Group-living Birds Research Articles

Audience effects in a group-living bird: How contact call rate is affected by vegetation and group size and composition.

Contact calling is a ubiquitous behavior of group-living animals. Yet in birds, beyond a general connection with group cohesion, its precise function is not well-understood, nor is it clear what stimulates changes in contact call rate. In an aviary experiment, we asked whether Swinhoe's White-eyes, Zosterops simplex, would regulate their own production of contact calls to maintain a specific rate at the group level. Specifically, we hypothesized that the sudden cessation of the group-level call rate could indicate an immediate predation threat, and we predicted that birds in smaller groups would call more to maintain a high call rate. We also investigated the effects of environmental characteristics, such as vegetation density, and social stimuli, such as the presence of certain individuals, on the rate of three different contact call types. To calculate mean individual-level rates, we measured the group-level rate and divided it by the number of birds in the aviary. We found that the individual-level rate of the most common call types increased with a greater group size, the opposite pattern to what would be expected if birds were maintaining a specific group-level rate. Vegetation density did not affect any call rate. However, individual-level rates of all call types decreased when birds were in subgroups with individuals of differing dominance status, and the rate of some call types increased when birds were with affiliated individuals. Our results do not support the hypothesis that contact calls are related to habitat structure or immediate predation risk. Rather, they appear to have a social function, used for communication within or between groups depending on the call type. Increases in call rates could recruit affiliated individuals, whereas subordinates could withhold calls so that dominants are unable to locate them, leading to fluctuations in contact calling in different social contexts.

Behavioral Strategies and the Spatial Pattern Formation of Nesting.

AbstractNesting in dense aggregations is common in central place foragers, such as group-living birds and insects. Both environmental heterogeneity and behavioral interactions are known to induce clustering of nests, but their relative importance remains unclear. We developed an individual-based model that simulated the spatial organization of nest building in a gregarious digger wasp, Bembix rostrata. This process-based model integrates environmental suitability, as derived from a microhabitat model, and relevant behavioral mechanisms related to local site fidelity and conspecific attraction. The drivers behind the nesting were determined by means of inverse modeling in which the emerging spatial and network patterns from simulations were compared with those observed in the field. Models with individual differences in behavior that include the simultaneous effect of a weak environmental cue and strong behavioral mechanisms yielded the best fit to the field data. The nest pattern formation of a central place foraging insect cannot be considered as the sum of environmental and behavioral mechanisms. We demonstrate the use of inverse modeling to understand complex processes that underlie nest aggregation in nature.

Seasonality impacts collective movements in a wild group-living bird

BackgroundA challenge faced by animals living in groups with stable long-term membership is to effectively coordinate their actions and maintain cohesion. However, as seasonal conditions alter the distribution of resources across a landscape, they can change the priority of group members and require groups to adapt and respond collectively across changing contexts. Little is known about how stable group-living animals collectively modify their movement behaviour in response to environment changes, such as those induced by seasonality. Further, it remains unclear how environment-induced changes in group-level movement behaviours might scale up to affect population-level properties, such as a population’s footprint.MethodsHere we studied the collective movement of each distinct social group in a population of vulturine guineafowl (Acryllium vulturinum), a largely terrestrial and non-territorial bird. We used high-resolution GPS tracking of group members over 22 months, combined with continuous time movement models, to capture how and where groups moved under varying conditions, driven by seasonality and drought.ResultsGroups used larger areas, travelled longer distances, and moved to new places more often during drier seasons, causing a three-fold increase in the area used at the population level when conditions turned to drought. By contrast, groups used smaller areas with more regular movements during wetter seasons.ConclusionsThe consistent changes in collective outcomes we observed in response to different environments raise questions about the role of collective behaviour in facilitating, or impeding, the capacity for individuals to respond to novel environmental conditions. As droughts will be occurring more often under climate change, some group living animals may have to respond to them by expressing dramatic shifts in their regular movement patterns. These shifts can have consequences on their ranging behaviours that can scale up to alter the footprints of animal populations.

Forest management affects seasonal source-sink dynamics in a territorial, group-living bird

The persistence of wildlife populations is under threat as a consequence of human activities, which are degrading natural ecosystems. Commercial forestry is the greatest threat to biodiversity in boreal forests. Forestry practices have degraded most available habitat, threatening the persistence of natural populations. Understanding population responses is, therefore, critical for their conservation. Population viability analyses are effective tools to predict population persistence under forestry management. However, quantifying the mechanisms driving population responses is complex as population dynamics vary temporally and spatially. Metapopulation dynamics are governed by local dynamics and spatial factors, potentially mediating the impacts of forestry e.g., through dispersal. Here, we performed a seasonal, spatially explicit population viability analysis, using long-term data from a group-living territorial bird (Siberian jay, Perisoreus infaustus). We quantified the effects of forest management on metapopulation dynamics, via forest type-specific demography and spatially explicit dispersal, and how forestry impacted the stability of metapopulation dynamics. Forestry reduced metapopulation growth and stability, through negative effects on reproduction and survival. Territories in higher quality natural forest contributed more to metapopulation dynamics than managed forests, largely through demographic processes rather than dispersal. Metapopulation dynamics in managed forest were also less resilient to disturbances and consequently, may be more vulnerable to environmental change. Seasonal differences in source-sink dynamics observed in managed forest, but not natural forests, were caused by associated seasonal differences in dispersal. As shown here, capturing seasonal source-sink dynamics allows us to predict population persistence under human disturbance and to provide targeted conservation recommendations.

Who do you trust? Wild birds use social knowledge to avoid being deceived.

Many species give deceptive warning calls, enabled by the high risk of ignoring them. In Siberian jays, a territorial, group-living bird, individuals give warning calls toward perched predators and mob them. However, intruding neighbors can emit these warning calls in the absence of predators to access food, but breeders often ignore these calls. Playback field experiments show that breeders flee sooner and return later after warning calls of former group members than those of neighbors or unknown individuals. Thus, breeders respond appropriately only to warning calls of previous cooperation partners. This mechanism facilitates the evolution and maintenance of communication vulnerable to deceptive signaling. This conclusion also applies to human language because of its cooperative nature and thus, its vulnerability to deception.

Larger group sizes facilitate the emergence and spread of innovations in a group-living bird

The benefits of group living have traditionally been attributed to risk dilution or the efficient exploitation of resources; individuals in social groups may therefore benefit from access to valuable information. If sociality facilitates access to information, then individuals in larger groups may be predicted to solve novel problems faster than individuals in smaller groups. Additionally, larger group sizes may facilitate the subsequent spread of innovations within animal groups, as has been proposed for human societies. We presented a novel foraging task (where a food reward could be accessed by pushing a self-shutting sliding door) to 16 groups of wild, cooperatively breeding Australian magpies, Cracticus tibicen dorsalis, ranging in size from two to 11 individuals. We found a nonlinear decline in the time taken for the innovative behaviour to emerge with increasing group size, and social information use facilitated the transmission of novel behaviour, with it spreading more quickly in larger than smaller groups. This study provides important evidence for a nonlinear relationship between group size and the emergence of innovation (and its subsequent transmission) in a wild population of animals. Further work investigating the scope and strength of group size–innovation relationships, and the mechanisms underpinning them, will help us understand the potential advantages of living in larger social groups.

Wild acorn woodpeckers recognize associations between individuals in other groups.

According to the social intelligence hypothesis, understanding the cognitive demands of the social environment is key to understanding the evolution of intelligence. Many important socio-cognitive abilities, however, have primarily been studied in a narrow subset of the social environment-within-group social interactions-despite the fact that between-group social interactions often have a substantial effect on fitness. In particular, triadic awareness (knowledge about the relationships and associations between others) is critical for navigating many types of complex social interactions, yet no existing study has investigated whether wild animals can track associations between members of other social groups. We investigated inter-group triadic awareness in wild acorn woodpeckers (Melanerpes formicivorus), a socially complex group-living bird. We presented woodpeckers with socially incongruous playbacks that simulated two outsiders from different groups calling together, and socially congruous playbacks that simulated two outsiders from the same group calling together. Subjects responded more quickly to the incongruous playbacks, suggesting that they were aware that the callers belonged to two different groups. This study provides the first demonstration that animals can recognize associations between members of other groups under natural circumstances, and highlights the importance of considering how inter-group social selection pressures may influence the evolution of cognition.

The interacting effects of forestry and climate change on the demography of a group-living bird population.

Anthropogenic degradation of natural habitats is a global driver of wildlife population declines. Local population responses to such environmental perturbations are generally well understood, but in socially structured populations, interactions between environmental and social factors may influence population responses. Thus, understanding how habitat degradation affects the dynamics of these populations requires simultaneous consideration of social and environmental mechanisms underlying demographic responses. Here we investigated the effect of habitat degradation through commercial forestry on spatiotemporal dynamics of a group-living bird, the Siberian jay, Perisoreus infaustus, in boreal forests of northern Sweden. We assessed the interacting effects of forestry, climate and population density on stage-specific, seasonal life-history rates and population dynamics, using long-term, individual-based demographic data from 70 territories in natural and managed forests. Stage-specific survival and reproductive rates, and consequently population growth, were lower in managed forests than in natural forests. Population growth was most sensitive to breeder survival and was more sensitive to early dispersing juveniles than those delaying dispersal. Forestry decreased population growth in managed forests by reducing reproductive success and breeder survival. Increased snow depth improved winter survival, and warmer spring temperatures enhanced reproductive success, particularly in natural forests. Population growth was stable in natural forests but it was declining in managed forests, and this difference accelerated under forecasted climate scenarios. Thus, climatic change could exacerbate the rate of forestry-induced population decline through reduced snow cover in our study species, and in other species with similar life-history characteristics and habitat requirements.

Coupling random inputs for parameter estimation in complex models

Complex stochastic models, such as individual-based models, are becoming increasingly popular. However this complexity can often mean that the likelihood is intractable. Performing parameter estimation on the model can then be difficult. One way of doing this when the complex model is relatively quick to simulate from is approximate Bayesian computation (ABC). Rejection-ABC algorithm is not always efficient so numerous other algorithms have been proposed. One such method is ABC with Markov chain Monte Carlo (ABC---MCMC). Unfortunately for some models this method does not perform well and some alternatives have been proposed including the fsMCMC algorithm (Neal and Huang, in: Scand J Stat 42:378---396, 2015) that explores the random inputs space as well unknown model parameters. In this paper we extend the fsMCMC algorithm and take advantage of the joint parameter and random input space in order to get better mixing of the Markov Chain. We also introduce a Gibbs step that conditions on the current accepted model and allows the parameters to move as well as the random inputs conditional on this accepted model. We show empirically that this improves the efficiency of the ABC---MCMC algorithm on a queuing model and an individual-based model of the group-living bird, the woodhoopoe.

Strategic use of allopreening in family-living wrens

A prominent interaction in the lives of social mammals is allogrooming. Individuals allogroom strategically and preferentially, grooming high-quality individuals that control access to resources. This results in distinct patterns of allogrooming within social groups, such that some individuals are groomed more than the others, some dyads express symmetry in allogrooming, and others exchange allogrooming for other benefits. Allopreening, the avian equivalent of allogrooming, occurs commonly in group-living birds, providing the opportunity to test whether social birds also use allopreening strategically in their social relationships. I examined this hypothesis in family-living buff-breasted wrens (Cantorchilus leucotis) by examining allopreening initiation, reciprocation and rate during interactions involving breeding adults and offspring. Buff-breasted wrens exhibit a nuclear family structure in which pairs display long-term partnerships, and juveniles delay dispersing from natal territories for about 10 months. Allopreening was symmetrical between breeding partners: males and females who initiated and reciprocated allopreening of partners with similar frequency, and within reciprocated bouts, allopreening was time-matched. Pairs in which allopreening was not observed were more likely to divorce, but allopreening patterns did not change in successive years of partnerships. Parent–offspring allopreening, by contrast, was asymmetrical in pattern; parents initiated allopreening more than their offspring, and reciprocation never occurred. The different patterns of allopreening observed within buff-breasted wren families suggest the strategic use of allopreening, possibly for the maintenance of long-term partnerships and in exchange for social benefits from offspring.

Breeding synchrony through social stimulation in a spatially segregated population of European starlings

Breeding synchrony occurs when individuals simultaneously perform portions of the reproductive cycle more often than expected by chance. We tested for the role of social stimulation in driving breeding synchrony of European starlings, Sturnus vulgaris, in a spatially-structured population in New Zealand over 11 years. Natural spreads in laying in groups were consistently tighter relative to aggregations of individuals chosen at random. Rather than a decrease in synchrony with increasing density, as would be expected if competition reduces breeding condition, we found that birds in dense breeding areas showed tighter synchrony. As the number of active nests within 500 m of a breeding location (i.e. human-constructed concrete bunkers) increased, the spread in clutch initiation date decreased. There was a correlation between synchrony and fledging success; less synchronous locations had lower fledging success. These results indicate that local and larger-scale levels of social stimulation may help coordinate reproductive timing in group-living birds. While the link between fitness and fledging success may not be causal, our results suggest that factors driving synchrony may also be related to individual quality and reproductive success.

Population regulation in group-living birds: predictive models of the Seychelles warbler.

A major challenge for population ecology is to predict population responses to novel conditions, such as habitat loss. This frequently involves understanding dispersal decisions, in terms of their consequences for fitness. However, this approach requires detailed data, and is thus often inappropriate for urgent problems on poorly known species. This may be resolved by developing a predictive framework based on well-studied species, for applying to those that are less well understood. Population size, group sizes and habitat occupancy of the Seychelles warbler (Acrocephalus sechellensis) can be predicted by determining the evolutionary stable dispersal strategy. For densities near to demographic equilibrium, regulation results from the combined effects of non-breeding and use of sink habitats. In the Seychelles warbler, resident male non-breeders compete for breeding vacancies on neighbouring territories. The resulting kin competition is a key process for predicting the observed balance between regulation by non-breeding and regulation by sink use. Family groups, in which offspring delay dispersal, hoping to fill a vacancy on a local territory, are common among group-living species. This suggests that kin competition may frequently play a central role in the population regulation of socially complex species. Although all the model variants considered are complex, predictions are shown to be insensitive to a range of simplifications, illustrating that, despite significant evolutionary import at the individual level, some behaviour can be unimportant when considering population level questions. Identifying which behavioural strategies have significant demographic consequences is key to the further development of population models based on fitness maximizing behaviour.

Evidence for long-distance dispersal in a sedentary passerine, Gymnorhina tibicen (Artamidae)

Australian magpies (Gymnorhina tibicen) are group-living birds found across much of mainland Australia. Adults commonly remain in a breeding territory until death. Young of the year either remain on the natal (birth) site or are forced by their parents to disperse. Observational studies in south-eastern Australia suggest that most dispersing juveniles settle within 7 km of their natal territory. Therefore, despite potential for considerable gene flow (via flight), social organization predisposes magpies towards local population structuring. In this study, we measured genetic variation at both nuclear (allozyme) and mitochondrial loci and found evidence of substantial gene flow over very large distances (up to 1599 km). Thus, some juvenile magpies may disperse much greater distances than was previously thought. For mtDNA, geographic and genetic distance were strongly correlated, consistent with a pattern of isolation by distance. Therefore, although female gene flow is substantial it is apparently geographically restricted over large distances, in approximately a stepping-stone fashion. We conclude that a strong relationship between gene flow and geographic distance can develop even over large distances if populations have experienced no major historical disturbances to gene flow.

Characteristic Features of Group Living Life of the Azure-winged Magpie Cyanopica cyana

1) The social system of Azure-winged Magpies Cyanopica cyana was studied at Hizume, Nagano City, from 1983 to 1984, and at Kawanakajima, Nagano City, from 1962 to 1965 and inwinter of 1983.2) The six months from April to September were defined as the breeding season, because pairs with or without their fledglings were observed during this period.3) In the non-breeding season, flocks consisting of about 20 magpies were formed and the size of each flock was stable throughout the day.4) In the non-breeding season, all members of a flock always roosted together. In winter and spring, different flocks sometimes roosted together without any antagonistic behavior.5) The home range of Hizume flock was 1.2 km long and 0.9 km wide, and included 18 units of premises consisting of some houses and surrounding woods. Magpies spent longer time in premises with closed woods than those with sparse woods.6) Home ranges of flocks seemed to coincide nearly with territories, because the home ranges overlapped little with each other and chases were observed around the boundaries of the ranges.7) Flocks broke up into some pairs, non-breeding adults and subadults in the breeding season. At Hizume, nests of pairs belonging to the same winter flock were built 10-20 m apart from each other in a unit of premises, and the flock members visited the nests without any antagonistic behavior.8) The fledging rate was nearly 50%. Families consisting of pairs and their fledglings were observed from mid-May to mid-September. Family members moved and roosted together in the nesting woods about one month after fledging.9) There was no antagonistic behavior between families of the same flock. Some adults which had failed in breeding joined families, and one subadult fed fledglings as a helper.10) Even in the breeding season, not only pairs but also some non-breeding adults and subadults of the same flock sometimes moved or roosted together. They showed defensive behavior against conspecific intruders or natural enemies such as domestic cats and crows.11) As a conclusion of comparison among group-living birds, the Azure-winged Magpie was considered as a typical species which formed stable flocks throughout the year.