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Telomeres are a popular biomarker of senescence, as telomere dynamics are linked with survival and lifespan. However, the evolutionary potential of telomere dynamics, and the selection pattern that gives rise to senescence, are not well known. To better understand this, it is necessary to quantify genetic variation in telomere length, and how such variation changes with age. Here, we analysed a longitudinal dataset (2,156 samples from 1,267 individuals across 15 years) from a wild, insular house sparrow (Passer domesticus) population with complete life-history and genetic relatedness data. Using a series of ‘animal’ models, we confirmed that telomere length (TL): (1) changes with age, reflecting senescence in this population. We then showed that (2) TL is repeatable (15.7%, 95% CrI: 10.2–21.0%) and heritable (14.0%, 95% CrI: 8.7–19.5%); and, for the first time in the wild, (3) that TL shows a genotype-by-age interaction, meaning that genotypes differ in their rate of TL change, where additive genetic variance increases at older ages. Our findings provide empirical evidence from a wild population that supports hypotheses explaining the evolution of senescence, and highlights the importance of telomere dynamics as a key biomarker of body physiology for the evolution of senescence.

Aim: Parental care improves the survival of offspring and therefore has a major impact on reproductive success. It is increasingly recognized that coordinated biparental care is necessary to ensure the survival of offspring in hostile environments, but little is known about the influence of environmental fluctuations on parental cooperation. Assessing the impacts of environmental stochasticity, however, is essential for understanding how populations will respond to climate change and the associated increasing frequencies of extreme weather events. Here we investigate the influence of environmental stochasticity on biparental incubation in a cosmopolitan ground-nesting avian genus. Location: Global. Methods: We assembled data on biparental care in 36 plover populations (Charadrius spp.) from six continents, collected between 1981 and 2012. Using a space-for-time approach we investigate how average temperature, temperature stochasticity (i.e. year-to-year variation) and seasonal temperature variation during the breeding season influence parental cooperation during incubation. Results: We show that both average ambient temperature and its fluctuations influence parental cooperation during incubation. Male care relative to female care increases with both mean ambient temperature and temperature stochasticity. Local climatic conditions explain within-species population differences in parental cooperation, probably reflecting phenotypic plasticity of behaviour. Main conclusions: The degree of flexibility in parental cooperation is likely to mediate the impacts of climate change on the demography and reproductive behaviour of wild animal populations.

We have developed a new marker (Z43B) that can be successfully used to identify the sex of most birds (69%), including species difficult or impossible to sex with other markers. We utilized the zebra finch Taeniopygia guttata EST microsatellite sequence (CK309496) which displays sequence homology to the 5' untranslated region (UTR) of the avian spindlin gene. This gene is known to be present on the Z and W chromosomes. To maximize cross-species utility, the primer set was designed from a consensus sequence created from homologs of CK309496 that were isolated from multiple distantly related species. Both the forward and reverse primer sequences were 100% identical to 14 avian species, including the Z chromosome of eight species and the chicken Gallus gallus W chromosome, as well as the saltwater crocodile Crocodylus porosus. The Z43B primer set was assessed by genotyping individuals of known sex belonging to 61 non-ratite species and a single ratite. The Z and W amplicons differed in size making it possible to distinguish between males (ZZ) and females (ZW) for the majority (69%) of non-ratite species tested, comprising 10 orders of birds. We predict that this marker will be useful for obtaining sex-typing data for ca 6,869 species of birds (69% of non-ratites but not galliforms). A wide range of species could be sex-typed including passerines, shorebirds, eagles, falcons, bee-eaters, cranes, shags, parrots, penguins, ducks, and a ratite species, the brown kiwi, Apteryx australis. Those species sexed include species impossible or problematic to sex-type with other markers (magpie, albatross, petrel, eagle, falcon, crane, and penguin species). Zoo Biol. 35:533-545, 2016. © 2016 The Authors. Zoo Biology published by Wiley Periodicals, Inc.

A new primer set was developed for sex-typing birds, Z37B. This primer set was designed to amplify al- leles of small size to render it suitable for sex-typing de- graded samples, including shed feathers. This marker successfully sex-typed 50 % of the species tested, includ- ing passerines, shorebirds, rails, seabirds, eagles and the brown kiwi Apteryx australis (allele size range =81-103 bp), and is therefore expected to be suitable for sex-typing a wide range of species. Z37B sex-typed non- degraded samples (blood), degraded tissue (dead unhatched embryos, dead nestlings and museum specimens) and samples of low quantity DNA (plucked feathers and buccal swabs). The small amplicon sizes in birds suggest that this marker will be of utility for sex-typing feathers, swabs and degraded samples from a wide range of avian species.

Adult sex ratio (ASR) is a central concept in population biology and a key factor in sexual selection, but why do most demographic models ignore sex biases? Vital rates often vary between the sexes and across life history, but their relative contributions to ASR variation remain poorly understood—an essential step to evaluate sex ratio theories in the wild and inform conservation. Here, we combine structured two-sex population models with individual-based mark–recapture data from an intensively monitored polygamous population of snowy plovers. We show that a strongly male-biased ASR (0.63) is primarily driven by sex-specific survival of juveniles rather than adults or dependent offspring. This finding provides empirical support for theories of unbiased sex allocation when sex differences in survival arise after the period of parental investment. Importantly, a conventional model ignoring sex biases significantly overestimated population viability. We suggest that sex-specific population models are essential to understand the population dynamics of sexual organisms: reproduction and population growth are most sensitive to perturbations in survival of the limiting sex. Overall, our study suggests that sex-biased early survival may contribute toward mating system evolution and population persistence, with implications for both sexual selection theory and biodiversity conservation.

The Chestnut‐banded Plover Charadrius pallidus is a Near‐Threatened shorebird species endemic to mainland Africa. We examined levels of genetic differentiation between its two morphologically and geographically distinct subspecies, C. p. pallidus in southern Africa (population size 11 000–16 000) and C. p. venustus in eastern Africa (population size 6500). In contrast to other plover species that maintain genetic connectivity over thousands of kilometres across continental Africa, we found profound genetic differences between remote sampling sites. Phylogenetic network analysis based on four nuclear and two mitochondrial gene regions, and population genetic structure analyses based on 11 microsatellite loci, indicated strong genetic divergence, with 2.36% mitochondrial sequence divergence between individuals sampled in Namibia (southern Africa) and those of Kenya and Tanzania (eastern Africa). This distinction between southern and eastern African populations was also supported by highly distinct genetic clusters based on microsatellite markers (global FST = 0.309, GST′ = 0.510, D = 0.182). Behavioural factors that may promote genetic differentiation in this species include habitat specialization, monogamous mating behaviour and sedentariness. Reliance on an extremely small number of saline lakes for breeding and limited dispersal between populations are likely to promote reproductive and genetic isolation between eastern and southern Africa. We suggest that the two Chestnut‐banded Plover subspecies may warrant elevation to full species status. To assess this distinction fully, additional sample collection will be needed, with analysis of genetic and phenotypic traits from across the species’ entire breeding range.

Sex-related variation in survival is common in birds and, as it influences effective population size and population growth, is important for conservation and species management. Here we assessed incubation behaviour and sex-related survival in a threatened sexually monomorphic shorebird, the St Helena Plover Charadrius sanctaehelenae. Males incubated at night, the period of highest activity of cats, which are likely to be predators of breeding birds. In spite of behavioural differences between the sexes, adult survival was not significantly different between males and females, suggesting that sex-biased behaviour need not lead to sex-related survival, and thus behavioural differences may not impact upon the adult sex ratio.

Sexual selection may act as a promotor of speciation since divergent mate choice and competition for mates can rapidly lead to reproductive isolation. Alternatively, sexual selection may also retard speciation since polygamous individuals can access additional mates by increased breeding dispersal. High breeding dispersal should hence increase gene flow and reduce diversification in polygamous species. Here, we test how polygamy predicts diversification in shorebirds using genetic differentiation and subspecies richness as proxies for population divergence. Examining microsatellite data from 79 populations in 10 plover species (Genus: Charadrius) we found that polygamous species display significantly less genetic structure and weaker isolation‐by‐distance effects than monogamous species. Consistent with this result, a comparative analysis including 136 shorebird species showed significantly fewer subspecies for polygamous than for monogamous species. By contrast, migratory behavior neither predicted genetic differentiation nor subspecies richness. Taken together, our results suggest that dispersal associated with polygamy may facilitate gene flow and limit population divergence. Therefore, intense sexual selection, as occurs in polygamous species, may act as a brake rather than an engine of speciation in shorebirds. We discuss alternative explanations for these results and call for further studies to understand the relationships between sexual selection, dispersal, and diversification.

We have developed a new marker (Z43B) that can be successfully used to identify the sex of most birds (69%), including species difficult or impossible to sex with other markers. We utilized the zebra finch Taeniopygia guttata EST microsatellite sequence (CK309496) which displays sequence homology to the 5' untranslated region (UTR) of the avian spindlin gene. This gene is known to be present on the Z and W chromosomes. To maximize cross-species utility, the primer set was designed from a consensus sequence created from homologs of CK309496 that were isolated from multiple distantly related species. Both the forward and reverse primer sequences were 100% identical to 14 avian species, including the Z chromosome of eight species and the chicken Gallus gallus W chromosome, as well as the saltwater crocodile Crocodylus porosus. The Z43B primer set was assessed by genotyping individuals of known sex belonging to 61 non-ratite species and a single ratite. The Z and W amplicons differed in size making it possible to distinguish between males (ZZ) and females (ZW) for the majority (69%) of non-ratite species tested, comprising 10 orders of birds. We predict that this marker will be useful for obtaining sex-typing data for ca 6,869 species of birds (69% of non-ratites but not galliforms). A wide range of species could be sex-typed including passerines, shorebirds, eagles, falcons, bee-eaters, cranes, shags, parrots, penguins, ducks, and a ratite species, the brown kiwi, Apteryx australis. Those species sexed include species impossible or problematic to sex-type with other markers (magpie, albatross, petrel, eagle, falcon, crane, and penguin species). Zoo Biol. 35:533-545, 2016. © 2016 The Authors. Zoo Biology published by Wiley Periodicals, Inc.

Three strikingly different alternative male mating morphs (aggressive 'independents', semicooperative 'satellites' and female-mimic 'faeders') coexist as a balanced polymorphism in the ruff, Philomachus pugnax, a lek-breeding wading bird(1-3). Major differences in body size, ornamentation, and aggressive and mating behaviors are inherited as an autosomal polymorphism(4,5). We show that development into satellites and faeders is determined by a supergene(6-8) consisting of divergent alternative, dominant and non-recombining haplotypes of an inversion on chromosome 11, which contains 125 predicted genes. Independents are homozygous for the ancestral sequence. One breakpoint of the inversion disrupts the essential CENP-N gene (encoding centromere protein N), and pedigree analysis confirms the lethality of homozygosity for the inversion. We describe new differences in behavior, testis size and steroid metabolism among morphs and identify polymorphic genes within the inversion that are likely to contribute to the differences among morphs in reproductive traits.

Exploring the patterns of genetic structure in the context of geographical and phenotypic variation is important to understand the evolutionary processes involved in speciation. We investigated population and subspecies differentiation in the Common Ringed Plover Charadrius hiaticula, a high latitude wader that breeds in arctic and temperate zones from northeast Canada across Eurasia to the Russian Far East. Three subspecies, hiaticula, tundrae and psammodromus, are currently widely recognised, whereas a fourth subspecies, kolymensis, has been proposed based on geographic isolation and phenotypic differences. We genotyped 173 samples from eleven Common Ringed Plover breeding sites, representing all four putative subspecies, at eight polymorphic microsatellite loci to examine the patterns of population and subspecies differentiation. Bayesian clustering identified three genetic clusters among samples, corresponding to the breeding sites of the three currently recognised subspecies. The existence of the subspecies kolymensis was not supported. We also detected the presence of a previously unknown hybridisation zone extending from Northern Scandinavia to Belarus. Differentiation of the subspecies tundrae and hiaticula most likely occurred in allopatry on the Eurasian continent during past glaciation events, followed by population expansion leading to colonisation of Iceland and Greenland. The lack of genetic differentiation within the tundrae subspecies is consistent with ongoing range expansion and high gene flow maintained through migratory behaviour. We discuss the importance of historic climate changes, migratory behaviour and mating system on shaping the observed pattern of genetic differentiation.

Establishing phylogenetic relationships within a clade can help to infer ancestral origins and indicate how widespread species reached their current biogeographic distributions. The small plovers, genus Charadrius, are cosmopolitan shorebirds, distributed across all continents except Antarctica. Here we present a global, species-level molecular phylogeny of this group based on four nuclear (ADH5, FIB7, MYO2 and RAG1) and two mitochondrial (COI and ND3) genes, and use the phylogeny to examine the biogeographic origin of the genus. A Bayesian multispecies coalescent approach identified two major clades (CRD I and CRD II) within the genus. Clade CRD I contains three species (Thinornis novaeseelandiae, Thinornis rubricollis and Eudromias morinellus), and CRD II one species (Anarhynchus frontalis), that were previously placed outside the Charadrius genus. In contrast to earlier work, ancestral area analyses using parsimony and Bayesian methods supported an origin of the Charadrius plovers in the Northern hemisphere. We propose that major radiations in this group were associated with shifts in the range of these ancestral plover species, leading to colonisation of the Southern hemisphere.