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Even in well-studied organisms, it is often challenging to uncover the social and environmental determinants of fitness. Typically, fitness is determined by a variety of factors that act in concert, thus forming complex networks of causal relationships. Moreover, even strong correlations between social and environmental conditions and fitness components may not be indicative of direct causal links, as the measured variables may be driven by unmeasured (or unmeasurable) causal factors. Standard statistical approaches, like multiple regression analyses, are not suited for disentangling such complex causal relationships. Here, we apply structural equation modeling (SEM), a technique that is specifically designed to reveal causal relationships between variables, and which also allows to include hypothetical causal factors. Therefore, SEM seems ideally suited for comparing alternative hypotheses on how fitness differences arise from differences in social and environmental factors. We apply SEM to a rich data set collected in a long-term study on the Seychelles warbler (Acrocephalus sechellensis), a bird species with facultatively cooperative breeding and a high rate of extra-group paternity. Our analysis reveals that the presence of helpers has a positive effect on the reproductive output of both female and male breeders. In contrast, per capita food availability does not affect reproductive output. Our analysis does not confirm earlier suggestions on other species that the presence of helpers has a negative effect on the reproductive output of male breeders. As such, both female and male breeders should tolerate helpers in their territories, irrespective of food availability.

Migratory birds must make complex decisions to use wind to their advantage during flight and increasing flight performance is particularly important while crossing ecological barriers. Age-related differences in how birds deal with wind have suggested experience improves necessary skills in gaining positive wind support. However, differences in wind support between age groups over ecological barriers have rarely been tested, and our understanding of how birds acquire related skills is lacking. We compared wind support achieved by adult and subadult Caspian terns, Hydroprogne caspia, during southward and northward crossings of the Sahara Desert by quantifying air-to-groundspeed ratios (AGR). We also tested possible underlying causes of lower subadult wind support in comparison to adults by calculating optimal AGR altitudes and fitting step selection functions in response to wind direction and speed. We found no difference between age groups in autumn, when young were flying with adults, but subadults had lower wind support during their first solo northward crossings. Adults departed northwards from wintering areas earlier in the year and encountered more favourable wind conditions than subadults, yet both age groups made similar movement decisions in relation to wind. Consequently, differences in performance are better explained by timing of passage rather than movement skills. Our findings highlight the influence of wind seasonality over the Sahara on migratory behaviour and raise questions about the evolution and ontogeny of migratory timing in relation to wind patterns and other factors that may determine departure decisions.(c) 2022 The Author(s). Published by Elsevier Ltd on behalf of The Association for the Study of Animal Behaviour. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/lice nses/by-nc-nd/4.0/).

Within socially monogamous breeding systems, levels of extra-pair paternity can vary not only between species, populations, and individuals, but also across time. Uncovering how different extrinsic conditions (ecological, demographic, and social) influence this behavior will help shed light on the factors driving its evolution. Here, we simultaneously address multiple socio-ecological conditions potentially influencing female infidelity in a natural population of the cooperatively breeding Seychelles warbler, Acrocephalus sechellensis. Our contained study population has been monitored for more than 25 years, enabling us to capture variation in socio-ecological conditions between individuals and across time and to accurately assign parentage. We test hypotheses predicting the influence of territory quality, breeding density and synchrony, group size and composition (number and sex of subordinates), and inbreeding avoidance on female infidelity. We find that a larger group size promotes the likelihood of extra-pair paternity in offspring from both dominant and subordinate females, but this paternity is almost always gained by dominant males from outside the group (not by subordinate males within the group). Higher relatedness between a mother and the dominant male in her group also results in more extra-pair paternity—but only for subordinate females—and this does not prevent inbreeding occurring in this population. Our findings highlight the role of social conditions favoring infidelity and contribute toward understanding the evolution of this enigmatic behavior., Infidelity is common in animals that form breeding pairs. Why this behavior occurs—and varies in frequency between species, populations, and individuals—remains unresolved. We investigate the role of social, demographic, and ecological factors on the occurrence of infidelity in Seychelles warblers. In this species, dominant pairs are often joined by subordinates to form breeding groups. We find that social conditions—group size and pair relatedness—promote female infidelity, but this infidelity almost always occurs with extra-group males.

Across socially monogamous species – where individuals form pairs to breed – infidelity is a common occurrence. However, the forces underlying its evolution are still unclear. In this thesis, I investigated several potential drivers of infidelity in the Seychelles warbler. This is a socially monogamous species with high rates of infidelity and individuals living in either pairs or groups (consisting of a breeding pair and subordinates that can help rear offspring). Moreover, I addressed a consequence of infidelity in the Seychelles warbler, i.e. the contribution of extra-group paternity (EGP) to the variance in male reproductive success. I showed that a social condition – group size – was positively associated to female infidelity. Moreover, patterns of male and female extra-group reproduction were age-dependent (changing within-individuals with age). In males, such age-related changes probably altered the age-specific contribution of EGP to the variance in reproductive success. Upon addressing whether infidelity enabled females to gain indirect genetic benefits – i.e. to increase the genetic quality of their offspring – I provided limited evidence that females use extra-group reproduction to avoid inbreeding (indirect non-additive genetic benefits). Further, I found that the age-related patterns of infidelity did not offer evidence that females use extra-group reproduction to acquire high-quality paternal alleles in offspring (indirect additive genetic benefits). Moreover, I showed that heritability of female extra-group reproduction was moderately low. These findings suggest that infidelity is unlikely to evolve via indirect additive genetic benefits in the Seychelles warbler. Other mechanisms may be at play in the evolution of this enigmatic mating strategy.

Extra-pair paternity (EPP) is often linked to male age in socially monogamous vertebrates; that is, older males are more likely to gain EPP and less likely to be cuckolded. However, whether this occurs because males improve at gaining paternity as they grow older, or because “higher quality” males that live longer are preferred by females, has rarely been tested, despite being central to our understanding of the evolutionary drivers of female infidelity. Moreover, how extra-pair reproduction changes with age within females has received even less attention. Using 18 years of longitudinal data from an individually marked population of Seychelles warblers (Acrocephalus sechellensis), we found considerable within-individual changes in extra-pair reproduction in both sexes: an early-life increase and a late-life decline. Furthermore, males were cuckolded less as they aged. Our results indicate that in this species age-related patterns of extra-pair reproduction are determined by within-individual changes with age, rather than differences among individuals in longevity. These results challenge the hypothesis—based on longevity reflecting intrinsic quality—that the association between male age and EPP is due to females seeking high-quality paternal genes for offspring. Importantly, EPP accounted for up to half of male reproductive success, emphasizing the male fitness benefits of this reproductive strategy. Finally, the occurrence of post-peak declines in extra-pair reproduction provides explicit evidence of senescence in infidelity in both males and females.

Inhibitory control enables subjects to quickly react to unexpectedly changing external demands. We assessed the ability of young (8 weeks old) pheasants Phasianus colchicus to exert inhibitory control in a novel response-inhibition task that required subjects to adjust their movement in space in pursuit of a reward across changing target locations. The difference in latencies between trials in which the target location did and did not change, the distance travelled towards the initially indicated location after a change occurred, and the change-signal reaction time provided a consistent measure that could be indicative of a pheasant’s inhibitory control. Between individuals, there was a great variability in these measures; these differences were not correlated with motivation either to access the reward or participate in the test. However, individuals that were slower to reach rewards in trials when the target did not change exhibited evidence of stronger inhibitory control, as did males and small individuals. This novel test paradigm offers a potential assay of inhibitory control that utilises a natural feature of an animal’s behavioural repertoire, likely common to a wide range of species, specifically their ability to rapidly alter their trajectory when reward locations switch. Electronic supplementary material The online version of this article (doi:10.1007/s10071-017-1120-8) contains supplementary material, which is available to authorized users.