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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.

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.

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.

Genomes retain evidence of the demographic history and evolutionary forces that have shaped populations and drive speciation. Across island systems, contemporary patterns of genetic diversity reflect population demography, including colonization events, bottlenecks, gene flow and genetic drift. Here, we investigate genome-wide diversity and the distribution of runs of homozygosity (ROH) using whole-genome resequencing of individuals (>22x coverage) from six populations across three archipelagos of Berthelot's pipit (Anthus berthelotii)-a passerine that has recently undergone island speciation. We show the most dramatic reduction in diversity occurs between the mainland sister species (the tawny pipit) and Berthelot's pipit and is lowest in the populations that have experienced sequential bottlenecks (i.e., the Madeiran and Selvagens populations). Pairwise sequential Markovian coalescent (PSMC) analyses estimated that Berthelot's pipit diverged from its sister species similar to 2 million years ago, with the Madeiran archipelago founded 50,000 years ago, and the Selvagens colonized 8000 years ago. We identify many long ROH (>1 Mb) in these most recently colonized populations. Population expansion within the last 100 years may have eroded long ROH in the Madeiran archipelago, resulting in a prevalence of short ROH (250 kb. These findings highlight the importance of demographic history, as well as selection and genetic drift, in shaping contemporary patterns of genomic diversity across diverging populations.

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.

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.

Crustaceans are notoriously difficult to age because of their indeterminate growth and the moulting of their exoskeleton throughout life. The poor knowledge of population age structure in crustaceans therefore hampers accurate assessment of population dynamics and consequently sustainable fisheries management. Quantification of DNA methylation of the evolutionarily conserved ribosomal DNA (rDNA) may allow for age prediction across diverse species. However, the rDNA epigenetic clock remains to be tested in crustaceans, despite its potential to inform both ecological and evolutionary understanding, as well as conservation and management practices. Here, patterns of rDNA methylation with age were measured across 5154 bp of rDNA corresponding to 355 quality‐filtered loci in the economically important European lobster (Homarus gammarus). Across 0‐ to 51‐month‐old lobsters (n = 155), there was a significant linear relationship between age and percentage rDNA methylation in claw tissue at 60% of quality‐filtered loci (n = 214). An Elastic Net regression model using 46 loci allowed for the accurate and precise age estimation of individuals (R 2 = 0.98; standard deviation = 1.6 months). Applying this ageing model to antennal DNA from wild lobsters of unknown age (n = 38) resulted in predicted ages that are concordant with estimates of minimum size at age in the wild (mean estimated age = 40.1 months; range 32.8–55.7 months). Overall, the rDNA epigenetic clock shows potential as a novel, nonlethal ageing technique for European lobsters. However, further validation is required across a wider range of known‐age individuals and tissue types before the model can be used in fisheries management.

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) genes). 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, or TLR3 genotype were associated with fungal GM differences in Seychelles warblers. However, the presence of four MHC-I alleles and one MHC-II allele were 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 seven 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 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 upon host evolution.

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 expectedper capitacontribution of the members of that category to the gene pool of future generations. Life history theory reveals how reproductive values can be determined via the estimation of life-history parameters, but this requires intricate algebraic calculations. Recently, a more intuitive, pedigree-based method has become popular, which estimates the genetic contributions of individuals to future generations by tracking their descendants down the pedigree. Here we compare both methods. We implement various life-history scenarios (for which the “true” reproductive values can be calculated) in individual-based simulations, use the simulation data to estimate reproductive values with both methods, and compare the results with the true target 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. In contrast, the traditional algebraic method estimates reproductive values with high accuracy and precision.Lay SummaryTo study evolution in empirical systems it is important to accurately and precisely measure fitness. Here, using simulations, we compare two methods for estimating fitness and test their accuracy and precision. One estimates life-history parameters and calculates a fitness proxy using complex algebra. The other method estimates genetic contribution to future generations by tracking descendants down the pedigree. We conclude that the pedigree method is unreliable, while the algebraic method is accurate and precise.

A key aim of observational campaigns is to sample atmosphere–ocean phenomena to improve understanding of these phenomena, and in turn, numerical weather prediction. In early 2018 and 2019, the Atmospheric River Reconnaissance (AR Recon) campaign released dropsondes and radiosondes into atmospheric rivers (ARs) over the northeast Pacific Ocean to collect unique observations of temperature, winds, and moisture in ARs. These narrow regions of water vapor transport in the atmosphere—like rivers in the sky—can be associated with extreme precipitation and flooding events in the midlatitudes. This study uses the dropsonde observations collected during the AR Recon campaign and the European Centre for Medium-Range Weather Forecasts (ECMWF) Integrated Forecasting System (IFS) to evaluate forecasts of ARs. Results show that ECMWF IFS forecasts 1) were colder than observations by up to 0.6 K throughout the troposphere; 2) have a dry bias in the lower troposphere, which along with weaker winds below 950 hPa, resulted in weaker horizontal water vapor fluxes in the 950–1000-hPa layer; and 3) exhibit an underdispersiveness in the water vapor flux that largely arises from model representativeness errors associated with dropsondes. Four U.S. West Coast radiosonde sites confirm the IFS cold bias throughout winter. These issues are likely to affect the model’s hydrological cycle and hence precipitation forecasts.

The strength of the stratospheric polar vortex influences the surface weather in the Northern Hemisphere in winter; a weaker (stronger) than average stratospheric polar vortex is connected to negative (positive) Arctic Oscillation (AO) and colder (warmer) than average surface temperatures in northern Europe within weeks or months. This holds the potential for forecasting in that timescale. We investigate here if the strength of the stratospheric polar vortex at the start of the forecast could be used to improve the extended-range temperature forecasts of the European Centre for Medium-Range Weather Forecasts (ECMWF) and to find periods with higher prediction skill scores. For this, we developed a stratospheric wind indicator (SWI) based on the strength of the stratospheric polar vortex and the phase of the AO during the following weeks. We demonstrate that there was a statistically significant difference in the observed surface temperature in northern Europe within the 3–6 weeks, depending on the SWI at the start of the forecast. When our new SWI was applied in post-processing the ECMWF's 2-week mean temperature reforecasts for weeks 3–4 and 5–6 in northern Europe during boreal winter, the skill scores of those weeks were slightly improved. This indicates there is some room for improving the extended-range forecasts, if the stratosphere–troposphere links were better captured in the modelling. In addition to this, we found that during the boreal winter, in cases where the polar vortex was weak at the start of the forecast, the mean skill scores of the 3–6 weeks' surface temperature forecasts were higher than average.

Spatial variability of precipitation is analyzed to characterize to what extent precipitation observed at a single location is representative of precipitation over a larger area. Characterization of precipitation representativeness is made in probabilistic terms using a parametric approach, namely, by fitting a censored shifted gamma distribution to observation measurements. Parameters are estimated and analyzed for independent precipitation datasets, among which one is based on high-density gauge measurements. The results of this analysis serve as a basis for accounting for representativeness error in an ensemble verification process. Uncertainty associated with the scale mismatch between forecast and observation is accounted for by applying a perturbed-ensemble approach before the computation of scores. Verification results reveal a large impact of representativeness error on precipitation forecast reliability and skill estimates. The parametric model and estimated coefficients presented in this study could be used directly for forecast postprocessing to partly compensate for the limitation of any modeling system in terms of precipitation subgrid-scale variability.

The performance of an ensemble forecast, as measured by scoring rules, depends on its number of members. Under the assumption of ensemble member exchangeability, ensemble-adjusted scores provide unbiased estimates of the ensemble-size effect. In this study, the concept of ensemble-adjusted scores is revisited and exploited in the general context of multi-model ensemble forecasting. In particular, an ensemble-size adjustment is proposed for the continuous ranked probability score in a multi-model ensemble setting. The method requires that the ensemble forecasts satisfy generalised multi-model exchangeability conditions. These conditions do not require the models themselves to be exchangeable. The adjusted scores are tested here on a dual-resolution ensemble, an ensemble which combines members drawn from the same numerical model but run at two different grid resolutions. It is shown that performance of different ensemble combinations can be robustly estimated based on a small subset of members from each model. At no additional cost, the ensemble-size effect is investigated not only considering the pooling of potential extra-members but also including the impact of optimal weighting strategies. With simple and efficient tools, the proposed methodology paves the way for predictive verification of multi-model ensemble forecasts; the derived statistics can provide guidance for the design of future operational ensemble configurations without having to run additional ensemble forecast experiments for all the potential configurations.

The MolProbity web service provides macromolecular model validation to help correct local errors, for the structural biology community worldwide. Here we highlight new validation features, and also describe how we are fighting back against outside developments which compromise that mission. Our new tool called UnDowser analyzes the properties and context of clashing HOH “waters” to diagnose what they might actually represent; a dozen distinct scenarios are illustrated and described. We now treat alternate conformations more thoroughly, and switching to the Neo4j database (graphical rather than relational) enables cleaner, more comprehensive, and much larger reference datasets. A problematic outside change is that refinement software now increasingly restrains traditional validation criteria (geometry, clashes, rotamers, and even Ramachandran) in order to supplement the sparser experimental data at 3–4 Å resolutions typical of modern cryoEM. But unfortunately the broad density allows model optimization without fixing underlying problems, which means these structures often score much better on validation than they really are. CaBLAM, our tool designed for evaluating peptide orientations at lower resolutions, was described in the previous Tools issue, and here we demonstrate its effectiveness in diagnosing local errors even when other validation outliers have been artificially removed. Sophisticated hacking of the MolProbity server has required continual monitoring and various security measures short of restricting user access. The deprecation of Java applets now prevents KiNG interactive online display of outliers on the 3D model during a MolProbity run, but that important functionality has now been recaptured with a modified version of the Javascript NGL Viewer.

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.

Understanding the mechanisms and genes that enable animal populations to adapt to pathogens is important from an evolutionary, health and conservation perspective. Berthelot's pipit (Anthus berthelotii) experiences extensive and consistent spatial heterogeneity in avian pox infection pressure across its range of island populations, thus providing an excellent system with which to examine how pathogen-mediated selection drives spatial variation in immunogenetic diversity. Here, we test for evidence of genetic variation associated with avian pox at both an individual and population-level. At the individual level, we find no evidence that variation in MHC class I and TLR4 (both known to be important in recognising viral infection) was associated with pox infection within two separate populations. However, using genotype-environment association (Bayenv) in conjunction with genome-wide (ddRAD-seq) data, we detected strong associations between population-level avian pox prevalence and allele frequencies of single nucleotide polymorphisms (SNPs) at a number of sites across the genome. These sites were located within genes involved in cellular stress signalling and immune responses, many of which have previously been associated with responses to viral infection in humans and other animals. Consequently, our analyses indicate that pathogen-mediated selection may play a role in shaping genomic variation among relatively recently colonised island bird populations and highlight the utility of genotype-environment associations for identifying candidate genes potentially involved in host-pathogen interactions., We are grateful to the Regional Governments of the Canary Islands (Ref.: 2019/5555), the Cabildo of Lanzarote (Ref.: 101/2019), the Cabildo of Tenerife (Ref.: 2019-01740), and the National Parks of El Teide (Ref.: MDV/amp) and Madeira (Ref.: 06/IFCN/2020) for their permission to sample and ring Berthelot’s pipits for this research, the Regional Governments of the Canary Islands and Madeira for providing accommodation, and the Portuguese Navy for transport to Selvagem Grande and Deserta Grande. The Bayenv analysis and associated bioinformatics performed in this study were carried out on the High Performance Computing Cluster supported by the Research and Specialist Computing Support service at the University of East Anglia. This work was supported by Natural Environment Research Council (NERC) studentships awarded to ECS through the ARIES DTP (NE/S007334/1), and to CAM and CA through the EnvEAST DTP (NE/L002582/1), a PhD grant awarded to CG-Q from COLCIENCIAS (Departamento Administrativo de Ciencia, Tecnología e Innovación, Colombia), and a Heredity Fieldwork Grant from the Genetics Society awarded to CAM.

We have curated a high-quality, "best parts" reference dataset of about 3 million protein residues in about 15,000 PDB-format coordinate files, each containing only residues with good electron density support for a physically acceptable model conformation. The resulting pre-filtered data typically contains the entire core of each chain, in quite long continuous fragments. Each reference file is a single protein chain, and the total set of files were selected for low redundancy, high resolution, good MolProbity score, and other chain-level criteria. Then each residue was critically tested for adequate local map quality to firmly support its conformation, which must also be free of serious clashes or covalent-geometry outliers. The resulting Top2018 pre-filtered datasets have been released on the Zenodo online web service and are freely available for all uses under a Creative Commons license. Currently, one dataset is residue-filtered on mainchain plus Cβ atoms, and a second dataset is full-residue filtered; each is available at 4 different sequence-identity levels. Here, we illustrate both statistics and examples that show the beneficial consequences of residue-level filtering. That process is necessary because even the best of structures contain a few highly disordered local regions with poor density and low-confidence conformations that should not be included in reference data. Therefore the open distribution of these very large, pre-filtered reference datasets constitutes a notable advance for structural bioinformatics and the fields that depend upon it. This article is protected by copyright. All rights reserved.

We have curated a high-quality, “best parts” reference dataset of about 3 million protein residues in about 15,000 PDB-format coordinate files, each containing only residues with good electron density support for a physically acceptable model conformation. The resulting pre-filtered data typically contains the entire core of each chain, in quite long continuous fragments. Each reference file is a single protein chain, and the total set of files were selected for low redundancy, high resolution, good MolProbity score, and other chain-level criteria. Then each residue was critically tested for adequate local map quality to firmly support its conformation, which must also be free of serious clashes or covalent-geometry outliers. The resulting Top2018 pre-filtered datasets have been released on the Zenodo online web service and is freely available for all uses under a Creative Commons license. Currently, one dataset is residue-filtered on mainchain plus Cβ atoms, and a second dataset is full-residue filtered; each is available at 4 different sequence-identity levels. Here, we illustrate both statistics and examples that show the beneficial consequences of residue-level filtering. That process is necessary because even the best of structures contain a few highly disordered local regions with poor density and low-confidence conformations that should not be included in reference data. Therefore the open distribution of these very large, pre-filtered reference datasets constitutes a notable advance for structural bioinformatics and the fields that depend upon it.The Top2018 dataset provides the first representative sample of 3D protein structure for which excellence of experimental data constrains the detailed local conformation to be correct for essentially all 3 million residues included. Earlier generations of residue-filtered datasets were central in developing MolProbity validation used worldwide, and now Zenodo has enabled anyone to use out latest version as a sound basis for structural bioinformatics, protein design, prediction, improving biomedically important structures, or other applications.

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.

SummaryThe preservation of genetic diversity is an important aspect of conservation biology. Low genetic diversity within a population can lead to inbreeding depression and a reduction in adaptive potential, which may increase extinction risk. Here we report changes in genetic diversity over 12 years in a declining population of the CorncrakeCrex crex, a grassland bird species of high conservation concern throughout Europe. Despite a twofold demographic decline during the same period, we found no evidence for a reduction of genetic diversity. The gradual genetic differentiation observed among populations of Corncrake across Europe suggests that genetic diversity is maintained in western populations by constant gene flow from the larger and more productive populations in eastern Europe and Asia. The maintenance of genetic diversity in this species is an opportunity that may help the implementation of effective conservation actions across the Corncrake’s European range.

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.

In cooperatively breeding species, helper aid may affect dominant breeders’ investment trade-offs between current and future reproduction. By compensating for the care provided by helpers, breeders can reduce the costs of reproduction and improve chances of survival. Also, helper care can be additive to that of dominants, resulting in higher success of the current brood.However, the influence of helpers on offspring care itself may be the by-product of group size and territory quality. Therefore to make conclusive inferences about causation of additive and compensatory care as a result of helpper serequires disentangling the impact of helping from other factors determining parental investment.In this study, we use 20 years of offspring provisioning data to investigate the effect of helping on breeder and overall offspring provisioning rates in the facultative cooperatively breeding Seychelles warbler (Acrocephalus sechellensis). Our extensive dataset allowed us to effectively control for the effects of living in a larger group and in territories with higher food availability.We show compensatory and additive care in response to helper aid. Helpers lightened the provisioning load of the dominant male and female and increased the total provisioning to the nestlings. This was irrespective of group size or territory quality (food availability).Our results illustrate how multiple benefits of helping behaviour can simultaneously be fundamental to the evolutionary maintenance of cooperative behaviour.

Genetic bottlenecks can limit the success of populations colonizing new ranges. However, successful colonizations can occur despite bottlenecks, a phenomenon known as the genetic paradox of invasion. Eusocial Hymenoptera such as bumblebees (Bombusspp.) should be particularly vulnerable to genetic bottlenecks, since homozygosity at the sex-determining locus leads to costly diploid male production (DMP). The Tree Bumblebee (Bombus hypnorum) has rapidly colonized the UK since 2001 and has been highlighted as exemplifying the genetic paradox of invasion. Using microsatellite genotyping, combined with the first genetic estimates of DMP in UKB. hypnorum, we tested two alternative genetic hypotheses (‘bottleneck’ and ‘gene flow’ hypotheses) forB. hypnorum's colonization of the UK. We found that the UK population has not undergone a recent severe genetic bottleneck and exhibits levels of genetic diversity falling between those of widespread and range-restrictedBombusspecies. Diploid males occurred in 15.4% of reared colonies, leading to an estimate of 21.5 alleles at the sex-determining locus. Overall, the findings show that this population is not bottlenecked, instead suggesting that it is experiencing continued gene flow from the continental European source population with only moderate loss of genetic diversity, and does not exemplify the genetic paradox of invasion.

The population of the Mauritian pink pigeon (Nesoenas mayeri) fell to fewer than 20 individuals in the 1970s. Following intensive conservation efforts, the free-living population is now estimated to be 470 individuals. However, because of the population bottleneck the species remains at risk of extinction because of genetic loss and inbreeding depression. A European captive population was established in 1977 and a European Endangered Species Program (EEP) was formalized in 1992. As birds in the EEP captive population possess unique alleles not observed in the surviving free-living birds, the EEP management plan recommends transferring EEP birds to Mauritius to improve genetic diversity. Health screening of the current EEP population to identify circulating pathogens was performed. Forty-two birds from three collections in the United Kingdom and one in Jersey were screened for a wide range of pathogens, present clinically or subclinically, including important viruses, bacteria, protozoa, and helminths. Eleven birds tested positive for at least one pathogen: Trichomonas spp. (5), Yersinia kristensenii (2), Yersinia aleksiciae (1), coccidial oocysts (3), and strongyle ova (3). None of the positive birds showed overt signs of clinical disease, although two birds with Trichomonas spp. had suboptimal body condition. Genotyping of one Trichomonas gallinae sample revealed a type-C strain (low pathogenicity). The results from this screening will contribute towards a disease risk assessment, to create a pre-export protocol for translocation of captive EEP birds to Mauritius.

Understanding trade‐offs in wild populations is difficult, but important if we are to understand the evolution of life histories and the impact of ecological variables upon them. Markers that reflect physiological state and predict future survival would be of considerable benefit to unraveling such trade‐offs and could provide insight into individual variation in senescence. However, currently used markers often yield inconsistent results. One underutilized measure is hematocrit, the proportion of blood comprising erythrocytes, which relates to the blood's oxygen‐carrying capacity and viscosity, and to individual endurance. Hematocrit has been shown to decline with age in cross‐sectional studies (which may be confounded by selective appearance/disappearance). However, few studies have tested whether hematocrit declines within individuals or whether low hematocrit impacts survival in wild taxa. Using longitudinal data from the Seychelles warbler (Acrocephalus sechellensis), we demonstrated that hematocrit increases with age in young individuals (, Using long‐term data we investigated the association between haematocrit, longitudinal age and survival in a wild vertebrate population—the Seychelles warbler (Acrocephalus sechellensis). Haematocrit exhibited a distinctive age‐dependent pattern within‐individuals, and young individuals with high haematocrit had poorer survival prospects. This study highlights a novel physiological change with age that is indicative of individual condition in wild populations.