Reproductive performance of resident and migrant males, females and pairs in a partially migratory bird

We thank everyone from the Centre for Ecology & Hydrology (CEH) who contributed to data collection, and Scottish Natural Heritage for access to the Isle of May National Nature Reserve. We thank the Scottish Ornithologists’ Club (SOC) for their support, and all volunteer observers, particularly Raymond Duncan, Moray Souter and Bob Swann. HG was funded by a Natural Environment Research Council (NERC) CASE studentship supported by CEH and SOC, FD, SW, MPH, MN and SB were funded by NERC and the Joint Nature Conservation Committee, and JMR was part-funded by the Royal Society. Finally, we thank the Associate Editor and two reviewers for constructive comments on the manuscript. The data are available from the Dryad Digital Repository https://doi.org/10.5061/dryad.532j0 (Grist et al., 2017)Peer reviewedPublisher PD

Assessing the vulnerability of the marine bird community in the western North Sea to climate change and other anthropogenic impacts

Ocean warming and anthropogenic activities such as fishing, shipping and marine renewable developments are affecting marine top predators. Research has focussed on the impacts of single stressors on single species, yet understanding cumulative effects of multiple stressors on communities is vital for effective conservation management. We studied a marine bird community (45 species; 11 families) that used the Forth and Tay region of the North Sea for breeding, overwintering or migration between 1980 and 2011. Local sea surface temperature (SST) increased significantly over this period, with concomitant changes in lower trophic levels. Simultaneously, the region has been subject to fishing pressure and shipping disturbance and is a priority area for renewable energy developments. We used colony-based and at-sea data to quantitatively assess relationships between SST and counts, productivity and survival of 25 species for which sufficient data were available for analysis. For the remaining species, we applied a qualitative approach using published population trends, published climate relationships and foraging sensitivity. In total, 53% of species showed negative relationships with SST. Trends in counts and demography were combined with climate vulnerability to give an index of population concern to future climate warming, and 44% of species were classified as high or very high concern, notably cormorants, grebes, skuas, shearwaters, terns and auks, as well as species breeding in the region. Qualitative assessments of vulnerability to fisheries, pollutants, disturbance (including introduced predators), marine renewables and climate found that 93% of species were vulnerable to ≥2 threats, and 58% to ≥4. Our results indicate that the majority of birds in this region of the North Sea face an uncertain future, potentially threatening the resilience of this important marine bird community

Among-individual and within-individual variation in seasonal migration covaries with subsequent reproductive success in a partially migratory bird

Acknowledgements: We thank Raymond Duncan (Grampian Ringing Group), whose insatiable enthusiasm for ringing was inspirational. We thank everyone who contributed to fieldwork, especially Mike Harris and David Hunter; Roger Pradel for generous advice on CMR mixture models; and UK Natural Environment Research Council for funding (NE/R000859/1, NE/P009719/1). Data accessibility Data are available from the Dryad Digital Repository: https://doi.org/10.5061/dryad.pvmcvdnhv [46]. Electronic supplementary material is available online at https://doi.org/10.6084/m9.figshare.c.5054649.Peer reviewedPostprin

Contrasting responses of male and female foraging effort to year-round wind conditions

There is growing interest in the effects of wind on wild animals, given evidence that wind speeds are increasing and becoming more variable in some regions, particularly at temperate latitudes. Wind may alter movement patterns or foraging ability, with consequences for energy budgets and, ultimately, demographic rates. These effects are expected to vary among individuals due to intrinsic factors such as sex, age or feeding proficiency. Furthermore, this variation is predicted to become more marked as wind conditions deteriorate, which may have profound consequences for population dynamics as the climate changes. However, the interaction between wind and intrinsic effects has not been comprehensively tested. In many species, in particular those showing sexual size dimorphism, males and females vary in foraging performance. Here, we undertook year-round deployments of data loggers to test for interactions between sex and wind speed and direction on foraging effort in adult European shags Phalacrocorax aristotelis, a pursuit-diving seabird in which males are c. 18% heavier. We found that foraging time was lower at high wind speeds but higher during easterly (onshore) winds. Furthermore, there was an interaction between sex and wind conditions on foraging effort, such that females foraged for longer than males when winds were of greater strength (9% difference at high wind speeds vs. 1% at low wind speeds) and when winds were easterly compared with westerly (7% and 4% difference, respectively). The results supported our prediction that sex-specific differences in foraging effort would become more marked as wind conditions worsen. Since foraging time is linked to demographic rates in this species, our findings are likely to have important consequences for population dynamics by amplifying sex-specific differences in survival rates

Sublethal effects of natural parasitism act through maternal, but not paternal; reproductive success in a wild population

Parasites are a major component of all animal populations. Males and females often differ in their levels of parasite prevalence, potentially leading to sex differences in the impact of parasitism on fitness, with important implications for the evolution of parasite and host traits including resistance, tolerance, and virulence. However, quantitative measures of the impact of parasitism under free‐living conditions are extremely rare, as they require detailed host demographic data with measures of parasite burden over time. Here, we use endoscopy for direct quantification of natural‐parasite burdens and relate these to reproductive success over 7 yr in a wild population of seabirds. Contrary to predictions, only female burdens were associated with negative impacts of parasitism on breeding success, despite males having significantly higher burdens. Female reproductive success declined by 30% across the range of natural parasite burdens. These effects persisted when accounting for interannual population differences in breeding success. Our results provide quantitative estimates of profound sub‐lethal effects of parasitism on the population. Importantly, they highlight how parasites act unpredictably to shape ecological and evolutionary processes in different components of the same population, with implications for demography and selection on host and parasite traits

Parental age influences offspring telomere loss

1. The age of the parents at the time of offspring production can influence offspring longevity, but the underlying mechanisms remain poorly understood. The effect of parental age on offspring telomere dynamics (length and loss rate) is one mechanism that could be important in this context. 2. Parental age might influence the telomere length that offspring inherit or age-related differences in the quality of parental care could influence the rate of offspring telomere loss. However, these routes have generally not been disentangled. 3. Here, we investigated whether parental age was related to offspring telomere dynamics using parents ranging in age from 2 to 22 years old in a free-living population of a long-lived seabird, the European shag (Phalacrocorax aristotelis). By measuring the telomere length of offspring at hatching and towards the end of the post-natal growth period, we could assess whether any potential parental age effect was confined to the post-natal rearing period. 4. There was no effect of maternal or paternal age on the initial telomere length of their chicks. However, chicks produced by older mothers and fathers experienced significantly greater telomere loss during the post-natal nestling growth period. We had relatively few nests in which the ages of both parents were known, and individuals in this population mate assortatively with respect to age. Thus, we could not conclusively determine whether the parental age effect was due to maternal age, paternal age, or both; however, it appears that the effect is stronger in mothers. 5. These results demonstrate that in this species, there was no evidence that parental age was related to offspring hatching telomere length. However, telomere loss during nestling growth was reduced in the offspring of older parents. This could be due to an age-related deterioration in the quality of the environment that parents provide, or because parents that invest less in offspring rearing live to an older age

The genome sequence of the European shag, Gulosus aristotelis (previously Phalacrocorax aristotelis) (Linnaeus, 1761)

We present a genome assembly from an individual female Gulosus aristotelis, previously known as Phalacrocorax aristotelis, (the European shag; Chordata; Aves; Pelecaniformes; Phalacrocoracidae). The genome sequence is 1,279.1 megabases in span. Most of the assembly is scaffolded into 36 chromosomal pseudomolecules, including the Z and W sex chromosomes. The mitochondrial genome has also been assembled and is 18.61 kilobases in length. Gene annotation of this assembly on Ensembl identified 16,474 protein coding genes

The energetic cost of parasitism in a wild population

Parasites have profound fitness effects on their hosts, yet these are often sub-lethal, making them difficult to understand and quantify. A principal sub-lethal mechanism that reduces fitness is parasite-induced increase in energetic costs of specific behaviours, potentially resulting in changes to time and energy budgets. However, quantifying the influence of parasites on these costs has not been undertaken in free-living animals. We used accelerometers to estimate energy expenditure on flying, diving and resting, in relation to a natural gradient of endo-parasite loads in a wild population of European shags Phalacrocorax aristotelis. We found that flight costs were 10% higher in adult females with higher parasite loads and these individuals spent 44% less time flying than females with lower parasite loads. There was no evidence for an effect of parasite load on daily energy expenditure, suggesting the existence of an energy ceiling, with the increase in cost of flight compensated for by a reduction in flight duration. These behaviour specific costs of parasitism will have knock-on effects on reproductive success, if constraints on foraging behaviour detrimentally affect provisioning of young. The findings emphasize the importance of natural parasite loads in shaping the ecology and life-history of their hosts, which can have significant population level consequences

Additive genetic and environmental variation interact to shape the dynamics of seasonal migration in a wild bird population

We thank everyone who contributed to long-term field data collection, particularly Raymond Duncan, Sarah Fenn, Hannah Grist, Calum Scott, Jenny Sturgeon, Moray Souter, John Anderson, and Harry Bell; and thank NatureScot for allowing work on the Isle of May National Nature Reserve, and Isle of May Bird Observatory Trust for supporting the longterm ringing of shags. We thank Stefanie Muff for helpful discussions, and Rita Fortuna and Thomas R. Haaland for useful comments on a manuscript draft. The current study was funded by Natural Environment Research Council (NERC; awards NE/M005186/1, NE/R000859/1, and NE/R016429/1 as part of the UK-SCaPE program delivering National Capability), Norwegian Research Council (SFF-III grant 223257, FRIPRO grant 313570), NTNU and University of Aberdeen. Analyses were performed using the IDUN cluster of NTNUPeer reviewedPublisher PD

Carry‐over effects of seasonal migration on reproductive success through breeding site retention in a partially migratory bird

•1. Understanding the maintenance and dynamics of phenotypic polymorphisms requires unpicking key ecological mechanisms shaping the fitness costs and benefits of expressing alternative phenotypes, generating selection. Seasonal migration versus year-round residence expressed in partially migratory populations represents one common polymorphism that can experience strong selection through differential reproductive success. Yet, key hypothesised pathways that could generate such selection remain to be empirically tested. •2. One hypothesis is that migratory tactics affect subsequent reproductive success through carry-over effects on breeding site retention and resulting breeding dispersal. By remaining in breeding areas all year round, residents could retain their preferred breeding site between years, and consequently have higher reproductive success. Conversely, migrants that escape harsh non-breeding season conditions could return in better condition, with high resource holding potential, and outcompete residents to retain their site. Such effects could further depend on migration timing and vary between years. Yet, such pathways have not been quantified, precluding empirical parameterisation of partial migration theory. •3. We used 4 years of breeding and non-breeding season data from partially migratory European shags (Gulosus aristotelis) to test whether the three most frequent migratory tactics in this population (full resident, early migrant departing soon after breeding, and late migrant departing in late autumn) differed in their breeding site retention; whether site retention predicted reproductive success; and hence whether effects of migratory tactic on reproductive success were explicable through site retention. •4. Overall, residents were much more likely to retain their breeding site between years than both early and late migrants, and site retention was associated with increased reproductive success. Yet, these effects varied somewhat among years: late migrants were always least likely to retain their site but had variable relative reproductive success. Path analyses revealed that effects of migratory tactic on reproductive success were only partly attributable to breeding site retention. •5. These results indicate that multiple mechanisms underlie reproductive selection on migratory tactics, potentially contributing to maintaining behavioural polymorphisms. Yet, the clear associations between migratory tactics and local breeding dispersal reveal that these movements can be strongly interlinked across seasons, shaping overall spatioseasonal dynamics in partially migratory systems