Ecosystem structure and biophysical processes mediate biological responses to climate changes, but few studies have examined impacts of this dynamic among upper trophic levels. We investigated ecosystem differences in how diverse seabird populations across the northern hemisphere have responded to changes in regional mixed layer temperature and water column stratification. Using 138 time series of breeding productivity over the past half-century, we show that seabird reproductive productivity has declined in the Arctic and North Atlantic but not in the Pacific during a period of ubiquitous mixed layer warming and regionally-variable stratification trends. Models of breeding productivity and ocean drivers show that seabird responses to climate change vary by ecosystem. Additionally, ecosystems in which seabirds exhibit detectibly declining productivity tend to have lower overall diet diversity across seabird species. These findings emphasize the importance of ecosystem processes and structure in determining the vulnerability of marine predators to climate change.

change affects timing of reproduction in many

Climate change affects timing of reproduction in many bird species, but few studies have investigated its influence on annual reproductive output. Here, we assess changes in the annual production of young by female breeders in 201 populations of 104 bird species (N = 745,962 clutches) covering all continents between 1970 and 2019. Overall, average offspring production has declined in recent decades, but considerable differences were found among species and populations. A total of 56.7% of populations showed a declining trend in offspring production (significant in 17.4%), whereas 43.3% exhibited an increase (significant in 10.4%). The results show that climatic changes affect offspring production through compounded effects on ecological and life history traits of species. Migratory and larger-bodied species experienced reduced offspring production with increasing temperatures during the chick-rearing period, whereas smaller-bodied, sedentary species tended to produce more offspring. Likewise, multi-brooded species showed increased breeding success with increasing temperatures, whereas rising temperatures were unrelated to reproductive success in single-brooded species. Our study suggests that rapid declines in size of bird populations reported by many studies from different parts of the world are driven only to a small degree by changes in the production of young.

Timing of breeding, an important driver of fitness in many populations, is widely studied in the context of global change, yet despite considerable efforts to identify environmental drivers of seabird nesting phenology, for most populations we lack evidence of strong drivers. Here we adopt an alternative approach, examining the degree to which different populations positively covary in their annual phenology to infer whether phenological responses to environmental drivers are likely to be (a) shared across species at a range of spatial scales, (b) shared across populations of a species or (c) idiosyncratic to populations. We combined 51 long-term datasets on breeding phenology spanning 50 years from nine seabird species across 29 North Atlantic sites and examined the extent to which different populations share early versus late breeding seasons depending on a hierarchy of spatial scales comprising breeding site, small-scale region, large-scale region and the whole North Atlantic. In about a third of cases, we found laying dates of populations of different species sharing the same breeding site or small-scale breeding region were positively correlated, which is consistent with the hypothesis that they share phenological responses to the same environmental conditions. In comparison, we found no evidence for positive phenological covariation among populations across species aggregated at larger spatial scales. In general, we found little evidence for positive phenological covariation between populations of a single species, and in many instances the inter-year variation specific to a population was substantial, consistent with each population responding idiosyncratically to local environmental conditions. Black-legged kittiwake Rissa tridactyla was the exception, with populations exhibiting positive covariation in laying dates that decayed with the distance between breeding sites, suggesting that populations may be responding to a similar driver. Our approach sheds light on the potential factors that may drive phenology in our study species, thus furthering our understanding of the scales at which different seabirds interact with interannual variation in their environment. We also identify additional systems and phenological questions to which our inferential approach could be applied.

Climate change and other human activities are causing profound effects on marine ecosystem productivity. We show that the breeding success of seabirds is tracking hemispheric differences in ocean warming and human impacts, with the strongest effects on fish-eating, surface-foraging species in the north. Hemispheric asymmetry suggests the need for ocean management at hemispheric scales. For the north, tactical, climate-based recovery plans for forage fish resources are needed to recover seabird breeding productivity. In the south, lower-magnitude change in seabird productivity presents opportunities for strategic management approaches such as large marine protected areas to sustain food webs and maintain predator productivity. Global monitoring of seabird productivity enables the detection of ecosystem change in remote regions and contributes to our understanding of marine climate impacts on ecosystems.

Telomeres are protective caps at the end of eukaryotic chromosomes that shorten with age and in response to stressful or resource-demanding conditions. Their length predicts individual health and lifespan across a wide range of animals, but whether the observed positive association between telomere length and lifespan is environmentally induced, or set at conception due to a shared genetic basis, has not been tested in wild animals. We applied quantitative genetic "animal models" to longitudinal telomere measurements collected over a 10-year period from individuals of a wild seabird (common tern; Sterna hirundo) with known pedigree. We found no variation in telomere shortening with age among individuals at the phenotypic and genetic level, and only a small permanent environmental effect on adult telomere length. Instead, we found telomere length to be highly heritable and strongly positively genetically correlated with lifespan. Such heritable differences between individuals that are set at conception may present a hitherto underappreciated component of variation in somatic state.

Telomeres are protective caps at the end of chromosomes, and their length is positively correlated with individual health and lifespan across taxa. Longitudinal studies have provided mixed results regarding the within‐individual repeatability of telomere length. While some studies suggest telomere length to be highly dynamic and sensitive to resource‐demanding or stressful conditions, others suggest that between‐individual differences are mostly present from birth and relatively little affected by the later environment. This dichotomy could arise from differences between species, but also from methodological issues. In our study, we used the highly reliable Terminal Restriction Fragment analysis method to measure telomeres over a 10‐year period in adults of a long‐lived seabird, the common tern (Sterna hirundo). Telomeres shortened with age within individuals. The individual repeatability of age‐dependent telomere length was high (>0.53), and independent of the measurement interval (i.e., one vs. six years). A small (R2 = .01), but significant part of the between‐individual variation in telomere length was, however, explained by the number of fledglings produced in the previous year, while reproduction in years prior to the previous year had no effect. We confirmed that age‐dependent telomere length predicted an individual's remaining lifespan. Overall, our study suggests that the majority of between‐individual variation in adult telomere length is consistent across adult life, and that a smaller part of the variation can be explained by dynamic factors, such as reproduction.

Kleptoparasitism is an exploitative foraging strategy used across taxa, but factors underlying variation in the foraging performance of individuals using it have not often been addressed. Using longitudinal data on Common Terns stealing food from conspecifics during breeding, we show that variation in the energetic reward of kleptoparasitic behaviour is explained by interactive effects of sex and the attack tactic used by the parasite, as well as by age. Males obtain a higher reward when using an aerial than a terrestrial attack tactic, with decomposition analyses showing that this is due to the energy content of chased after prey items being higher in their aerial attacks. On the other hand, females obtain a higher reward when attacking terrestrially, which is due to their success rate being higher on land than in the air. In addition, the birds show decelerating within-individual improvement with age, which is due to individuals chasing after prey with a higher energy content as they grow older. Our study not only pinpoints factors underlying variation in the foraging performance of kleptoparasites, but also illustrates the importance of modelling individual variation when analyzing foraging performance.

Maternal and environmental effects can have profound effects on offspring performance by generating variation in offspring phenotypes, independent of genetic effects. Within avian broods, differential maternal investment of resources across the laying sequence is thought to be an adaptive strategy to modulate competitive hierarchies induced by hatching asynchrony. In this study, we evaluated the relative importance of maternally derived within-clutch variation and the asymmetric post-hatching environment for growth and survival of common tern (Sterna hirundo) siblings. We experimentally manipulated hatching order, resulting in chicks from last-laid eggs hatching first and vice versa. Although both initial age and size asymmetries were larger within experimental than control broods, the early survival of last-hatched chicks was similar between groups. Initial positive effects of egg size disappeared as siblings approached fledging. Ultimately at fledging, both within-brood growth and cumulative survival patterns were similar between experimental and control broods, suggesting that the effects of systematic variation of egg constituents (e.g., maternally derived yolk hormones) and egg size are too subtle and largely overwhelmed by the effects of hatching asynchrony. Therefore, we conclude that variation in offspring phenotypes is pre-dominantly determined by the social environment experienced post-hatching. Maternal effects may further fine-tune phenotypic variation in response to varying environmental conditions, but this needs to be tested through empirical studies in which multiple maternal effects are measured simultaneously under different environmental conditions.

Selection is a central force underlying evolutionary change and can vary in strength and direction, for example across time and space. The fitness consequences of individual genetic diversity have often been investigated by testing for multilocus heterozygosity-fitness correlations (HFCs), but few studies have been able to assess HFCs across life stages and in both sexes. Here, we test for HFCs using a 26-year longitudinal individual-based data set from a large population of a long-lived seabird (the common tern, Sterna hirundo), where 7,974 chicks and breeders of known age were genotyped at 15 microsatellite loci and sampled for life-history traits over the complete life cycle. Heterozygosity was not correlated with fledging or post-fledging prospecting probabilities, but was positively correlated with recruitment probability. For breeders, annual survival was not correlated with heterozygosity, but annual fledgling production was negatively correlated with heterozygosity in males and highest in intermediately heterozygous females. The contrasting HFCs among life stages and sexes indicate differential selective processes and emphasize the importance of assessing fitness consequences of traits over complete life histories.

Reproductive timing in many taxa plays a key role in determining breeding productivity 1 , and is often sensitive to climatic conditions 2 . Current climate change may alter the timing of breeding at different rates across trophic levels, potentially resulting in temporal mismatch between the resource requirements of predators and their prey 3 . This is of particular concern for higher-trophic-level organisms, whose longer generation times confer a lower rate of evolutionary rescue than primary producers or consumers 4 . However, the disconnection between studies of ecological change in marine systems makes it difficult to detect general changes in the timing of reproduction 5 . Here, we use a comprehensive meta-analysis of 209 phenological time series from 145 breeding populations to show that, on average, seabird populations worldwide have not adjusted their breeding seasons over time (−0.020 days yr−1) or in response to sea surface temperature (SST) (−0.272 days °C−1) between 1952 and 2015. However, marked between-year variation in timing observed in resident species and some Pelecaniformes and Suliformes (cormorants, gannets and boobies) may imply that timing, in some cases, is affected by unmeasured environmental conditions. This limited temperature-mediated plasticity of reproductive timing in seabirds potentially makes these top predators highly vulnerable to future mismatch with lower-trophic-level resources 2 . Time of reproduction may be altered as the climate changes. For seabirds, it is shown that there has not been an adjustment in timing as the climate changes and the sea surface warms. This lack of plasticity could result in a mismatch with food resources.

In a monogamous species two partners contribute to the breeding process. We study pair formation as well as the effect of pair bond length and age on breeding performance, incorporating individual heterogeneity, based on a high-quality dataset of a long-lived seabird, the common tern (Sterna hirundo). To handle missing information and model the complicated processes driving reproduction, we use a hierarchical Bayesian model of the steps that lead to the number of fledglings, including processes at the individual and the pair level. The results show that the age of both partners is important for reproductive performance, with similar patterns for both sexes and individual heterogeneity in reproductive performance, but pair bond length is not. The terns are more likely to choose a former partner independent of the previous breeding outcome with that partner, which suggests a tendency to retain the partner chosen at the beginning of the breeding career.

Biparental care is found across taxa, but sex-specific contributions vary greatly both between and within species. Factors underlying intraspecific variation may include the phase of the reproductive cycle and sex-specific duties resulting in physiological or environmental constraints that differ between the sexes. For pair members to achieve high breeding success while maintaining good condition, parental duties should be well coordinated between mates. Avian breeding behavior is mediated by hormones such as prolactin and corticosterone, which link the internal and environmental parameters of individuals and regulate resource allocation. Using automated recording at the nest, we analyzed parental attendance by Common Terns (Sterna hirundo) in relation to sex, reproductive phase (incubation, chick rearing), time of day, breeding experience, and hormone levels, and related it to reproductive success. Nest attendance of females exceeded that of males, especially at night and during chick rearing. Greater breeding experience was linked to increased attendance at night by males and during incubation by females. Parental nest attendance during incubation was not related to duration of the incubation phase, nor to hatching success. However, fledging success increased with greater parental attendance during the early chick rearing period, reflecting a higher vulnerability of small chicks to mortality compared with eggs. Prolactin values of male parents were significantly lower during chick rearing than incubation and in comparison with values of female parents. Female breeding experience was positively correlated with increased hormone values during chick rearing. Male attendance during chick rearing covaried with hormone levels: moderately increased corticosterone could support males in their new role as feeders by facilitating high activity and foraging effort, whereas lowered prolactin might indicate less contact with chicks and reduced body weight. We show that Common Terns exhibit a flexible system of sex-specific parental care during incubation and chick rearing, reflected in hormone levels, which mediate breeding behavior and ultimately affect reproductive success.

Kleptoparasitism is a well-known foraging strategy used by many colonial seabirds. The expression of this behavior can reduce the cost of foraging during the breeding season and ultimately this can be reflected in physiological parameters. Here we analyzed the relationship between the trophic strategy of Common Terns Sterna hirundo and two hormones, that are known to be involved in decision-making during the reproductive cycle and foraging activities: corticosterone (CORT) and prolactin (PRL). The study was conducted during the breeding seasons 2012-2014 at the Banter See colony in northern Germany where individuals with different foraging strategies were identified and monitored since 2008. We studied the concentration of hormones in two groups of birds: kleptoparasitic individuals i.e. individuals who steal the food captured by another individual, and honest individuals i.e. individuals who obtain the food on their own. Blood samples were obtained during the mid-incubation period using a non-invasive technique. Both hormones showed no significant relationship to each other. CORT concentration was similar in both groups of birds. However, the concentration of PRL was higher in parasitic than honest individuals in 2014. In addition, CORT modelling indicated a year effect (higher CORT in 2013), and PRL modelling indicated a significant effect of year (higher PRL in 2012) and a significant interaction between year and the trophic strategy in 2014. The results show no clear relationship between the feeding strategy and studied corticosterone and prolactin levels.

Abstract A major question for conservationists and evolutionary biologists is whether natural populations can adapt to rapid environmental change through micro‐evolution or phenotypic plasticity. Making use of 17 years of data from a colony of a long‐distant migratory seabird, the common tern (Sterna hirundo), we examined phenotypic plasticity and the evolutionary potential of breeding phenology, a key reproductive trait. We found that laying date was strongly heritable (0.27 ± 0.09) and under significant fecundity selection for earlier laying. Paradoxically, and in contrast to patterns observed in most songbird populations, laying date became delayed over the study period, by about 5 days. The discrepancy between the observed changes and those predicted from selection on laying date was explained by substantial phenotypic plasticity. The plastic response in laying date did not vary significantly among individuals. Exploration of climatic factors showed individual responses to the mean sea surface temperature in Senegal in December prior to breeding: Common terns laid later following warmer winters in Senegal. For each 1°C of warming of the sea surface in Senegal, common terns delayed their laying date in northern Germany by 6.7 days. This suggests that warmer waters provide poorer wintering resources. We therefore found that substantial plastic response to wintering conditions can oppose natural selection, perhaps constraining adaptation.

Telomere length (TL) is a candidate biomarker of ageing and phenotypic quality, but little is known of the (physiological) causes of TL variation. We previously showed that individual common terns Sterna hirundo with high reproductive success had short telomeres independent of age, and this pattern was particularly strong in the longer telomeres of the within-individual TL distribution. To test whether this relation can be attributed to effects of reproductive effort, we investigated baseline corticosterone in relation to reproductive success (number of fledglings) and TL. In this context, we assume that variation in baseline corticosterone can be interpreted as index of energy expenditure and allostatic load. Males with higher corticosterone levels during incubation, compared between and within individuals, achieved higher reproductive success and had shorter telomeres. The effect on telomeres was more pronounced in corticosterone measured later in incubation and in the longer telomeres of the within-individual TL distribution. Female corticosterone level during incubation was neither related to reproductive success nor to TL. That we observed these effects only in males mirrors different parental roles during reproduction in the common tern, where males do most of the chick provisioning. The negative association between reproductive success and TL suggests individual differences in reproductive effort as reflected in, or mediated by, baseline corticosterone. We see this result as a promising step towards unravelling the physiological causes of variation in TL and the costs of reproduction.

Long term studies, focusing on population- and socio-biology research, require the unequivocal identification of individuals. DNA studies with Short Tandem Repeats (STR loci) became a widespread tool in population genetics. We used the next-generation sequencing (NGS) approach with 454 shot-gun pyrosequencing to identify 13 new polymorphic STR loci for the Common Tern,Sterna hirundo. To enlarge the marker set we added two more loci originally developed for Black-legged Kittiwake (Rissa tridactyla) and Red-billed Gull (Chroicocephalus scopulinus) and arranged these 15 loci into three multiplex PCR panels for high throughput genotyping. Loci characterization demonstrated that our marker set is of high quality. A PIC value of about 0.67 and a power of exclusion value of 0.99 were reached. Deviation from Hardy-Weinberg expectations of some loci and low frequencies for null alleles are interpreted as a result of inbreeding and founder effect in the investigated tern colony. We used a test data set of this well-studied breeding colony of Common Tern at Banter Lake, Wilhelmshaven, Germany, to perform a parentage test. Parent-chick relationships, known from the social pedigree of that colony, were compared with genetically calculated ones. In order to test our markers and the used parentage program COLONY, we conducted six competing data sets with varying completeness of included parental genotypes. By including fully sampled parent pairs of known family assignment, results were correct for nest mates, single parents and parent pairs. Our marker set provides a powerful tool to investigate life-time reproductive success and other issues of population and socio-biology for Common Terns,e.g. in the aforementioned colony monitored for decades.

El objetivo de este trabajo es actualizar la descripcion de una tecnica de obtencion de muestras de sangre en aves y mostrar su eficiencia en una colonia de Gaviotin Golondrina (Sterna hirundo) situada en el lago Banter, Alemania. Las tecnicas utilizadas actualmente involucran la captura y manejo de las aves y la extraccion de sangre con agujas hipodermicas, lo cual puede causar estres en el individuo estudiado. En este trabajo se utilizo una tecnica minimamente invasiva usando insectos hematofagos del orden Heteroptera. Se describen las modificaciones del diseno original de la tecnica y se muestra que existio una alta eficiencia en el muestreo, en terminos de tiempo y numero de muestras tomadas.

We studied the temporal–spatial distribution of Common Terns Sterna hirundo along the East Atlantic Flyway. In 2009 and 2010 experienced adults from a colony on the German North Sea coast were tagged with geolocators recording light intensity and saltwater contact. Main objectives were the inter-individual temporal–spatial variation of migration routes and wintering areas, wintering site fidelity, and time spent at sea across the annual cycle. Geolocators had no effects on various traits of breeders, but their reproductive output suffered from egg breakage. This can be avoided by artificially incubating the eggs. Twelve routes of nine individuals were tracked. Transponder readings at the breeding site showed that birds left the colony 4 weeks before starting autumn migration. In spring and autumn, Common Terns stopped over around the Canary Islands. Main wintering distribution was the upwelling seas alongside the West African coast and similar between years, but different among individuals. Three females wintered further north and more offshore than six males. Pair mates wintered at different locations. Spring migration was longer (56 ± 8 days) than autumn migration (37 ± 17 days). During both migration and wintering the terns spent more time on salt water than during breeding and post-breeding. In most individuals saltwater contact was higher during the day than at night, reduced at sunrise and sunset likely due to foraging, and peaked about noon possibly related to resting or thermoregulation. Detailed ecological and behavioral studies of common terns during wintering are needed to clarify the results based on geolocators.

Optimal sex allocation is frequency-dependent, but senescence may cause behaviour at old age to be suboptimal. We investigated whether sex allocation changes with parental age, using 16 years of data comprising more than 2500 molecularly sexed offspring of more than 600 known-age parents in common terns ( Sterna hirundo ), slightly sexually size-dimorphic seabirds. We decomposed parental age effects into within-individual change and sex allocation-associated selective (dis)appearance. Individual parents did not differ consistently in sex allocation, but offspring sex ratios at fledging changed from female- to male-biased as parents aged. Sex ratios at hatching were not related to parental age, suggesting sons to outperform daughters after hatching in broods of old parents. Our results call for the integration of sex allocation theory with theory on ageing and demography, as a change in sex allocation with age per se will cause the age structure of a population to affect the frequency-dependent benefits and the age-specific strength of selection on sex allocation.