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Individuals respond adaptively to their environment. Yet, they may differ in their responses even when confronted with the same environmental challenge. Several complementary conceptual frameworks suggest that within populations among-individual variation in life history strategies aligns not only with individuals' propensities to take risks across different situations but also with their sensitivity to variation in environmental cues. Risk-prone individuals, suggested to invest more in current reproduction at the cost of their future reproductive prospects, are predicted to be less sensitive to environmental variation than risk-averse individuals. We tested this prediction in a population of breeding blue tits, _Cyanistes caeruleus_, by confronting them with different levels of predation threat at their nests and recording their latency to resume brood provisioning after the removal of the predator stimulus. We presented taxidermic woodpecker, _Dendrocopos major_ (a common brood predator) and sparrowhawk, _Accipiter nisus_ (a common adult predator) mounts at each nest, respectively representing low and high levels of threat to adult blue tits. As a nonpredator control stimulus, we presented a blackbird, _Turdus merula_, mount. We found that on average parents took longer to resume provisioning after presentation of a sparrowhawk than a woodpecker or blackbird. Furthermore, individual latency responses across all threat levels taken together were repeatable. However, despite the population level plastic adjustment to the level of predation threat, we found no evidence for among-individual variation in plasticity. Instead, individual differences in responses were roughly maintained across all levels of threat. While our findings show that individuals differ in their level of risk taking, in the high-stakes and ecologically relevant context of predation risk during parental care, commonly held expectations about among-individual variation in behavioural plasticity were not met.

Road mortality can have a significant negative impact on amphibian population survival. Amphibian roadkill and how to avoid it are therefore widely studied, mostly on car roads but limitedly on bicycle paths. We investigated whether amphibian mortality on bicycle paths in Bargerveen, a Dutch Natura 2000 site, was affected by the number of passing cyclists and crossing amphibians. We investigated four transects on a daily basis during most of the amphibian spring migration in 2021. We counted and identified (to species level) all killed amphibians; further, we used cyclist counters and toad fences to assess the number of passing bicycles and crossing amphibians, respectively. We found 11 killed smooth newts Lissotriton vulgaris, out of 5,037 that crossed the bicycle paths. Although 11,453 anurans crossed, we found no killed anurans. The occurrence of killed smooth newts was not affected by the number of passing bicycles or crossing newts. The probability of being killed was extremely low for crossing smooth newts (0.22 %) and anurans (0 %), possibly because cyclists successfully avoid cycling over amphibians. Future monitoring should occur from early February to late November to include the complete active period of amphibians including juvenile dispersal, and across multiple successive years because amphibian numbers can vary largely between years. During our study period, however, amphibian mortality on bicycle paths in Bargerveen seems no threat to populations, despite the high numbers of cyclists. Keywords: road ecology, animal-cyclist collisions, smooth newts, anurans, conservation

Telomeres have been advocated to be important markers of biological age in evolutionary and ecological studies. Telomeres usually shorten with age, and shortening is frequently associated with environmental stressors and increased subsequent mortality. Telomere lengthening – an apparent increase in telomere length between repeated samples from the same individual – also occurs. However, the exact circumstances, and consequences, of telomere lengthening are poorly understood. Using longitudinal data from the Seychelles warbler (Acrocephalus sechellensis), we tested whether telomere lengthening – which occurs in adults of this species – is associated with specific stressors (reproductive effort, food availability, malarial infection and cooperative breeding) and predicts subsequent survival. In females, telomere shortening was observed under greater stress (i.e. low food availability, malaria infection), while telomere lengthening was observed in females experiencing lower stress (i.e. high food availability, assisted by helpers, without malaria). The telomere dynamics of males were not associated with the key stressors tested. These results indicate that, at least for females, telomere lengthening occurs in circumstances more conducive to self-maintenance. Importantly, both females and males with lengthened telomeres had improved subsequent survival relative to individuals that displayed unchanged, or shortened, telomeres – indicating that telomere lengthening is associated with individual fitness. These results demonstrate that telomere dynamics are bidirectionally responsive to the level of stress that an individual faces, but may poorly reflect the accumulation of stress over the lifetime. This study challenges how we think of telomeres as a marker of biological age.

Parental age can have considerable effects on offspring phenotypes and health. However, intergenerational effects may also have longer term effects on offspring fitness. Few studies have investigated parental age effects on offspring fitness in natural populations while also testing for sex- and environment-specific effects. Further, longitudinal parental age effects may be masked by population-level processes such as the selective disappearance of poor-quality individuals. Here, we used multigenerational data collected on individually marked Seychelles warblers (Acrocephalus sechellensis) to investigate the impact of maternal and paternal age on offspring life span and lifetime reproductive success. We found negative effects of maternal age on female offspring life span and lifetime reproductive success, which were driven by within-mother effects. There was no difference in annual reproductive output of females born to older versus younger mothers, suggesting that the differences in offspring lifetime reproductive success were driven by effects on offspring life span. In contrast, there was no association between paternal age and female offspring life span or either maternal or paternal age and male offspring life span. Lifetime reproductive success, but not annual reproductive success, of male offspring increased with maternal age, but this was driven by between-mother effects. No paternal age effects were found on female offspring lifetime reproductive success but there was a positive between-father effect on male offspring lifetime reproductive success. We did not find strong evidence for environment-dependent parental age effects. Our study provides evidence for parental age effects on the lifetime fitness of offspring and shows that such effects can be sex dependent. These results add to the growing literature indicating the importance of intergenerational effects on long-term offspring performance and highlight that these effects can be an important driver of variation in longevity and fitness in the wild.

Individual variation in telomere length is predictive of health and mortality risk across a range of species. However, the relative influence of environmental and genetic variation on individual telomere length in wild populations remains poorly understood. Heritability of telomere length has primarily been calculated using parent-offspring regression which can be confounded by shared environments. To control for confounding variables, quantitative genetic "animal models" can be used, but few studies have applied animal models in wild populations. Furthermore, parental age at conception may also influence offspring telomere length, but most studies have been cross-sectional. We investigated within- and between-parental age at conception effects and heritability of telomere length in the Seychelles warbler using measures from birds caught over 20 years and a multigenerational pedigree. We found a weak negative within-paternal age at conception effect (as fathers aged, their offspring had shorter telomeres) and a weak positive between-maternal age at conception effect (females that survived to older ages had offspring with longer telomeres). Animal models provided evidence that heritability and evolvability of telomere length were low in this population, and that variation in telomere length was not driven by early-life effects of hatch period or parental identities. Quantitative polymerase chain reaction plate had a large influence on telomere length variation and not accounting for it in the models would have underestimated heritability. Our study illustrates the need to include and account for technical variation in order to accurately estimate heritability, as well as other environmental effects, on telomere length in natural populations.

'Evolvability' - the ability to undergo adaptive evolution - is a key concept for understanding and predicting the response of biological systems to environmental change. Evolvability has various facets and is applied in many ways, easily leading to misunderstandings among researchers. To clarify matters, we first categorize the mechanisms and organismal features underlying evolvability into determinants providing variation, determinants shaping the effect of variation on fitness, and determinants shaping the selection process. Second, we stress the importance of timescale when studying evolvability. Third, we distinguish between evolvability determinants with a broad and a narrow scope. Finally, we highlight two contrasting perspectives on evolvability: general evolvability and specific evolvability. We hope that this framework facilitates communication and guides future research.

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.

Variation in developmental conditions is known to affect fitness in later life, but the mechanisms underlying this relationship remain elusive. We previously found in jackdaws Corvus monedula that larger eggs resulted in larger nestlings up to fledging. Through a cross-foster experiment of complete clutches we tested whether this association can be attributed to egg size per se, or to more proficient parents producing larger eggs and larger nestlings, with the latter effect being more or less independent of egg size. Due to other manipulations post-hatching, we primarily investigated effects on nestling mass on day 5, which we show to predict survival until fledging. We introduce a new statistical approach to compare the competing hypotheses and discuss the multiple advantages of this approach over current practice of which we report the results for comparison. We conclude that 92% of the slope of the association between egg size and nestling mass can be attributed to a direct effect of egg size. The remaining 8% of the slope can be attributed to aspects of parental chick rearing ability as reflected in egg size, but this component did not deviate significantly from zero. Intriguingly, the effect of egg size on day 5 nestling mass was steeper (1.7 g cm−3) than the effect of egg size on day 1 hatchling mass (0.7 g cm−3). Early growth is exponential, and the difference in effect size may therefore be explained by hatchlings from large eggs being further in their development at hatching. The direct effect of egg size on nestling mass raises the question what causes egg size variation in jackdaws.

1. The social environment in which individuals live affects their fitness and in turn population dynamics as a whole. Birds with facultative cooperative breeding can live in social groups with dominants, subordinate helpers that assist with the breeding of others, and subordinate non-helpers. Helping behaviour benefits dominants through increased reproductive rates and reduced extrinsic mortality, such that cooperative breeding might have evolved in response to unpredictable, harsh conditions affecting reproduction and/or survival of the dominants. Additionally, there may be different costs and benefits to both helpers and non-helpers, depending on the time-scale. For example, early-life costs might be compensated by later-life benefits. These differential effects are rarely analysed in the same study. 2. We examined whether helping behaviour affects population persistence in a stochastic environment and whether there are direct fitness consequences of different life-history tactics adopted by helpers and non-helpers. 3. We parameterised a matrix population model describing the population dynamics of female Seychelles warblers Acrocephalus sechellensis, birds that display facultative cooperative breeding. The stochastic density-dependent model is defined by a (st)age structure that includes life-history differences between helpers and non-helpers and thus can estimate the demographic mechanisms of direct benefits of helping behaviour. 4. We found that population dynamics are strongly influenced by stochastic variation in the reproductive rates of the dominants, that helping behaviour promotes population persistence and that there are only early-life differences in the direct fitness of helpers and non-helpers. 5. Through a matrix population model, we captured multiple demographic rates simultaneously and analysed their relative importance in determining population dynamics of these cooperative breeders. Disentangling early-life versus lifetime effects of individual tactics sheds new light on the costs and benefits of helping behaviour. For example, the finding that helpers and non-helpers have similar lifetime reproductive outputs and that differences in reproductive values between the two life-history tactics arise only in early life suggests that overall, helpers and non-helpers have a similar balance of costs and benefits when analysing direct benefits. We recommend analysing the consequence of different life-history tactics, during both early life and over the lifetime, as analyses of these different time frames may produce conflicting results.

Background Considerable research has focussed on the importance of bacterial communities within the vertebrate gut microbiome (GM). However, studies investigating the significance of other microbial kingdoms, such as fungi, are notably lacking, despite their potential to influence host processes. Here, we characterise the fungal GM of individuals living in a natural population of Seychelles warblers (Acrocephalus sechellensis). We evaluate the extent to which fungal GM structure is shaped by environment and host factors, including genome-wide heterozygosity and variation at key immune genes (major histocompatibility complex (MHC) and Toll-like receptor (TLR)). Importantly, we also explore the relationship between fungal GM differences and subsequent host survival. To our knowledge, this is the first time that the genetic drivers and fitness consequences of fungal GM variation have been characterised for a wild vertebrate population. Results Environmental factors, including season and territory quality, explain the largest proportion of variance in the fungal GM. In contrast, neither host age, sex, genome-wide heterozygosity, nor TLR3 genotype was associated with fungal GM differences in Seychelles warblers. However, the presence of four MHC-I alleles and one MHC-II allele was associated with changes in fungal GM alpha diversity. Changes in fungal richness ranged from between 1 and 10 sequencing variants lost or gained; in some cases, this accounted for 20% of the fungal variants carried by an individual. In addition to this, overall MHC-I allelic diversity was associated with small, but potentially important, changes in fungal GM composition. This is evidenced by the fact that fungal GM composition differed between individuals that survived or died within 7 months of being sampled. Conclusions Our results suggest that environmental factors play a primary role in shaping the fungal GM, but that components of the host immune system—specifically the MHC—may also contribute to the variation in fungal communities across individuals within wild populations. Furthermore, variation in the fungal GM can be associated with differential survival in the wild. Further work is needed to establish the causality of such relationships and, thus, the extent to which components of the GM may impact host evolution.

BackgroundThe gut microbiome (GM) can influence many biological processes in the host, impacting its health and survival, but the GM can also be influenced by the host’s traits. In vertebrates, Major Histocompatibility Complex (MHC) genes play a pivotal role in combatting pathogens and are thought to shape the host’s GM. Despite this—and the documented importance of both GM and MHC variation to individual fitness—few studies have investigated the association between the GM and MHC in the wild.ResultsWe characterised MHC class I (MHC-I), MHC class II (MHC-II) and GM variation in individuals within a natural population of the Seychelles warbler (Acrocephalus sechellensis). We determined how the diversity and composition of the GM varied with MHC characteristics, in addition to environmental factors and other host traits. Our results show that the presence of specific MHC alleles, but not MHC diversity, influences both the diversity and composition of the GM in this population. MHC-I alleles, rather than MHC-II alleles, had the greatest impact on the GM. GM diversity was negatively associated with the presence of three MHC-I alleles (Ase-ua3, Ase-ua4, Ase-ua5), and one MHC-II allele (Ase-dab4), while changes in GM composition were associated with the presence of four different MHC-I alleles (Ase-ua1, Ase-ua7, Ase-ua10, Ase-ua11). There were no associations between GM diversity andTLR3genotype, but GM diversity was positively correlated with genome-wide heterozygosity and varied with host age and field period.ConclusionsThese results suggest that components of the host’s immune system play a role in shaping the GM of wild animals. Host genotype—specifically MHC-I and to a lesser degree MHC-II variation—can modulate the GM, although whether this occurs directly, or indirectly through effects on host health, is unclear. Importantly, if immune genes can regulate host health through modulation of the microbiome, then it is plausible that the microbiome could also influence selection on immune genes. As such, host–microbiome coevolution may play a role in maintaining functional immunogenetic variation within natural vertebrate populations.

Division of labour occurs in a broad range of organisms. Yet, how division of labour can emerge in the absence of pre-existing interindividual differences is poorly understood. Using a simple but realistic model, we show that in a group of initially identical individuals, division of labour emerges spontaneously if returning foragers share part of their resources with other group members. In the absence of resource sharing, individuals follow an activity schedule of alternating between foraging and other tasks. If non-foraging individuals are fed by other individuals, their alternating activity schedule becomes interrupted, leading to task specialisation and the emergence of division of labour. Furthermore, nutritional differences between individuals reinforce division of labour. Such differences can be caused by increased metabolic rates during foraging or by dominance interactions during resource sharing. Our model proposes a plausible mechanism for the self-organised emergence of division of labour in animal groups of initially identical individuals. This mechanism could also play a role for the emergence of division of labour during the major evolutionary transitions to eusociality and multicellularity.

**Abstract** Acoustic and visual signals are well known to play important roles in social communication in birds. Growing evidence suggests that many bird species, including species of songbirds, additionally have a well-developed sense of smell. However, we are still at the beginning of understanding the potential importance of chemical communication in the social lives of birds, for example in mate choice. The secretion of the preen gland may be an important contributor to the chemical phenotype of birds. Here, we report on a first characterisation of the chemical composition of the preen gland secretion of the Blue Tit (Cyanistes caeruleus), a common songbird which is an often used model species in animal behaviour and ecology, in particular also in studies of sexual selection and (extra-pair) mate choice. We found sex differences in the composition of the preen gland secretion in breeding Blue Tits. Females further tended to have a larger number of putative compounds in their secretions compared to males. We briefly discuss the possible implications of these findings and speculate that the chemical composition of the preen gland secretion may be a sexually selected trait in Blue Tits. Our preliminary findings warrant follow-up research into the patterns of within- and among individual variation in the chemical composition of the preen gland secretion as well as the identification of the main chemical compounds involved., **Zusammenfassung**Chemische Analyse zeigt Geschlechtsunterschiede im Bürzeldrüsensekret brütender BlaumeisenSingvögel sind vor allem für ihren Gesang und ihre Gefiedermerkmale, die bei der sozialen Kommunikation eine bedeutende Rolle spielen, bekannt. Immer mehr Hinweise deuten aber darauf hin, dass viele Vogelarten, darunter auch Singvogelarten, zusätzlich einen gut entwickelten Geruchssinn haben. Welche Bedeutung die geruchliche Kommunikation im sozialen Leben der Vögel, zum Beispiel bei der Partnerwahl, spielt ist allerdings weitgehend noch unbekannt. Das Bürzeldrüsensekret könnte in diesem Zusammenhang einen wichtigen Beitrag zum chemischen Phänotyp der Vögel leisten. In unserer Studie haben wir uns die chemische Zusammensetzung des Bürzeldrüsensekrets der Blaumeise (Cyanistes caeruleus) in der Brutzeit angeschaut. Blaumeisen sind eine viel genutzte Modellart in der Verhaltensforschung und Verhaltensökologie, insbesondere auch in Studien zur sexuellen Selektion und (außerpaarigen) Partnerwahl. Anhand von chemischen Analysen mittels Gas-Chromatographie fanden wir heraus, dass es bei brütenden Blaumeisen einen Geschlechtsunterschied in der Zusammensetzung des Bürzeldrüsensekrets gibt. Weibchen neigen außerdem dazu, eine größere Anzahl von vermeintlichen Substanzen in ihren Sekreten zu haben als Männchen. Wir diskutieren hier kurz die möglichen Implikationen dieser Ergebnisse und spekulieren, dass die chemische Zusammensetzung des Bürzeldrüsensekrets ein sexuell selektiertes Merkmal bei Blaumeisen sein könnte. Unsere vorläufigen Ergebnisse rechtfertigen Folgeuntersuchungen, in denen die Identifikation der wichtigsten chemischen Verbindungen des Bürzeldrüsensekrets und die Variation innerhalb und zwischen Individuen in der chemischen Zusammensetzung des Drüsensekrets im Fokus stehen.

Sexual conflict over parental investment is widespread among species with biparental care. Studies have indicated that a high degree of behavioural similarity between the two parents can increase offspring survival; however, it remains unclear how sexual conflict over parental care is resolved. In this study, we examined whether similarity of personality traits between the two parents plays an important role in affecting the provisioning behaviour of each sex in a wild population of the chestnut thrush, Turdus rubrocanus. First, as expected, the mating pairs with more similar personality traits had higher provisioning rates than those pairs with dissimilar traits. Moreover, we found that the similarity of personality traits can modulate the sexual conflict over provisioning in this species, as both parents with more similar partners had relatively higher and less divergent provisioning rates. A partner removal experiment revealed how the sole female or male parent responded when the level of conflict over care increased (the removed partner does not provide any care). The majority of males always reduced their provisioning investment, while females’ decisions depended on the degree of similarity with their partners. Females compensated by provisioning more frequently in pairs of similar personality traits (i.e. accepting a high level of conflict), but reduced their provisioning investment in extremely dissimilar pairs. Our results promote a better understanding of the resolution of sexual conflict over provisioning and highlight the evolutionary significance of mating with similar partners based on certain personality traits.

Behavioural patterns often differ consistently across individuals and are linked to fitness. In species with biparental care, the defence behaviour of both parents can affect reproductive success through offspring survival. In addition to the intensity of defence behaviour by both pair members, the similarity in this behaviour among parents may affect offspring survival. However, few studies have investigated the relative impact of both the intensity and similarity of defence behaviour. Here, we examined nest defence behaviour of males and females during the incubation stage in an Arctic population of barnacle geese Branta leucopsis. We calculated the repeatability of defence behaviour to test whether this behaviour is consistent within individuals and investigated how it is associated with age. In addition, we investigated how daily survival rate (DSR) of the nests until hatching is associated with nest defence behaviour and age of the parents, as well as the effect of parent similarity in nest defence behaviour as an emergent trait of the pair bond. Both male and female defence behaviour were highly repeatable. The ages of both partners within breeding pairs were positively related, but age was only significantly associated with defence behaviour in females. Further, we found high similarity in defence behaviour within breeding pairs, but the similarity and intensity of defence behaviour within breeding pairs did not predict DSR. Finally, male defence behaviour positively predicted DSR, but female defence behaviour and male and female age did not. Our results suggest that nest protection is adaptive in males but behavioural similarity of pair members does not enhance nest survival, indicating behavioural similarity itself is not adaptive but rather a by-product of different effects.

Quantifying fitness is important to understand adaptive evolution. Reproductive values are useful for making fitness comparisons involving different categories of individuals, like males and females. By definition, the reproductive value of a category is the expected per capita contribution of the members of that category to the gene pool of future generations. Life history theory reveals how reproductive values can be determined from life-history parameters, but this requires matrix algebra. Recently, a more intuitive, pedigree-based method has become popular. It estimates genetic contributions of individuals to future generations by tracking their descendants down through the pedigree. We compare both methods. We implement various life-history scenarios (for which the “true” reproductive values can be calculated) in individual-based simulations, use this data to estimate reproductive values with both methods, and compare the results with the true reproductive values. We show that the pedigree-based estimation of reproductive values is either systematically biased (and hence inaccurate), or very imprecise. This holds even for simple life histories and under idealized conditions (such as complete knowledge of the pedigree). The traditional algebraic method estimates reproductive values with accuracy, and the precision of estimates can be determined by sensitivity analyses. However, these estimates can be biased as well when they are based on inadequate life-history models.

According to Fisher’s principle of equal allocation, parents should invest equal amounts of their parental resources in the production of sons and daughters, implying that the ‘cheaper’ sex should be overproduced. Most sex ratio models consider the costs per son and daughter as externally given and fixed. Here, we consider an individual-based evolutionary model where the parental investment in sons and daughters is an evolvable trait that coevolves with the primary sex ratio (PSR). We consider various scenarios and show that the evolutionary outcomes deviate strongly from Fisher’s principle. (1) When the parental care level coevolves with the PSR in the absence of linkage between the two types of loci, the evolved PSR is more strongly biased than Fisher’s principle predicts. (2) In the presence of linkage, polymorphism in care strategies and sex ratios evolves; the average PSR in the population is even more strongly biased than before. (3) If biparental care can evolve and females control the PSR (without linkage), male-only care and an unbiased PSR evolve if daughters are more costly than sons; female-only care with a strongly biased PSR evolves if sons are more costly. (4) In the presence of linkage, uniparental evolves as before, but in case of female care polymorphism evolves with an even more strongly biased PSR. (5) In some scenarios biparental care can evolve, with a PSR that is biased to the more expensive sex. We conclude that details matter when predicting the joint evolution of parental effort and the primary sex ratio.

18. Life histories Mirjam Borger1 , Jan Komdeur1, David Richardson2, Franz Weissing1 1 Groningen Institute for Evolutionary Life Sciences, University of Groningen, P.O. Box 11103, 9700 CC, Groningen, The Netherlands 2 Centre for Ecology, Evolution and Conservation, School of Biological Sciences, University of East Anglia, Norwich Research Park, NR4 7TJ Norwich, UK Quantifying fitness is important to understand adaptive evolution. Reproductive values are useful for making fitness comparisons involving different categories of individuals, like males and females. By definition, the reproductive value of a category is the expected per capita contribution of the members of that category to the gene pool of future generations. Life history theory reveals how reproductive values can be determined from life-history parameters, but this requires matrix algebra. Recently, a more intuitive, pedigree-based method has become popular. It estimates genetic contributions of individuals to future generations by tracking their descendants down through the pedigree. We compare both methods. We implement various life-history scenarios (for which the “true” reproductive values can be calculated) in individual-based simulations, use this data to estimate reproductive values with both methods, and compare the results with the true reproductive values. We show that the pedigree-based estimation of reproductive values is either systematically biased (and hence inaccurate), or very imprecise. This holds even for simple life histories and under idealized conditions (such as complete knowledge of the pedigree). The traditional algebraic method estimates reproductive values with accuracy, and the precision of estimates can be determined by sensitivity analyses. However, these estimates can be biased as well when they are based on inadequate life-history models.

Variation in types and distribution of resources promotes the evolution of competition and cooperation in animals. Small vertebrate carcasses, which are bonanza and unpredictable available for carrion-use insects are essential resources for the survival and reproduction of burying beetles N. vespilloides, a facultative communal breeding insect with parental care. Burying beetles bury carcasses for breeding, lay their eggs in or around the buried carcass and hatched larvae are fed by their parents. Resource distribution is related to variation in amount and availability. The aim of the research is to investigate whether resource distribution influences the degree of communal breeding. We expect that centralized resource distribution (i.e. large but less carcasses) promotes communal breeding, while decentralize resource distribution (i.e. small but more carcasses) promotes pair breeding. We constructed four contrast experimental groups at a field site in De Vosbergen, The Netherlands. Each experimental group has access to the same total resource amount in a certain area, however the groups vary in the number and size of resource units (i.e. three 30g dead mice, vs six 15g dead mice, vs one 30g and four 15g dead mice, vs two 30g and two 15g dead mice). We predict that in the group that have access to large but less carcasses: (1) there are no differences in the interspecific competition (i.e. the number and species of competitors) between these four groups, (2) burying beetles will compete more for gaining access to carcasses; (3) communal breeding will be more likely to occur, and (4) the per-capita reproductive output(brood size and mass of the brood) will be lower as a consequence of a more pronounced trade-offs between investment for competition (fighting) and reproduction.

Aggression is expressed during intraspecific competition over finite resources (food, territories, mates)essential for survival and reproduction. Aggression likely increases individuals’ resource-holding potential.However, there are still knowledge gaps in our understanding of the trade-offs governing the eco-evolutionary causes and consequences of aggressive behaviour. Given that natural environments are typically heterogeneous, with high and low-quality patches in terms of their resources, correlations are expected between the competitiveness of individual phenotypes that may be related to their aggressiveness, and the quality of their breeding environments. Such phenotype–environment correlations may challenge the partitioning of phenotypic and environmental contributions to among-individual variation in fitness (reproductive performance) and drive the dynamics of dispersal and settlement within population. In this PhD project, we will investigate the combined effects of phenotype(aggression) and environmental quality in driving dispersal, settlement and fitness using a long-term individual-based population study of blue tits (Cyanistes caeruleus). The study has four main objectives:(1) Testing for phenotype–environment correlation between aggression and territory quality; (2)Assessing the role of individual aggression in mate selection and the fitness consequences for each sex;(3) Documenting the links between individual aggression, environmental quality and breeding dispersal;(4) Providing a meta-analysis on the trade-off between resource acquisition and resource utilization.

Presence data for some hummingbird species demonstrates that cities are good living spaces across the Americas. Monitoring hummingbird populations in big cities shows that some species could be very abundant in highly transformed environments. Individuals are managing to face interaction with human activities. On the other hand, forest specialists or species less tolerant to habitat disturbance have become rare or completely absent from cities or other environments modified by humans, such as peri urban suburbia or agricultural landscapes. Hummingbirds are particularly challenging animals to monitor with marking methods, their metabolic requirements and small bodies can not hold heavy targets or electronic devices for telemetry. Even ring marking could be considered not completely adequate. Xalapa is a medium size city located in central east Mexico where cloud montane forest used to extend. Two hummingbird species (Pampa curvipennis and Saucerottia cyanocephala) associated with cloud forest are becoming abundant city species and have been observed feeding from several exotic plants and nesting in highly perturbed environments such as human houses. The third aim of my PhD project is to gain a better understanding on how hummingbirds are coping with human environments when forests are highly transformed and cities become attractive providers of nectar. Some research questions to address are: How cloud forest hummingbirds respond to human presence: exotic plants and artificial feeders? Arecities becoming ecological traps? Can we identify if urban populations are behaving differently towards humans than forest ones? (Tolerant to noise, cars or people when feeding). At what point of environmental transformation will species disappear from their habitat forests? Are native individuals from cities also able to return to forest or an ecological segregation could be going on among our backyard gardens? I present preliminary ideas to discuss these topics in order to set a monitoring method to explain why forest hummingbirds are now living in cities.

Aggressive behaviour is common in nature and may be closely linked to individual fitness (i.e., survival and reproductive success). Aggressive individuals may gain priority access to resources such as food and mates. At the same time, they may also bear costs, including energy expenditure, risk of injury,exposure to predation, and physiological stress. At present, research on individual differences in aggressiveness and its ecological causes and consequences remains scarce. Moreover, much research has focused predominantly on aggressiveness in males rather than females. In this study, we tested the repeatability of aggressiveness in both sexes in a nestbox-breeding blue tit (Cyanistes caeruleus) population using simulated territorial intrusions. We investigated female and male aggressiveness towards intruders of both sexes during different breeding stages (nest-building and egg-laying), and with different modes of song playback (paused and continuous). We found that aggressiveness was repeatable in both sexes. Male blue tits showed greater aggressiveness towards male intruders than female intruders, and their aggressiveness levels decreased in the egg-laying stage compared to the nest-building stage. In contrast, we found no significant differences in female aggressiveness between female and male intruders or different breeding stages. As for song type, paused song triggered more aggressiveness in each breeding stage in both sexes. Furthermore, focal bird age influenced aggressiveness — first-year breeders showed stronger aggressive responses than older ones, but only in the nest-building stage. Our results show that intruder sex and breeding stage relate to male but not female aggressiveness, suggesting different ecological trade-offs may underlie the observed variation in female and male aggressiveness.

Among-individual variation in behaviours closely related to fitness, such as risk-taking behaviours, is found across the animal kingdom. However, its ecological drivers are poorly understood. Theoretical work suggests that individual differences in investment into current versus future reproduction may lead to a“slow-fast” continuum, known as the “pace-of-life syndrome” (POLS) hypothesis. Accordingly, populations may show variation among individuals in the level of risk-taking behaviours (i.e., "fast” individuals tend to take more risks and invest more in current breeding endangering their own survival as opposed to “slow”individuals). In addition, it has been suggested that consistent differences in individual risk-taking behaviours are repeatable across various threatening contexts. Yet, empirical studies in a natural setting remain scarce. The objectives of this study are: (1) to establish to what extent levels of risk-taking are repeatable in blue tits (males and females) across different contexts, and (2) to investigate how the variation in risk-taking relates to levels of parental investment. We will assess this hypothesis in a wild blue tit (Cyanistes caeruleus) population using two risky contexts during the nestling stage: (1) presenting a Eurasian sparrowhawk (Accipiter nisus) model as a predator stimulus, and (2) temporarily removing one parent bird to simulate a predation event. In both cases, to quantify the variation in risk-taking behaviour,we will use the latencies of birds to resume brood provisioning after having been exposed to the threat.Furthermore, we will collect fitness-related data (e.g., egg-laying date, clutch size, offspring survival) to investigate the fitness consequences of this behaviour. As predicted by the POLS hypothesis, we expect risk-taking behaviour to be consistent and repeatable in the individuals under different contexts.Moreover, we predict individuals that take more risk to have higher breeding success, as they invest more in their current reproduction (at the cost of their own survival).

The pace-of-life syndrome (POLS) hypothesis posits that consistent between-individual variation in behavioural traits (‘animal personalities’) facilitate trade-offs in life history. Individuals who display risk averse behaviours are expected to follow a ‘slow’ pace-of-life, including short lifespan and delayed reproduction, compared to their risk-taking, ‘fast’, counterparts. Despite a breadth of empirical research testing the POLS hypothesis, findings are equivocal. This is likely due to core assumptions of the POLS hypothesis not being met, including clear trade-offs in life history, resource limited conditions, and evidence for a functional role of the behavioural trait being investigated. Here, we use long-term data from a closed island population of individually-marked Seychelles warblers to determine the relationship between repeatable exploration–avoidance personalities and an individual’s age at first breeding, lifespan and lifetime reproductive success, separately in males and females. In this population, there is a clear trade-off in reproduction and survival: individuals with a higher age at first breeding display delayed survival senescence. Further, high intraspecific competition for breeding vacancies, as well as a functional role of exploratory traits in accessing these vacancies, also exist. Using multivariate models, we found no evidence for covariation between exploratory and life history traits in either sex. We thus found no support for the POLS hypothesis, despite accounting for core components that may have masked the prevalence of POLS. Our results demonstrate that whilst particular behavioural traits may play functional roles in certain aspects of life history, they may not necessarily be influential enough to have overarching effects on an individual’s pace-of-life.

The environment experienced during development, and its impact on intrinsic condition, can have lasting outcomes for individual phenotypes and could contribute to variation in adult senescence trajectories. However, the nature of this relationship in wild populations remains uncertain, owing to the difficulties in summarizing natal conditions and in long-term monitoring of individuals from free-roaming long-lived species. Utilizing a closely monitored, closed population of Seychelles warblers (Acrocephalus sechellensis), we determine whether juvenile body mass is associated with natal socioenvironmental factors, specific genetic traits linked to fitness in this system, survival to adulthood, and senescence-related traits. Juveniles born in seasons with higher food availability and into smaller natal groups (i.e., fewer competitors) were heavier. In contrast, there were no associations between juvenile body mass and genetic traits. Furthermore, size-corrected mass—but not separate measures of natal food availability, group size, or genetic traits—was positively associated with survival to adulthood, suggesting juvenile body mass is indicative of natal condition. Heavier juveniles had greater body mass and had higher rates of annual survival as adults, independent of age. In contrast, there was no association between juvenile mass and adult telomere length attrition (a measure of somatic stress) nor annual reproduction. These results indicate that juvenile body mass, while not associated with senescence trajectories, can influence the likelihood of surviving to old age, potentially due to silver-spoon effects. This study shows that measures of intrinsic condition in juveniles can provide important insights into the long-term fitness of individuals in wild populations.

In animals, species differ remarkably in parental care strategies. For instance, male-only care is prevalent in teleost fishes, while biparental care predominates in birds and female-only care is widespread in mammals. Understanding the origin and maintenance of diversified parental care systems is a key challenge in evolutionary ecology. It has been suggested that ecological factors and life-history traits play important roles in the evolution of parental care, but the generality of these predictions has not been investigated across a broad range of taxa. Using phylogenetic comparative analyses and detailed parental care data from 1101 avian species that represent 119 families of 26 orders, here we investigate whether parental strategies are associated with ecological variables (i.e., food type, nest structure, and coloniality) and life-history characteristics (i.e., chick development mode and body size). We show that parental care strategies are in relation to coloniality (solitary, semi-colonial, colonial) and chick development mode (altricial vs. precocial). Colonial and altricial species provide more biparental care than solitary and precocial species, respectively. In contrast, food type (plant, invertebrate, vertebrate), nest structure (open vs. closed), and body size do not covary systematically with parental care patterns in birds. Taken together, our results suggest that living in groups and/or having high-demand offspring are strongly associated with biparental care. Towards the end, we discuss future research directions for the study of parental care evolution. Significance statement: Animal species differ remarkably in the amount of care parents provide to their offspring and in the distribution of care tasks over the parents. In birds, for example, the young of some species are quite independent from the start, while the young of other species are helpless after hatching, requiring a lot of care. Moreover, either the female or the male does most of the caring in some species, while the parental tasks are shared equally in still other species. To understand the diversified parental care patterns, we applied advanced comparative methods to a large data set comprising over 1000 bird species. The analysis reveals that the parents tend to share their care duties equally when they live in groups and/or have offspring that are born helpless, and that parental care patterns are not associated with diet, nest type or body size. Hence, living in groups and having high-demand offspring seem to play important roles in the evolution of parental care.

Queens of eusocial species live extraordinarily long compared to their workers. So far, it has been argued that these lifespan divergences are readily explained by the classical evolutionary theory of ageing. As workers predominantly perform risky tasks, such as foraging and nest defense, and queens stay in the well-protected nests, selection against harmful genetic mutations expressed in old age should be weaker in workers than in queens due to caste differences in extrinsic mortality risk, and thus, lead to the evolution of longer queen and shorter worker lifespans. However, these arguments have not been supported by formal models. Here, we present a model for the evolution of caste-specific ageing in social insects, based on Williams’ antagonistic pleiotropy theory of ageing. In individual-based simulations, we assume that mutations with antagonistic fitness effects can act within castes, that is, mutations in early life are accompanied by an antagonistic effect acting in later life, or between castes, where antagonistic effects emerge due to caste antagonism or indirect genetic effects between castes. In monogynous social insect species with sterile workers, large lifespan divergences between castes evolved under all different scenarios of antagonistic effects, but regardless of the degree of caste-specific extrinsic mortality. Mutations with antagonistic fitness effects within castes reduced lifespans of both castes, while mutations with between-caste antagonistic effects decreased worker lifespans more than queen lifespans, and consequently increased lifespan divergences. Our results challenge the central explanatory role of extrinsic mortality for caste-specific ageing in eusocial organisms and suggest that antagonistic pleiotropy affects castes differently due to reproductive monopolization by queens, hence, reproductive division of labor. Finally, these findings provide new insights into the evolution of tissue-specific ageing in multicellular organisms in general.

Mutation provides novelty, and thus the raw material for selection to act on. Previous evidence has shown that increased mutation rates can indeed lead to higher evolutionary rates and faster adaptation. However, when organisms are well adapted, most mutations are likely to be maladaptive – depending on the current degree of adaptation, increased mutation rates may thus be either beneficial or harmful. Particularly interesting in this context is stress-induced mutagenesis: a condition-dependent upregulation of mutation rates under stress. As stress may be a signal for maladaptation, stress-induced mutagenesis may allow mutations to occur predominantly in times of maladaptation, i.e. when they are more likely to be adaptive. Whilst this has received much attention in empirical work, comparatively few theoretical models have explored this topic. Here, we use individual-based simulations to investigate the evolution of condition-dependent mutation rates in fluctuating environments, where environmental change repeatedly leads to maladaptation and thus to stress. We show that condition-dependent mutation rates are indeed adaptive in this scenario – however, they only evolve under specific circumstances. We explore how this relates to initial conditions, selection strength, genome structure and patterns of environmental change. Overall, we discuss the relevance of condition-dependent mutation rates for evolvability, i.e. for the capacity for adaptive evolution.

Environmental conditions experienced during early life may have long-lasting effects on later-life phenotypes and fitness. Individuals experiencing poor early-life conditions may suffer subsequent fitness constraints. Alternatively, individuals may use a strategic “Predictive Adaptive Response” (PAR), whereby they respond—in terms of physiology or life-history strategy—to the conditions experienced in early life to maximize later-life fitness. Particularly, the Future Lifespan Expectation (FLE) PAR hypothesis predicts that when poor early-life conditions negatively impact an individual's physiological state, it will accelerate its reproductive schedule to maximize fitness during its shorter predicted life span. We aimed to measure the impact of early-life conditions and resulting fitness across individual lifetimes to test predictions of the FLE hypothesis in a wild, long-lived model species. Using a long-term individual-based dataset, we investigated how early-life conditions are linked with subsequent fitness in an isolated population of the Seychelles warbler Acrocephalus sechellensis. How individuals experience early-life environmental conditions may vary greatly, so we also tested whether telomere length—shorter telomers are a biomarker of an individual's exposure to stress—can provide an effective measure of the individual-specific impact of early-life conditions. Specifically, under the FLE hypothesis, we would expect shorter telomeres to be associated with accelerated reproduction. Contrary to expectations, shorter juvenile telomere length was not associated with poor early-life conditions, but instead with better conditions, probably as a result of faster juvenile growth. Furthermore, neither juvenile telomere length, nor other measures of early-life conditions, were associated with age of first reproduction or the number of offspring produced during early life in either sex. We found no support for the FLE hypothesis. However, for males, poor early-life body condition was associated with lower first-year survival and reduced longevity, indicating that poor early-life conditions pose subsequent fitness constraints. Our results also showed that using juvenile telomere length as a measure of early-life conditions requires caution, as it is likely to not only reflect environmental stress but also other processes such as growth.

Breastfeeding (duration) can be positively associated with infant growth outcomes as well as improved cognitive functions during childhood and later life stages. (Prolonged) exposure to optimal lipid quantity and quality, i.e., the supramolecular structure of lipids, in mammalian milk, may contribute to these beneficial effects through nutritional early-life programming. In this pre-clinical study, we exposed male C57BL/6J mice from post-natal Days 16 to 42 (i.e., directly following normal lactation), to a diet with large lipid droplets coated with bovine milk fat globule membrane-derived phospholipids, which mimic more closely the supramolecular structure of lipid droplets in mammalian milk. We investigated whether exposure to this diet could affect growth and brain development-related parameters. As these outcomes are also known to be affected by the post-weaning social environment in mice, we included both individually housed and pair-wise housed animals and studied whether effects of diet were modulated by the social environment. After Day 42, all the animals were fed standard semi-synthetic rodent diet. Growth and body composition were assessed, and the mice were subjected to various behavioral tests. Individual housing attenuated adolescent growth, reduced femur length, and increased body fat mass. Adult social interest was increased due to individual housing, while cognitive and behavioral alterations as a result of different housing conditions were modest. The diet increased adolescent growth and femur length, increased lean body mass, reduced adolescent anxiety, and improved adult cognitive performance. These effects of diet exposure were comparable between individually and socially housed mice. Hence, early life exposure to a diet with lipid droplets that mimic the supramolecular structure of those in mammalian milk may improve adolescent growth and alters brain function in both socially and individually housed mice. These findings suggest that lipid structure in infant milk formula may be a relevant target for nutritional solutions, targeting both healthy infants and infants facing growth challenges.

Cooperatively breeding animals live longer than their solitary counterparts. This has been suggested for birds, mole rats, and social insects. A common explanation for these long lifespans is that cooperative breeding evolves more readily in long-lived species because lower mortality reduces the rate of territory turnover and thus leads to a limitation of breeding territories. Here, we reverse this argument and show that—rather than being a cause for its evolution—long lifespans are an evolutionary consequence of cooperative breeding. In evolutionary individual-based simulations, we show that natural selection favors a delayed onset of senescence in cooperative breeders, relative to solitary breeders, because cooperative breeders have a delayed age of first reproduction as helpers wait in a reproductive queue to obtain breeder status. Especially long lifespans evolve in cooperative breeders in which queue positions depend on the helpers’ age rank among the helpers within the breeding territory. Furthermore, we show that lower genetic relatedness among group members leads to the evolution of longer lifespans. This is because selection against higher mortality is weaker when mortality reduces competition for breeding between relatives. Our results link the evolutionary theory of ageing with kin selection theory, demonstrating that the evolution of ageing in cooperative breeders is driven by the timing of reproduction and kin structure within breeding territories.

After the breeding season, territorial adult Black-crested Titmouse (Baeolophus atricristatus) and residing juveniles form social groups that may persist until the following spring. Under the prolonged brood care hypothesis, one would expect these juveniles to be retained offspring with delayed dispersal of the breeding pair. To test if Black-crested Titmouse juveniles that reside in post-breeding territories are offspring of the territorial adult male, we performed microsatellite-based paternity analyses of 6 juvenile–adult male social dyads on 6 different territories. None of the juveniles were offspring of the adult male with which it shared a territory. We discuss several possible evolutionary explanations for this result.

Sex roles describe sex differences in courtship, mate competition, social pair-bonds and parental care. A key challenge is to identify associations among the components and the drivers of sex roles. Here, we investigate sex rolesusing data from over 1800 bird species. We found extensive variation and lability in proxies of sex roles, indicating remarkably independent evolution among sex role components. Climate and life history showed weak associations with sex roles. However, adult sex ratio is associated with sexual dimorphism, mating system and parental care, suggesting that social environment is central to explaining variation in sex roles among birds. Our results suggest that sex differences in reproductive behaviour are the result of diverse and idiosyncratic responses to selection. Further understanding of sex roles requires studies at the population level to test how local responses to ecology, life histories and mating opportunities drive processes that shape sex role variation among higher taxa. © 2022 John Wiley & Sons Ltd.

Communal breeding, wherein multiple conspecifics live and reproduce together, may generate short-term benefits in terms of defence and reproduction. However, its carry-over effects remain unclear. We experimentally tested the effects of communal breeding on parental care and reproduction in burying beetles (Nicrophorus vespilloides), which use carcasses as breeding resources and provide parental care to offspring. We subjected individuals to communal or non-communal breeding (i.e. pair breeding) during their first breeding event and to non-communal breeding during their second breeding event. We measured the parental care of individuals and of groups and the reproductive success of groups during both breeding events. In communal groups, large individuals became dominant and largely monopolized the carcass, whereas small individuals (i.e. subordinates) had restricted access to the carcass. At the first breeding event, large males in communal groups spent more time providing care than large males in non-communal groups, whereas such an effect was not observed for large females and small individuals. Reproductive successes were similar in communal and non-communal groups, indicating no short-term benefits of communal breeding in terms of reproduction. Compared with males from non-communal groups, males originating from communal groups produced a larger size of brood during their second breeding event, whereas such an effect was not observed for females. Our results demonstrate the sex-specific effects of communal breeding experience on parenting performance and fitness.

Success in sperm competition is an important determinant of male fitness in mating systems with female multiplemating. Thus, sperm competition risk represents a key dimension of the male social environment to which in-dividual males are expected to adaptively adjust their reproductive phenotype. Such adaptive phenotypicadjustment we here refer to as male social niche conformance. In this pre-registered study, we investigated howmale zebra finches, Taeniopygia guttata, adjust their behavior to sperm competition risk. We experimentallymanipulated the opportunity for extra-pair mating to create two levels of sperm competition risk: 1) Single-pair,no sperm competition risk; 2) Double-pair, sperm competition risk. We compared male courtship, mate guarding,copulation rates, and aggression between the treatment groups. To identify hormonal correlates of malebehavioral adjustment, we measured plasma testosterone and corticosterone levels before and after the socialtreatment started. Contrary to our pre-registered predictions, males from the Double-pair treatment groupdecreased courtship rates compared to those from the Single-pair group, and Double-pair males responded lessaggressively towards intruders than Single-pair males. Testosterone levels decreased over the breeding cycle, butsocial treatment had no effect on either testosterone or corticosterone levels. Our results indicate that male zebrafinches do not intensify courtship or competitive reproductive behaviors, or upregulate key hormones whenanother breeding pair is present. Although we found no evidence for the predicted adaptive behavioral responsesto sperm competition risk, we show that male zebra finches plastically adjust their behavior to their socialenvironment

Altered interactions between pathogens, their hosts and vectors have potential consequences for human disease risk. Notably, tick-borne pathogens, many of which are associated with growing deer abundance, show global increasing prevalence and pose increasing challenges for disease prevention. Human activities can largely affect the patterns of deer space-use and can therefore be potential management tools to alleviate human-wildlife conflicts. Here, we tested how deer space-use patterns are influenced by human recreational activities, and how this in turn affects the spatial distribution of the sheep tick (Ixodes ricinus), a relevant disease vector of zoonoses such as Lyme borrelioses. We compared deer dropping and questing tick density on transects near (20 m) and further away from (100 m) forest trails that were either frequently used (open for recreation) or infrequently used (closed for recreation, but used by park managers). In contrast to infrequently used trails, deer dropping density was 31% lower near (20 m) than further away from (100 m) frequently used trails. Similarly, ticks were 62% less abundant near (20 m) frequently used trails compared to further away from (100 m) these trails, while this decline in tick numbers was only 14% near infrequently used trails. The avoidance by deer of areas close to human-used trails was thus associated with a similar reduction in questing tick density near these trails. As tick abundance generally correlates to pathogen prevalence, the use of trails for recreation may reduce tick-borne disease risk for humans on and near these trails. Our study reveals an unexplored effect of human activities on ecosystems and how this knowledge could be potentially used to mitigate zoonotic disease risk.

Although biparental care is the most commonly observed pattern in bird species, the evolutionary causations of variable parental care patterns demand more scientific attention. I investigated the evolutionary ecology of a variable breeding system of the Chinese penduline tits (Remiz consobrinus), in which uniparental, biparental care and biparental desertion of nests occur, and care patterns differ within and between populations, ranging from mainly female-only care to mainly biparental care across populations. To explore the mechanisms of such a flexible parental care system, I used observational, experimental and theoretical methodologies. Within populations, I found that the higher the re-mating opportunities for males, the higher the rate of male desertion. However, re-mating opportunity for females was not associated with female desertion. Across populations, parental care pattern and breeding success of parental care types differed greatly, and revealed that higher male competition intensity and lower offspring survival success may cause biparental care evolved more in one population than the other. Field experiments testing functions of egg burial behaviour found that instead of resolving sexual conflict as that in Eurasian penduline tits, Chinese penduline tits bury eggs to prevent eggs from being blown out of nests. Parents of uniparental care have high parenting capabilities. By temporarily removing the partner of biparental cared nest, I discovered that the remaining partner compensated for the provisioning frequency but not for the brooding duration. The Individual-based simulation revealed that season length and uniparental care efficiency variably affect parental care and lead to polymorphic patterns in a population.

In many animals, parents provide care to their offspring, such as building nests, or feeding the young. Species differ considerably in how parental care is distributed between the male and the female parent. In my PhD thesis, I shed new light on the question how the baffling diversity of parental care patterns 1,101 bird species, in order to find out whether ecological factors (such as nest type) or life-history characteristics (such as body size) explain when species exhibit uniparental or biparental care. Although I could identify some significant factors (e.g., breeding in a colony), my main conclusion is that many hypotheses proposed in the literature are not confirmed by the data. Second, I constructed theoretical models in order to gain a better understanding of how and why diverse parental care patterns emerge in the course of evolution. I analysed these models by means of individual-based simulations, which are more versatile and based on fewer simplifying assumptions than standard mathematical approaches. My simulations provide surprising new insights. During evolution, regularly ‘care polymorphisms’ emerge where very different care strategies coexist in males and/or females. Although these polymorphisms may disappear again after a brief period of time, I could show that they are often decisive for the course and outcome of evolution. This finding has important implications for the evolution of parental sex roles, and it necessitates a reconsideration of many seemingly well-established predictions of sex role theory.

In socially monogamous species with bi-parental care, males may face a trade-off between providing parental care and pursuing extra-pair matings. The "parenting-mating trade-off" hypothesis predicts that high-quality males-who have greater potential to gain extra-pair matings, for example, larger males usually win the competition for extra-pair mating-should reduce parental care and spend more time looking for extra-pair matings. However, the trade-off between parenting and mating efforts may be complicated by variation in the availability of extra-pair mating opportunities. By using field data of hair-crested drongos (Dicrurus hottentottus), a species exhibiting bi-parental incubation behavior, collected in central China from 2010 to 2017, we tested whether the potential negative relationship between male quality and paternal care was dependent on the number of nearby fertile females. We found that male drongos mainly seek extra-pair matings during the incubation period and high-quality individuals (males with longer tarsi) are more likely to sire extra-pair offspring. In agreement with the "parenting-mating trade-off" hypothesis, high-quality males incubated less by recessing longer between incubation bouts. However, this was only the case when sufficient fertile females nearby for extra-pair mating opportunities. Females compensated for reduced male care, but this was independent of male quality. This suggests that the reduction in care by high-quality males might be a direct response to extra-pair mating opportunities rather than facilitated by differential allocation of females. Our results indicate that individual quality and available mating opportunities may shape the optimal trade-off between parental care and seeking additional matings for males.Lay Summary: How do individual quality and the number of potential extra-pair mates influence the optimal trade-off between parental care and engaging extra-pair courtship in animals? In hair-crested drongos, high-quality males, who are more successful in obtaining extra-pair fertilizations, reduced their share in incubation, but only when they had sufficient extra-pair mating opportunities. Females partially compensate for the reduced incubation of their partners, but the compensation was not affected by male quality.

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.

Background The vertebrate gut microbiome (GM) can vary substantially across individuals within the same natural population. Although there is evidence linking the GM to health in captive animals, very little is known about the consequences of GM variation for host fitness in the wild. Here, we explore the relationship between faecal microbiome diversity, body condition, and survival using data from the long-term study of a discrete natural population of the Seychelles warbler (Acrocephalus sechellensis) on Cousin Island. To our knowledge, this is the first time that GM differences associated with survival have been fully characterised for a natural vertebrate species, across multiple age groups and breeding seasons. Results We identified substantial variation in GM community structure among sampled individuals, which was partially explained by breeding season (5% of the variance), and host age class (up to 1% of the variance). We also identified significant differences in GM community membership between adult birds that survived, versus those that had died by the following breeding season. Individuals that died carried increased abundances of taxa that are known to be opportunistic pathogens, including several ASVs in the genus Mycobacterium. However, there was no association between GM alpha diversity (the diversity of bacterial taxa within a sample) and survival to the next breeding season, or with individual body condition. Additionally, we found no association between GM community membership and individual body condition. Conclusions These results demonstrate that components of the vertebrate GM can be associated with host fitness in the wild. However, further research is needed to establish whether changes in bacterial abundance contribute to, or are only correlated with, differential survival; this will add to our understanding of the importance of the GM in the evolution of host species living in natural populations.

Life-history theory predicts that increased resource allocation in current reproduction comes at the cost of survival and future reproductive fitness. In taxa with biparental care, each parent can adjust investment on current reproduction according to changes in their partner’s effort, but these adjustments may be different for males and females as they may have different reproductive strategies. Numerous theoretical and empirical studies have proposed the mechanism underlying such adjustments. In addition, the value of the brood or litter (brood size) has also been suggested to affect the amount of care through manipulation of brood size. While the two conditions have been studied independently, the impact of their interplay on potential sex-dependent future reproductive performance remains largely unknown. In this study, we simultaneously manipulated both care system (removal of either parent vs. no removal) and brood size in a burying beetle (Nicrophorus vespilloides) to understand their joint effect on reproductive allocation and trade-off between current and future reproduction. Our results show that males compensated for mate loss by significantly increasing the level of care regardless of brood size, while females exhibited such compensation only for small brood size. Additionally, with an increase in allocation to current reproduction, males showed decreased parental investment during the subsequent breeding event as a pair. These findings imply a dual influence of parental care system and brood size on allocation in current reproduction. Moreover, the impact of such adjustments on sex-dependent differences in future reproduction (parental care, larvae number, and average larval mass at dispersal) is also demonstrated. Our findings enhance the understanding of sex roles in parental investment and highlight their importance as drivers of reproductive allocation.

How can the stunning diversity of social systems and behaviours seen in nature be explained? Drawing on social evolution theory, experimental evidence and studies conducted in the field, this book outlines the fundamental principles of social evolution underlying this phenomenal richness.To succeed in the competition for resources, organisms may either 'race' to be quicker than others, 'fight' for privileged access, or 'share' their efforts and gains. The authors show how the ecology and intrinsic attributes of organisms select for each of these strategies, and how a handful of straightforward concepts explain the evolution of successful decision rules in behavioural interactions, whether among members of the same or different species. With a broad focus ranging from microorganisms to humans, this is the first book to provide students and researchers with a comprehensive account of the evolution of sociality by natural selection.

While it is universally recognised that environmental factors can cause phenotypic trait variation via phenotypic plasticity, the extent to which causal processes operate in the reverse direction has received less consideration. In fact individuals are often active agents in determining the environments, and hence the selective regimes, they experience. There are several important mechanisms by which this can occur, including habitat selection and niche construction, that are expected to result in phenotype–environment correlations (i.e. non-random assortment of phenotypes across heterogeneous environments). Here we highlight an additional mechanism – intraspecific competition for preferred environments – that may be widespread, and has implications for phenotypic evolution that are currently underappreciated. Under this mechanism, variation among individuals in traits determining their competitive ability leads to phenotype–environment correlation; more competitive phenotypes are able to acquire better patches. Based on a concise review of the empirical evidence we argue that competition-induced phenotype–environment correlations are likely to be common in natural populations before highlighting the major implications of this for studies of natural selection and microevolution. We focus particularly on two central issues. First, competition-induced phenotype–environment correlation leads to the expectation that positive feedback loops will amplify phenotypic and fitness variation among competing individuals. As a result of being able to acquire a better environment, winners gain more resources and even better phenotypes – at the expense of losers. The distinction between individual quality and environmental quality that is commonly made by researchers in evolutionary ecology thus becomes untenable. Second, if differences among individuals in competitive ability are underpinned by heritable traits, competition results in both genotype–environment correlations and an expectation of indirect genetic effects (IGEs) on resource-dependent life-history traits. Theory tells us that these IGEs will act as (partial) constraints, reducing the amount of genetic variance available to facilitate evolutionary adaptation. Failure to recognise this will lead to systematic overestimation of the adaptive potential of populations. To understand the importance of these issues for ecological and evolutionary processes in natural populations we therefore need to identify and quantify competition-induced phenotype–environment correlations in our study systems. We conclude that both fundamental and applied research will benefit from an improved understanding of when and how social competition causes non-random distribution of phenotypes, and genotypes, across heterogeneous environments.

Evolutionary theory suggests that individuals can benefit from deferring the fitness cost of developing under poor conditions to later in life. Although empirical evidence for delayed fitness costs of poor developmental conditions is abundant, individuals that die prematurely have not often been incorporated when estimating fitness, such that age-specific fitness costs, and therefore the relative importance of delayed fitness costs are actually unknown.We developed a Bayesian statistical framework to estimate age-specific reproductive values in relation to developmental conditions. We applied it to data obtained from a long-term longitudinal study of common terns Sterna hirundo, using sibling rank to describe variation in developmental conditions. Common terns have a maximum of three chicks, and later hatching chicks acquire less food, grow more slowly and have a lower fledging probability than their earlier hatched siblings.We estimated fitness costs in adulthood to constitute c. 45% and 70% of the total fitness costs of hatching third and second, respectively, compared to hatching first. This was due to third-ranked hatchlings experiencing especially high pre-fledging mortality, while second-ranked hatchlings had lower reproductive success in adulthood. Both groups had slightly lower adult survival. There was, however, no evidence for sibling rank-specific rates of senescence.We additionally found years with low fledgling production to be associated with particularly strong pre-fledging selection on sibling rank, and with increased adult survival to the next breeding season. This suggests that adults reduce parental allocation to reproduction in poor years, which disproportionately impacts low-ranked offspring.Interpreting these results, we suggest that selection at the level of the individual offspring for delaying fitness costs is counteracted by selection for parental reduction in brood size when resources are limiting.

Senescence is widespread in nature, often resulting in diminishing survival or reproduction with age, but its role in age-dependent variation in sexual traits is often poorly understood. One reason is that few studies of sexual traits consider non-linear relationships with age, or only consider a narrow range of years relative to the life span of the species. Birdsong has evolved to allow assessment of conspecific quality in numerous bird species. Whilst theory and empirical work suggests that song may become more elaborate with age, there are a paucity of long-term studies testing whether song is associated with age or longevity. In particular, the occurrence of song senescence has rarely been demonstrated. Using an exceptional long-term dataset for the Seychelles warbler (Acrocephalus sechellensis), we analysed relationships between male song, age, survival, and longevity. This species is a long-lived songbird with early life increases, followed by senescent declines, in survival and reproduction. The study population (Cousin Island, Seychelles) is a closed population, with no depredation of adults, providing an excellent opportunity to study senescence in free-living animals. We tested whether song traits were related to age at recording, future survival, longevity, and territory quality. We found age-dependent changes in five song traits (duration, maximum frequency, peak frequency of songs, and duration and frequency bandwidth of trills). Relationships with age were quadratic, indicating reversal in the expression of song coinciding with the onset of senescence in reproduction and survival in this species. One song trait (trill bandwidth) had a quadratic relationship with future survival, but no song traits were related to longevity, suggesting age-related patterns were not the result of selective disappearance. Our study provides one of the first examples of functional senescence in song, offering new insights into avian senescence. Late-life declines in avian song, and possibly other sexual traits, may be more common than currently known, and may play a fundamental role in age-dependent changes in reproductive success.

Numerous cases of evolutionary trait loss and regain have been reported over the years. Here, we argue that such reverse evolution can also become apparent when trait expression is plastic in response to the environment. We tested this idea for the loss and regain of fat synthesis in parasitic wasps. We first show experimentally that the wasp Leptopilina heterotoma switches lipogenesis on in a fat-poor environment, and completely off in a fat-rich environment. Plasticity suggests that this species did not regain fat synthesis, but that it can be switched off in some environmental settings. We then compared DNA sequence variation and protein domains of several more distantly related parasitoid species thought to have lost lipogenesis, and found no evidence for non-functionality of key lipogenesis genes. This suggests that other parasitoids may also show plasticity of fat synthesis. Last, we used individual-based simulations to show that a switch for plastic expression can remain functional in the genome for thousands of generations, even if it is only used sporadically. The evolution of plasticity could thus also explain other examples of apparent reverse evolution.

Sleep is a behavioral and physiological state that is thought to serve important functions. Many animals go through phases in the annual cycle where sleep time might be limited, for example, during the migration and breeding phases. This leads to the question whether there are seasonal changes in sleep homeostasis. Using electroencephalogram (EEG) data loggers, we measured sleep in summer and winter in 13 barnacle geese (Branta leucopsis) under semi-natural conditions. During both seasons, we examined the homeostatic regulation of sleep by depriving the birds of sleep for 4 and 8 h after sunset. In winter, barnacle geese showed a clear diurnal rhythm in sleep and wakefulness. In summer, this rhythm was less pronounced, with sleep being spread out over the 24-h cycle. On average, the geese slept 1.5 h less per day in summer compared with winter. In both seasons, the amount of NREM sleep was additionally affected by the lunar cycle, with 2 h NREM sleep less during full moon compared to new moon. During summer, the geese responded to 4 and 8 h of sleep deprivation with a compensatory increase in NREM sleep time. In winter, this homeostatic response was absent. Overall, sleep deprivation only resulted in minor changes in the spectral composition of the sleep EEG. In conclusion, barnacle geese display season-dependent homeostatic regulation of sleep. These results demonstrate that sleep homeostasis is not a rigid phenomenon and suggest that some species may tolerate sleep loss under certain conditions or during certain periods of the year., Sleep, 44 (4), ISSN:1550-9109, ISSN:0161-8105

Nest predation is a common cause of reproductive failure for many bird species, and various antipredator defense behaviors have evolved to reduce the risk of nest predation. However, trade-offs between current reproductive duties and future reproduction often limit the parent’s ability to respond to nest predation risk. Individual responses to experimentally increased nest predation risk can give insights into these trade-offs. Here, we investigate whether social and ecological factors affect individual responses to predation risk by experimentally manipulating the risk of nest predation using taxidermic mounts in the cooperative breeding Seychelles warbler (Acrocephalus sechellensis). Our results show that dominant females, but not males, alarm called more often when they confront a nest predator model alone than when they do so with a partner, and that individuals that confront a predator together attacked more than those that did so alone. Dominant males increased their antipredator defense by spending more time nest guarding after a presentation with a nest predator, compared with a nonpredator control, but no such effect was found for females, who did not increase the time spent incubating. In contrast to incubation by females, nest guarding responses by dominant males depended on the presence of other group members and food availability. These results suggest that while female investment in incubation is always high and not dependent on social and ecological conditions, males have a lower initial investment, which allows them to respond to sudden changes in nest predation risk., Parental investment in antipredator defense depends on its relative costs and benefits. By experimentally manipulating nest predation risk in the Seychelles warbler, we show that sex-specific antipredator defenses are modified by social and ecological factors. Females show the highest investment in the nest, while males invest less, enabling them to increase their investment when predation risk increases.