Extra-pair parentage and personality in a cooperatively breeding bird

Why so much variation in extra-pair parentage occurs within and among populations remains unclear. Often the fitness costs and benefits of extra-pair parentage are hypothesised to explain its occurrence; therefore, linking extra-pair parentage with traits such as personality (behavioural traits that can be heritable and affect reproductive behaviour) may help our understanding. Here, we investigate whether reproductive outcomes and success are associated with exploratory behaviour in a natural population of cooperatively breeding Seychelles warblers (Acrocephalus sechellensis) on Cousin Island. Exploratory behaviour correlates positively with traits such as risk-taking behaviour and activity in other wild bird species and might promote extra-pair mating by increasing the rate at which potential extra-pair partners are encountered. We therefore predicted that fast-exploring individuals would have more extra-pair offspring. There is also a potential trade-off between pursuing extra-pair parentage and mate guarding in males. We therefore also predicted that fast-exploring males would be more likely to pursue extra-pair parentage and that this would increase the propensity of their mate to gain extra-pair parentage. We found that neither the total number of offspring nor the number of extra-pair offspring were associated with a male’s or female’s exploratory behaviour. However, there was a small but significant propensity for females to have extra-pair fertilisations in pairs that were behaviourally disassortative. Overall, we conclude that, due to the small effect size, the association between exploratory behaviour and extra-pair paternity is unlikely to be biologically relevant. Significance statement: True genetic monogamy is rare, even in socially monogamous systems, and multiple factors, such as behaviour, social structure, morphology and physiology, determined by the biological system can cause variation in extra-pair parentage (EPP). Therefore, investigating the inherent differences in these factors among individuals could be informative. We investigated whether reproductive outcomes/success are associated with differences in the propensity to explore novel environments/objects in a promiscuous, island-dwelling cooperatively breeding bird, the Seychelles warbler. Our results showed that exploratory behaviour was not associated with the number of offspring produced by an individual, and thus the long-term fitness consequences of different exploratory tendencies did not differ. We also found that the propensity to engage in EPP in females was higher in dissimilar behavioural pairs, but due to the small effect size, we hesitate to conclude that there are personality-dependent mating outcomes in the population

Multinomial analysis of behavior: statistical methods

Behavioral ecologists frequently use observational methods, such as instantaneous scan sampling, to record the behavior of animals at discrete moments in time. We develop and apply multilevel, multinomial logistic regression models for analyzing such data. These statistical methods correspond to the multinomial character of the response variable while also accounting for the repeated observations of individuals that characterize behavioral datasets. Correlated random effects potentially reveal individual-level trade-offs across behaviors, allowing for models that reveal the extent to which individuals who regularly engage in one behavior also exhibit relatively more or less of another behavior. Using an example dataset, we demonstrate the estimation of these models using Hamiltonian Monte Carlo algorithms, as implemented in the RStan package in the R statistical environment. The supplemental files include a coding script and data that demonstrate auxiliary functions to prepare the data, estimate the models, summarize the posterior samples, and generate figures that display model predictions. We discuss possible extensions to our approach, including models with random slopes to allow individual-level behavioral strategies to vary over time and the need for models that account for temporal autocorrelation. These models can potentially be applied to a broad class of statistical analyses by behavioral ecologists, focusing on other polytomous response variables, such as behavior, habitat choice, or emotional states

Environmental factors shaping ungulate abundances in Poland

Population densities of large herbivores are determined by the diverse effects of density-dependent and independent environmental factors. In this study, we used the official 1998–2003 inventory data on ungulate numbers from 462 forest districts and 23 national parks across Poland to determine the roles of various environmental factors in shaping country-wide spatial patterns of ungulate abundances. Spatially explicit generalized additive mixed models showed that different sets of environmental variables explained 39 to 50 % of the variation in red deer Cervus elaphus, wild boar Sus scrofa, and roe deer Capreolus capreolus abundances. For all of the studied species, low forest cover and the mean January temperature were the most important factors limiting their numbers. Woodland cover above 40–50 % held the highest densities for these species. Wild boar and roe deer were more numerous in deciduous or mixed woodlands within a matrix of arable land. Furthermore, we found significant positive effects of marshes and water bodies on wild boar abundances. A juxtaposition of obtained results with ongoing environmental changes (global warming, increase in forest cover) may indicate future growth in ungulate distributions and numbers

Influences of mating group composition on the behavioral time-budget of male and female Alpine Ibex (Capra ibex) during the rut.

During the rut, polygynous ungulates gather in mixed groups of individuals of different sex and age. Group social composition, which may vary on a daily basis, is likely to have strong influences on individual's time-budget, with emerging properties at the group-level. To date, few studies have considered the influence of group composition on male and female behavioral time budget in mating groups. Focusing on a wild population of Alpine ibex, we investigated the influence of group composition (adult sex ratio, the proportion of dominant to subordinate males, and group size) on three behavioral axes obtained by Principal Components Analysis, describing male and female group time-budget. For both sexes, the first behavioral axis discerned a trade-off between grazing and standing/vigilance behavior. In females, group vigilance behavior increased with increasingly male-biased sex ratio, whereas in males, the effect of adult sex ratio on standing/vigilance behavior depended on the relative proportion of dominant males in the mating group. The second axis characterized courtship and male-male agonistic behavior in males, and moving and male-directed agonistic behavior in females. Mating group composition did not substantially influence this axis in males. However, moving and male-directed agonistic behavior increased at highly biased sex ratios (quadratic effect) in females. Finally, the third axis highlighted a trade-off between moving and lying behavior in males, and distinguished moving and female-female agonistic behavior from lying behavior in females. For males, those behaviors were influenced by a complex interaction between group size and adult sex ratio, whereas in females, moving and female-female agonistic behaviors increased in a quadratic fashion at highly biased sex ratios, and also increased with increasing group size. Our results reveal complex behavioral trade-offs depending on group composition in the Alpine ibex, and emphasize the importance of social factors in influencing behavioral time-budgets of wild ungulates during the rut

Sociospatial structure explains marked variation in brucellosis seroprevalence in an Alpine ibex population

Abstract In a context of (re)emerging infectious diseases with wildlife reservoirs, understanding how animal ecology shapes epidemiology is a key issue, particularly in wild ungulates that share pathogens with domestic herbivores and have similar food requirements. For the first time in Europe, brucellosis (Brucella melitensis), a virulent zoonosis, persisted in an Alpine ibex (Capra ibex) population and was transmitted to cattle and humans. To better understand disease dynamics, we investigated the relationships between the spatial ecology of ibex and the epidemiology of brucellosis. Combining home range overlap between 37 GPS-collared individuals and visual observations of 148 visually-marked individuals monitored during the 2013–2016 period, we showed that females were spatially segregated in at least 4 units all year round, whereas males were more prone to move between female units, in particular during the rutting period. In addition to ibex age, the spatial structure in females largely contributed to variation in seroprevalence in the whole population. These results suggest that non-sexual routes are the most likely pathways of intraspecific transmission, crucial information for management. Accounting for wildlife spatial ecology was hence decisive in improving our ability to better understand this health challenge involving a wildlife reservoir

The weather dictates the rhythms: Alpine chamois activity is well adapted to ecological conditions

Activity rhythms play an important role in the ecological relations of a species and form part of its evolutionary adaptation. Such rhythms are strongly synchronised with the annual cyclic changes by environmental stimuli, the so-called zeitgebers. Animals’ reliance on environmental stimuli is highly species-specific and allows behavioural adjustments to be made in preparation for the conditions expected in each season. We investigated daily and annual activity rhythms of Alpine chamois (Rupicapra rupicapra) by analysing high-resolution data of animals monitored with GPS collars. This first detailed field study of chamois activity showed that this species exhibited clear daily and annual activity rhythms entrained to the light-dark cycle. Chamois were more active during spring-summer and less active during winter, likely in response to the variation in the availability of food resources: both sexes appeared to maximise energy intake during the season offering the highest amount of food resources to compensate for poor food supply during winter. Daily activity was influenced by the climatic factors considered. We showed a negative correlation between daily activity and adverse climatic conditions (i.e. precipitation and, during winter, snow depth). As activity was strongly influenced by the interplay between temperature and wind throughout the year and by radiation and wind in winter, we conjectured that it was critically dependent upon animals’ thermal balance. In conclusion, our study highlighted that chamois is well adapted to the Alpine environment and seasonality but also raised questions about its ability to adapt to future climate change. Significance statement: In this study, we investigated the effects of ecological factors on Alpine chamois activity. Thanks to radio collars with accelerometers, we obtained highly detailed information on activity levels of wild animals. We found that chamois were more active during spring-summer (i.e. the seasons with the highest quality and quantity of food) and less active during winter. Our results showed that chamois activity was strongly influenced by such climatic factors as temperature, precipitation and wind speed. In winter time, chamois activity increased during the days with high solar radiation and decreased with high snow depth. Given their wide distribution in the Alps, chamois can be considered as a sentinel species of Alpine habitats. Thus, our results on the current relationship between climate and chamois behaviour may shed light on the animals’ ability to track and adapt to climate change

Determinants and Patterns of Reproductive Success in the Greater Horseshoe Bat during a Population Recovery

An individual's reproductive success will depend on traits that increase access to mates, as well as the number of mates available. In most well-studied mammals, males are the larger sex, and body size often increases success in intra-sexual contests and thus paternity. In comparison, the determinants of male success in species with reversed sexual size dimorphism (RSD) are less well understood. Greater horseshoe bats (Rhinolophus ferrumequinum) exhibit RSD and females appear to exert mate choice when they visit and copulate with males in their underground territories. Here we assessed putative determinants of reproductive success in a colony of greater horseshoe bats during a 19-year period of rapid population growth. We genotyped 1080 bats with up to 40 microsatellite loci and assigned maternity to 99.5% of pups, and paternity to 76.8% of pups. We found that in spite of RSD, paternity success correlated positively with male size, and, consistent with our previous findings, also with age. Female reproductive success, which has not previously been studied in this population, was also age-related and correlated positively with individual heterozygosity, but not with body size. Remarkable male reproductive skew was detected that initially increased steadily with population size, possibly coinciding with the saturation of suitable territories, but then levelled off suggesting an upper limit to a male's number of partners. Our results illustrate that RSD can occur alongside intense male sexual competition, that male breeding success is density-dependent, and that male and female greater horseshoe bats are subject to different selective pressures