known about the time frame over which specific song types are preserved, in the face of copy errors and corresponding modifications to song structure. Here we investigate cultural evolution in songs of male Geospiza fortis, at Academy Bay, Santa Cruz Island, comparing songs recorded in 1961 by R. Bowman (20 individuals) to those recorded in 1999 by J. Podos (16 individuals). For each individual, we characterized four timing and six frequency parameters, and assessed inter-individual variation in song structure using multivariate analysis. Several 1961 song types persisted into 1999, some with remarkable fidelity. Variation among song types was extensive during both years, and we detected no changes in 10 vocal parameters across the sampling period. These results illustrate temporal continuity in a culturally acquired trait, and raise questions about mechanisms that promote stability in song structure.

Bird fruit preferences match the frequency of fruit colours in tropical Asia

While many factors explain the colour of fleshy fruits, it is thought that black and red fruits are common in part because frugivorous birds prefer these colours. We examined this still controversial hypothesis at a tropical Asian field site, using artificial fruits, fresh fruits, four wild-caught resident frugivorous bird species, and hand-raised naïve birds from three of the same species. We demonstrate that all birds favored red artificial fruits more than yellow, blue, black and green, although the artificial black colour was found subsequently to be similar to the artificial blue colour in its spectral reflectance. Wild-caught birds preferred both black and red fleshy natural fruits, whereas hand-raised naïve birds preferred black to red natural fleshy fruits and to those of other colours. All birds avoided artificial and naturally ripe green fruits. The inter-individual variation in colour choice was low and the preferences were constant over time, supporting the hypothesis that bird colour preferences are a contributing factor driving fruit colour evolution in tropical Asia.

The innate responses of bumble bees to flower patterns: separating the nectar guide from the nectary changes bee movements and search time

Nectar guides can enhance pollinator efficiency and plant fitness by allowing pollinators to more rapidly find and remember the location of floral nectar. We tested if a radiating nectar guide around a nectary would enhance the ability of naïve bumble bee foragers to find nectar. Most experiments that test nectar guide efficacy, specifically radiating linear guides, have used guides positioned around the center of a radially symmetric flower, where nectaries are often found. However, the flower center may be intrinsically attractive. We therefore used an off-center guide and nectary and compared “conjunct” feeders with a nectar guide surrounding the nectary to “disjunct” feeders with a nectar guide separated from the nectary. We focused on the innate response of novice bee foragers that had never previously visited such feeders. We hypothesized that a disjunct nectar guide would conflict with the visual information provided by the nectary and negatively affect foraging. Approximately, equal numbers of bumble bees (Bombus impatiens) found nectar on both feeder types. On disjunct feeders, however, unsuccessful foragers spent significantly more time (on average 1.6-fold longer) searching for nectar than any other forager group. Successful foragers on disjunct feeders approached these feeders from random directions unlike successful foragers on conjunct feeders, which preferentially approached the combined nectary and nectar guide. Thus, the nectary and a surrounding nectar guide can be considered a combination of two signals that attract naïve foragers even when not in the floral center.

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 structure of mixed-species bird flocks, and their response to anthropogenic disturbance, with special reference to East Asia

Mixed-species flocks of birds are distributed world-wide and can be especially dominant in temperate forests during the non-breeding season and in tropical rainforests year-round. We review from a community ecology perspective what is known about the structure and organization of flocks, emphasizing that flocking species tend to be those particularly vulnerable to predation, and flocks tend to be led by species that are able to act as sources of information about predators for other species. Studies on how flocks respond to fragmentation and land-use intensification continue to accumulate, but the question of whether the flock phenomenon makes species more vulnerable to anthropogenic change remains unclear. We review the literature on flocks in East Asia and demonstrate there is a good foundation of knowledge on which to build. We then outline potentially fruitful future directions, focusing on studies that can investigate how dependent species are on each other in flocks, and how such interdependencies might affect avian habitat selection in the different types of human-modified environments of this region.

Effects of forests, roads and mistletoe on bird diversity in monoculture rubber plantations.

Rising global demand for natural rubber is expanding monoculture rubber (Hevea brasilensis) at the expense of natural forests in the Old World tropics. Conversion of forests into rubber plantations has a devastating impact on biodiversity and we have yet to identify management strategies that can mitigate this. We determined the life-history traits that best predict bird species occurrence in rubber plantations in SW China and investigated the effects of surrounding forest cover and distance to roads on bird diversity. Mistletoes provide nectar and fruit resources in rubber so we examined mistletoe densities and the relationship with forest cover and rubber tree diameter. In rubber plantations, we recorded less than half of all bird species extant in the surrounding area. Birds with wider habitat breadths and low conservation value had a higher probability of occurrence. Species richness and diversity increased logarithmically with surrounding forest cover, but roads had little effect. Mistletoe density increased exponentially with rubber tree diameters, but was unrelated to forest cover. To maximize bird diversity in rubber-dominated landscapes it is therefore necessary to preserve as much forest as possible, construct roads through plantations and not forest, and retain some large rubber trees with mistletoes during crop rotations.

Flight initiation distance as behavioral indicator of hunting pressure: a case study of the Sooty-headed Bulbul (Pycnonotus aurigaster) in Xishuangbanna, SW China

Traditional assessments of anthropogenic impacts on biodiversity often ignore hunting pressure or use subjective categories (e.g. high, medium or low) that cannot be readily understood by readers or replicated in other studies. Although animals often appear tame in habitats without hunting compared to habitats with hunting, few studies have demonstrated such effects. We determined the flight initiation distance (FID; i.e. human-animal distance when the animal begins to flee) of a common frugivorous bird of Southeast Asia, Sooty-headed Bulbul (Pycnonotus aurigaster) across a gradient of hunting pressures in Xishuangbanna, Yunnan, SW China. Controlling for confounding effects, we show that FID increased with hunting pressure, which was quantitatively measured through encounters with hunters. As FIDs respond more specifically to hunting than other defaunation metrics, we suggest they can be used as behavioral indicators of hunting pressure in developing conservation strategies.