Alarm communication networks as a driver of community structure in African savannah herbivores.

Social information networks have the potential to shape the spatial structure of ecological communities by promoting the formation of mixed-species groups. However, what actually drives social affinity between species in the wild will depend on the characteristics of the species available to group. Here we first present an agent-based model that predicts trait-related survival benefits from mixed-species group formation in a multi-species community and we then test the model predictions in a community-wide field study of African savannah herbivores using multi-layered network analysis. We reveal benefits from information transfer about predators as a key determinant of mixed-species group formation, and that dilution benefits alone are not enough to explain patterns in interspecific sociality. The findings highlight the limitations of classical ecological approaches focusing only on direct trophic interactions when analysing community structure and suggest that declines in species occupying central social network positions, such as key informants, can have significant repercussions throughout communities.

Using social network analysis of mixed-species groups in African savannah herbivores to assess how community structure responds to environmental change.

The dynamics of wildlife populations often depend heavily on interspecific interactions and understanding the underlying principles can be an important step in designing conservation strategies. Behavioural ecological studies can here provide useful insights into the structure and function of communities and their likely response to environmental changes. In this study of the Masai Mara herbivore community, we use a social network approach to investigate social affinities between species and how these change over the year in response to seasonal changes in ecological conditions. We find that even though social networks were correlated across different ecological conditions, for half the species dyads in the community, the strength of social affinities responded to changes in rainfall and/or the presence of migratory wildebeest. Several species consequentially adopted more or less central positions in the network depending on the ecological conditions. The findings point out interspecific social links that are likely to be attenuated or strengthened as a consequence of human-induced environmental changes and therefore call for particular attention from conservation managers. The eco-evolutionary ramifications of the perturbations of social affinities still require further study. This article is part of the theme issue 'Linking behaviour to dynamics of populations and communities: application of novel approaches in behavioural ecology to conservation'.

Linking behaviour to dynamics of populations and communities: application of novel approaches in behavioural ecology to conservation.

The impact of environmental change on the reproduction and survival of wildlife is often behaviourally mediated, placing behavioural ecology in a central position to quantify population- and community-level consequences of anthropogenic threats to biodiversity. This theme issue demonstrates how recent conceptual and methodological advances in the discipline are applied to inform conservation. The issue highlights how the focus in behavioural ecology on understanding variation in behaviour between individuals, rather than just measuring the population mean, is critical to explaining demographic stochasticity and thereby reducing fuzziness of population models. The contributions also show the importance of knowing the mechanisms by which behaviour is achieved, i.e. the role of learning, reasoning and instincts, in order to understand how behaviours change in human-modified environments, where their function is less likely to be adaptive. More recent work has thus abandoned the 'adaptationist' paradigm of early behavioural ecology and increasingly measures evolutionary processes directly by quantifying selection gradients and phenotypic plasticity. To support quantitative predictions at the population and community levels, a rich arsenal of modelling techniques has developed, and interdisciplinary approaches show promising prospects for predicting the effectiveness of alternative management options, with the social sciences, movement ecology and epidemiology particularly pertinent. The theme issue furthermore explores the relevance of behaviour for global threat assessment, and practical advice is given as to how behavioural ecologists can augment their conservation impact by carefully selecting and promoting their study systems, and increasing their engagement with local communities, natural resource managers and policy-makers. Its aim to uncover the nuts and bolts of how natural systems work positions behavioural ecology squarely in the heart of conservation biology, where its perspective offers an all-important complement to more descriptive 'big-picture' approaches to priority setting. This article is part of the theme issue 'Linking behaviour to dynamics of populations and communities: application of novel approaches in behavioural ecology to conservation'.

Multiple adaptive and non-adaptive processes determine responsiveness to heterospecific alarm calls in African savannah herbivores.

Heterospecific alarm calls may provide crucial survival benefits shaping animal behaviour. Multispecies studies can disentangle the relative importance of the various processes determining these benefits, but previous studies have included too few species for alternative hypotheses to be tested quantitatively in a comprehensive analysis. In a community-wide study of African savannah herbivores, we here, for the first time to our knowledge, partition alarm responses according to distinct aspects of the signaller-receiver relationship and thereby uncover the impact of several concurrent adaptive and non-adaptive processes. Stronger responses were found to callers who were vulnerable to similar predators and who were more consistent in denoting the presence of predators of the receiver. Moreover, alarm calls resembling those of conspecifics elicited stronger responses, pointing to sensory constraints, and increased responsiveness to more abundant callers indicated a role of learning. Finally, responses were stronger in risky environments. Our findings suggest that mammals can respond adaptively to variation in the information provided by heterospecific callers but within the constraints imposed by a sensory bias towards conspecific calls and reduced learning of less familiar calls. The study thereby provides new insights central to understanding the ecological consequences of interspecific communication networks in natural communities.

On the Challenge of Interpreting Census Data: Insights from a Study of an Endangered Pinniped.

Population monitoring is vital for conservation and management. However, simple counts of animals can be misleading and this problem is exacerbated in seals (pinnipeds) where individuals spend much time foraging away from colonies. We analyzed a 13-year-series of census data of Galapagos sea lions (Zalophus wollebaeki) from the colony of Caamaño, an islet in the center of the Galapagos archipelago where a large proportion of animals was individually marked. Based on regular resighting efforts during the cold, reproductive (cold-R; August to January) and the warm, non-reproductive (warm-nR; February to May) season, we document changes in numbers for different sex and age classes. During the cold-R season the number of adults increased as the number of newborn pups increased. Numbers were larger in the morning and evening than around mid-day and not significantly influenced by tide levels. More adults frequented the colony during the warm-nR season than the cold-R season. Raw counts suggested a decline in numbers over the 13 years, but Lincoln-Petersen (LP-) estimates (assuming a closed population) did not support that conclusion. Raw counts and LP estimates were not significantly correlated, demonstrating the overwhelming importance of variability in attendance patterns of individuals. The probability of observing a given adult in the colony varied between 16% (mean for cold-R season) and 23% (warm-nR season) and may be much less for independent 2 to 4 year olds. Dependent juveniles (up to the age of about 2 years) are observed much more frequently ashore (35% during the cold-R and 50% during the warm-nR seasons). Simple counts underestimate real population size by a factor of 4-6 and may lead to erroneous conclusions about trends in population size.

Offspring Hormones Reflect the Maternal Prenatal Social Environment: Potential for Foetal Programming?

Females of many species adaptively program their offspring to predictable environmental conditions, a process that is often mediated by hormones. Laboratory studies have shown, for instance, that social density affects levels of maternal cortisol and testosterone, leading to fitness-relevant changes in offspring physiology and behaviour. However, the effects of social density remain poorly understood in natural populations due to the difficulty of disentangling confounding influences such as climatic variation and food availability. Colonially breeding marine mammals offer a unique opportunity to study maternal effects in response to variable colony densities under similar ecological conditions. We therefore quantified maternal and offspring hormone levels in 84 Antarctic fur seals (Arctocephalus gazella) from two closely neighbouring colonies of contrasting density. Hair samples were used as they integrate hormone levels over several weeks or months and therefore represent in utero conditions during foetal development. We found significantly higher levels of cortisol and testosterone (both P < 0.001) in mothers from the high density colony, reflecting a more stressful and competitive environment. In addition, offspring testosterone showed a significant positive correlation with maternal cortisol (P < 0.05). Although further work is needed to elucidate the potential consequences for offspring fitness, these findings raise the intriguing possibility that adaptive foetal programming might occur in fur seals in response to the maternal social environment. They also lend support to the idea that hormonally mediated maternal effects may depend more strongly on the maternal regulation of androgen rather than cortisol levels.

Applicability of single-camera photogrammetry to determine body dimensions of pinnipeds: Galapagos sea lions as an example.

Morphological features correlate with many life history traits and are therefore of high interest to behavioral and evolutionary biologists. Photogrammetry provides a useful tool to collect morphological data from species for which measurements are otherwise difficult to obtain. This method reduces disturbance and avoids capture stress. Using the Galapagos sea lion (Zalophus wollebaeki) as a model system, we tested the applicability of single-camera photogrammetry in combination with laser distance measurement to estimate morphological traits which may vary with an animal's body position. We assessed whether linear morphological traits estimated by photogrammetry can be used to estimate body length and mass. We show that accurate estimates of body length (males: ±2.0%, females: ±2.6%) and reliable estimates of body mass are possible (males: ±6.8%, females: 14.5%). Furthermore, we developed correction factors that allow the use of animal photos that diverge somewhat from a flat-out position. The product of estimated body length and girth produced sufficiently reliable estimates of mass to categorize individuals into 10 kg-classes of body mass. Data of individuals repeatedly photographed within one season suggested relatively low measurement errors (body length: 2.9%, body mass: 8.1%). In order to develop accurate sex- and age-specific correction factors, a sufficient number of individuals from both sexes and from all desired age classes have to be captured for baseline measurements. Given proper validation, this method provides an excellent opportunity to collect morphological data for large numbers of individuals with minimal disturbance.

Sources of acoustic variation: implications for production specificity and call categorization in chacma baboon (Papio ursinus) grunts.

Elucidating the information content of vocal signals is fundamental to the understanding of animal communication. Acoustically distinct calls produced in specific contexts allow listeners to predict future events and choose adequate responses. However, the vocal repertoires of most terrestrial mammals consist of a limited number of call types that vary within and between categories. These "graded signaling systems" are thought to be rich in information, at the cost of increasing uncertainty regarding call categorization. In this study, patterns of acoustic variation in grunts of wild chacma baboons (Papio ursinus) were assessed in relation to different contexts, callers' arousal, the presence of listeners, and individual identity. Although overall production specificity was low, and sensitive to the number of contexts under consideration, grunts given in three contexts could be statistically distinguished from each other. Contextual differences remained when controlling for caller arousal, suggesting that these differences cannot be explained by variation in arousal. No audience effect was detected, but individual identity was found to have an influence on acoustic structure. Overall, these results support the view that, in comparison to other signaling systems associated with hazardous conditions, lower production specificity might evolve under relaxed circumstances where unambiguous signaling is less important.