Does seasonality drive spatial patterns in demography? Variation in survival in African reed warblers Acrocephalus baeticatus across southern Africa does not reflect global patterns

Among birds, northern temperate species generally have larger clutches, shorter development periods and lower adult survival than similarly‐sized southern and tropical species. Even though this global pattern is well accepted, the driving mechanism is still not fully understood. The main theories are founded on the differing environmental seasonality of these zones (higher seasonality in the North). These patterns arise in cross‐species comparisons, but we hypothesized that the same patterns should arise among populations within a species if different types of seasonality select for different life histories. Few studies have examined this. We estimated survival of an azonal habitat specialist, the African reed warbler, across the environmentally diverse African subcontinent, and related survival to latitude and to the seasonality of the different environments of their breeding habitats. Data (1998–2010) collected through a public ringing scheme were analyzed with hierarchical capture‐mark‐recapture models to determine resident adult survival and its spatial variance across sixteen vegetation units spread across four biomes. The models were defined as state‐space multi‐state models to account for transience and implemented in a Bayesian framework. We did not find a latitudinal trend in survival or a clear link between seasonality and survival. Spatial variation in survival was substantial across the sixteen sites (spatial standard deviation of the logit mean survival: 0.70, 95% credible interval (CRI): 0.33–1.27). Mean site survival ranged from 0.49 (95% CRI: 0.18–0.80) to 0.83 (95% CRI: 0.62–0.97) with an overall mean of 0.67 (95% CRI: 0.47–0.85). A hierarchical modeling approach enabled us to estimate spatial variation in survival of the African reed warbler across the African subcontinent from sparse data. Although we could not confirm the global pattern of higher survival in less seasonal environments, our findings from a poorly studied region contribute to the study of life‐history strategies.     

Patterns and processes in shorebird survival rates: a global review

Changes in demographic rates underpin changes in population size, and understanding demographic rates can greatly aid the design and development of strategies to maintain populations in the face of environmental changes. However, acquiring estimates of demographic parameters at relevant spatial scales is difficult. Measures of annual survival rates can be particularly challenging to obtain because large‐scale, long‐term tracking of individuals is difficult and the resulting data contain many inherent biases. In recent years, advances in both tracking and analytical techniques have meant that, for some taxonomic groups, sufficient numbers of survival estimates are available to allow variation within and among species to be explored. Here we review published estimates of annual adult survival rates in shorebird species across the globe, and construct models to explore the phylogenetic, geographical, seasonal and sex‐based variation in survival rates. Models of 295 survival estimates from 56 species show that survival rates calculated from recoveries of dead individuals or from return rates of marked individuals are significantly lower than estimates from mark–recapture models. Survival rates also vary across flyways, largely as a consequence of differences in the genera that have been studied and the analytical methods used, with published estimates from the Americas and from smaller shorebirds (Actitis, Calidris and Charadrius spp.) tending to be underestimated. By incorporating the analytical method used to generate each estimate within a mixed model framework, we provide method‐corrected species‐specific and genus‐specific adult annual survival estimates for 52 species of 15 genera.     

A review of survival estimates for raptors and owls

This paper reviews the literature on survival estimates for different species of raptors and owls, examines the methods used to obtain the estimates, and draws out some general patterns arising. Estimating survival usually involves the marking of birds so that they can be recognized as individuals on subsequent encounters. Annual survival can then be estimated from: (1) birds ringed at known age (usually as nestlings) and subsequently reported by members of the public (usually as found dead), the ratio of recoveries at different ages being used to calculate annual survival; (2) marked breeding adults, trapped or re‐sighted in subsequent years in particular study areas, with the proportion re‐trapped (or re‐sighted) in each year being taken as the minimum annual survival; (3) live encounter (trapped or re‐sighted) of birds marked either as nestlings or breeding adults analysed using the capture–mark–recapture (or re‐sighting) methods to estimate annual survival; (4) a combination of reports of known‐age dead birds and re‐trapping/re‐sighting of live birds; (5) use of radio‐ or satellite‐tracking to follow the fates of individuals; and (6) the integration of these methods with other information, such as change in numbers between years, to derive estimates of survival and other demographic parameters. Studies confined to particular areas usually give estimates of ‘apparent annual survival’, because they take no account of birds that leave the area. However, radio‐ or satellite‐tracking makes it possible to estimate true survival, including survival of prebreeders that have low natal‐site fidelity (this usually requires satellite telemetry). As in other birds, the preferred method for estimating survival has changed over time, as new and more robust methods of estimation have been developed. Methods 1 and 2 were the first to be developed, but without statistical underpinning, while methods 3–6 were developed later on the basis of formal statistical models. This difference has to be borne in mind in comparing older with newer estimates for particular species. Published survival estimates were found for three species of Cathartidae, one of Pandionidae, 29 of Accipitridae, 12 of Falconidae, one of Tytonidae and nine of Strigidae, almost all from temperate Northern Hemisphere species. In most of these species more than one estimate was available, and in some separate estimates for different age or sex groups. The main patterns to emerge included: (1) a significant tendency for annual adult survival to increase with body weight, smaller species having annual survival rates mainly of 60–70%, medium‐sized species having rates mainly in the range 70–90% and the largest having rates of > 90%, in the absence of obvious human‐caused losses; (2) a lower survival in the first or prebreeding years of life than in subsequent years; (3) a lack of obvious or consistent differences in survival between the sexes, where these could be distinguished; and (4) in the few species for which enough data were available, a decline in annual survival rates in the later years of life.     

Apparent survival and return rate of breeders in the southern temperate White‐rumped Swallow Tachycineta leucorrhoa

Life‐histories and demographic parameters of southern temperate bird species have been little studied. We estimated return rates between years and sexes, and adult apparent survival and recapture probabilities with mark–recapture data on White‐rumped Swallows and found a lower return rate of unsuccessful females. There was little support for influences of sex or year on survival rates. The estimates were equivalent to the lowest value reported for a northern congener, in contrast to the prediction of geographical variation under life‐history theory.     

Climate history,human impacts and global body size of Carnivora (Mammalia: Eutheria) at multiple evolutionary scales

Aim One of the longest recognized patterns in macroecology, Bergmann’s rule, describes the tendency for homeothermic animals to have larger body sizes in cooler climates than their phylogenetic relatives in warmer climates. Here we provide an integrative process‐based explanation for Bergmann’s rule at the global scale for the mammal order Carnivora. Location Global. Methods Our database comprises the body sizes of 209 species of extant terrestrial Carnivora, which were analysed using phylogenetic autocorrelation and phylogenetic eigenvector regression. The interspecific variation in body size was partitioned into phylogenetic (P) and specific (S) components, and mean P‐ and S‐components across species were correlated with environmental variables and human occupation both globally and for regions glaciated or not during the last Ice Age. Results Three‐quarters of the variation in body size can be explained by phylogenetic relationships among species, and the geographical pattern of mean values of the P‐component is the opposite of the pattern predicted by Bergmann’s rule. Partial regression revealed that at least 43% of global variation in the mean phylogenetic component is explained by current environmental factors. In contrast, the mean S‐component of body size shows large positive deviations from ancestors across the Holarctic, and negative deviations in southern South America, the Sahara Desert, and tropical Asia. There is a moderately strong relationship between the human footprint and body size in glaciated regions, explaining 19% of the variance of the mean P‐component. The relationship with the human footprint and the P‐component is much weaker in the rest of the world, and there is no relationship between human footprint and S‐component in any region. Main conclusions Bergmannian clines are stronger at higher latitudes in the Northern Hemisphere because of the continuous alternation of glacial–interglacial cycles throughout the late Pliocene and Pleistocene, which generated increased species turnover, differential colonization and more intense adaptive processes soon after glaciated areas became exposed. Our analyses provide a unified explanation for an adaptive Bergmann’s rule within species and for an interspecific trend towards larger body sizes in assemblages resulting from historical changes in climate and contemporary human impacts.     

On the evolution of clutch size and nest size in passerine birds

Summary I examined the hypothesis that the clutch size of some altricial birds may be limited by over-crowding of the nestlings in the nest, by comparing data on different species of European passerines. Large-sized birds build, relative to the body, larger nests than small-sized birds, both as regards the inner and the outer nest widths and as regards edge breadth; only inner nestcup depth did not change relatively to body size. Nest size also varied in relation to nesting place. Birds with open nests built off the ground had a rather narrow nestcup, whereas those with a domed nest, or which nest in a cavity, had a wide nestcup. When only open-nesters were compared, birds nesting on, or close to, the ground tended to have a wider nestcup than birds nesting above the ground. Inner nestcup width was correlated with the amount of mosses and lichens used in building the nest; the more of such materials the narrower the nestcup. The three variables: standardised body size, nesting place, and type of nesting material used accounted for 92% of the overall variation observed in inner nestcup width. When controlling for adult body size, clutch size was positively correlated with the size of the nestcup. A multiple regression analysis showed that relative nestcup depth, nest site, and type of nesting materials used, accounted for 64% of the overall variation in clutch size.     

South temperate birds have higher apparent adult survival than tropical birds in Africa

Life history theory predicts an inverse relationship between annual adult survival and fecundity. Globally, clutch size shows a latitudinal gradient among birds, with south temperate species laying smaller clutches than north temperate species, but larger clutches than tropical species. Tropical birds often have higher adult survival than north temperate birds associated with their smaller clutches. However, the prediction that tropical birds should also have higher adult survival than south temperate birds because of smaller clutch sizes remains largely untested. We measured clutch size and apparent annual breeding adult survival for 17 south temperate African species to test two main predictions. First, we found strong support for a predicted inverse relationship between adult survival and clutch size among the south temperate species, consistent with life‐history theory. Second, we compared our clutch size and survival estimates with published estimates for congeneric tropical African species to test the prediction of larger clutch size and lower adult survival among south temperate than related tropical species. We found that south‐temperate species laid larger clutches, as predicted, but had higher, rather than lower, apparent adult survival than related tropical species. The latter result may be an artefact of different approaches to measuring survival, but the results suggest that adult survival is generally high in the south temperate region and raises questions about the importance of the cost of reproduction to adult survival.     

Do southern African songbirds live longer than their European counterparts?

Understanding life history evolution in tropical and southern hemisphere birds has been hindered by a paucity of knowledge of key life history traits and this is particularly true of African songbirds. Here we use a unique long-term mark-recapture data set collected over 16 years in Malawi (latitude 16°S) to estimate adult survival rates for 28 African passerine species. Survival of these and 11 other African songbirds (taken from the literature) showed a bi-modal distribution with annual survival of insectivores and nectarivores (bulbuls, thrushes, warblers and sunbirds) averaging 72% (quartiles 63–80%) compared to 54% (50–62%) in granivores (weavers, finches and canaries). The mean adult life expectancy of African insectivores and nectarivores (3.1 yr) was more than twice that of related European insectivores (1.4 yr) and nearly twice that of African granivores (1.6 yr). These marked differences in survival were highly significant after controlling for body mass and phylogeny. Among African songbirds there was a strong negative correlation between adult survival and clutch size with granivores laying relatively large clutches and living relatively short lives. We hypothesize that these differing life history trade-offs reflect variation in the seasonality of food resources whereby survival rates of northern temperate songbirds may be limited by food availability and cold weather during winter, while survival of southern African granivores may be limited by the influence of a variable and unpredictable rainfall regime on seed availability.     

Geographic variation in reproduction and survival of kelp gulls Larus dominicanus vetula in southern Africa

Different populations of a species tend to vary in survival and reproduction, but the extent and scale of such spatial variation are poorly known. We estimated survival and clutch size of kelp gulls Larus dominicanus vetula across their entire African range. At this large geographic scale, we found no evidence for spatial variation in survival, and there was no variation in clutch size. However, there was considerable variation in clutch size among colonies within regions. Over the whole study, mean annual survival of juvenile and adult birds was 0.44 and 0.84, and mean clutch size was 2.2 eggs. A matrix population model showed that population growth was least sensitive to variation in clutch size, and the observed variation in clutch size could not fully account for the observed variation in population growth among colonies and regions. Our results thus suggest that dispersal and/or variation in survival (including egg/nestling survival) at a small spatial scale are also important for the spatial pattern of kelp gull population dynamics. These results are consistent with a metapopulation approach to spatial population dynamics.     

Density‐dependent survival and recruitment in a long‐distance Palaearctic migrant,the Sand Martin Riparia riparia

Long‐term studies can provide powerful insights into the relative importance of different demographic and environmental factors determining avian population dynamics. Here we use 23 years of capture–mark–recapture data (1981–2003) to estimate recruitment and survival rates for a Sand Martin Riparia riparia population in Cheshire, NW England. Inter‐annual variation in recruitment and adult survival was positively related to rainfall in the sub‐Saharan wintering grounds, but unrelated to weather conditions on the breeding grounds. After allowing for the effects of African rainfall, both demographic rates were negatively density‐dependent: adult survival was related to the size of the western European Sand Martin population (probably reflecting competition for resources in the shared wintering grounds) while recruitment was related to the size of the local study population in Cheshire (potentially reflecting competition for nesting sites or food). Local population size was more sensitive to variation in adult survival than to variation in recruitment, and an increase in population size after 1995 was driven mainly by the impact of more favourable conditions in the African wintering grounds on survival rates of adults. Overwinter survival in this long‐distance Palaearctic migrant is determined partly by the amount of suitable wetland foraging habitat in the sub‐Saharan wintering grounds (which is limited by the extent of summer rainfall) and partly by the number of birds exploiting that habitat.     

‘Heaven’ for serpents? A mark–recapture study of tiger snakes (Notechis scutatus) on Carnac Island,Western Australia

Abstract Animals resident on small islands provide excellent opportunities to carry out detailed mark–recapture studies. Populations are closed and ecosystems are often simpler than those of mainland sites. These factors enable the study of cryptic species that have otherwise been neglected. Snakes are notable for their secretive nature and, as a result, few natural populations have been accurately described through long‐term mark–recapture monitoring. A population of tiger snakes (Notechis scutatus) was studied on Carnac Island, a small limestone island (16 ha) off the coast of Western Australia. Population estimates show that snake density is very high, with more than 20 adult snakes per ha. This equates to a biomass of more than 100 kg of a top predator concentrated in a very small area. Such a high predator density can be explained because adult snakes feed mainly on chicks of nesting birds that breed in large colonies on Carnac but forage elsewhere. Substantial annual growth rates in body size in most individuals suggest that food availability is high on Carnac. Growth rates decreased more sharply in adult females than in males, whereas annual changes in body mass were similar in both sexes, probably because of the high energetic costs of reproduction experienced by females. Surprisingly, the sex ratio was highly biased, with males largely outnumbering females.     

Estimability of migration survival rates from integrated breeding and winter capture–recapture data

Long‐distance migration is a common phenomenon across the animal kingdom but the scale of annual migratory movements has made it difficult for researchers to estimate survival rates during these periods of the annual cycle. Estimating migration survival is particularly challenging for small‐bodied species that cannot carry satellite tags, a group that includes the vast majority of migratory species. When capture–recapture data are available for linked breeding and non‐breeding populations, estimation of overall migration survival is possible but current methods do not allow separate estimation of spring and autumn survival rates. Recent development of a Bayesian integrated survival model has provided a method to separately estimate the latent spring and autumn survival rates using capture–recapture data, though the accuracy and precision of these estimates has not been formally tested. Here, I used simulated data to explore the estimability of migration survival rates using this model. Under a variety of biologically realistic scenarios, I demonstrate that spring and autumn migration survival can be estimated from the integrated survival model, though estimates are biased toward the overall migration survival probability. The direction and magnitude of this bias are influenced by the relative difference in spring and autumn survival rates as well as the degree of annual variation in these rates. The inclusion of covariates can improve the model's performance, especially when annual variation in migration survival rates is low. Migration survival rates can be estimated from relatively short time series (4–5 years), but bias and precision of estimates are improved when longer time series (10–12 years) are available. The ability to estimate seasonal survival rates of small, migratory organisms opens the door to advancing our understanding of the ecology and conservation of these species. Application of this method will enable researchers to better understand when mortality occurs across the annual cycle and how the migratory periods contribute to population dynamics. Integrating summer and winter capture data requires knowledge of the migratory connectivity of sampled populations and therefore efforts to simultaneously collect both survival and tracking data should be a high priority, especially for species of conservation concern.     

Covariation between clutch size, egg weight and egg shape: comparative evidence for chelonians

The variation in reproductive variables is documented both within and across species of chelonians. At both the generic and family levels, egg weight and clutch size show positive, significant correlations with adult carapace length. There is a negative correlation across both genera and families between clutch size and egg weight after removing the effects of body size, suggesting an evolutionary trade-off between these two life-history characteristics. However, the trade-off is not complete, since clutch size is positively correlated with clutch weight after removing the effects of body size. Terrestrial species lay fewer and larger eggs for their size than freshwater or marine species, but this association is statistically confounded by the fact that chelonian families form ecological groups. There is no significant association between habitat and clutch weight after removing the effects of body size, nor between latitude and either egg weight or clutch size, but temperate species have a heavier clutch weight after removing the effects of body size. Larger species lay eggs that are more spherical, but after controlling for body size, egg shape is not associated with clutch size. The patterns of covariation between adult weight, egg weight and clutch size contrast with those reported for birds and mammals, and some reasons for these differences are discussed.     

Mass gained during breeding positively correlates with adult survival because both reflect life history adaptation to seasonal food availability

Both mass (as a measure of body reserves) during breeding and adult survival should reflect variation in food availability. Those species that are adapted to less seasonally variable foraging niches and so where competition dominates during breeding, will tend to have a higher mass increase via an interrupted foraging response, because their foraging demands increase and so become more unpredictable. They will then produce few offspring per breeding attempt, but trade this off with higher adult survival. In contrast, those species that occupy a more seasonal niche will not gain mass because foraging remains predictable, as resources become superabundant during breeding. They can also produce more offspring per breeding attempt, but with a trade-off with reduced adult survival. We tested whether the then predicted positive correlation between levels of mass gained during seasonal breeding and adult survival was present across 40 species of tropical bird measured over a 10-year period in a West African savannah. We showed that species with a greater seasonal mass increase had higher adult survival, controlling for annual mass variation (i.e. annual variation in absolute food availability) and variation in the timing of peak mass (i.e. annual predictability of food availability), clutch size, body size, migratory status and phylogeny. Our results support the hypothesis that the degree of seasonal mass variation in birds is probably an indication of life history adaptation: across tropical bird species it may therefore be possible to use mass gain during breeding as an index of adult survival.     

Life-history and ecological correlates of geographic variation in egg and clutch mass among passerine species

Broad geographic patterns in egg and clutch mass are poorly described, and potential causes of variation remain largely unexamined. We describe interspecific variation in avian egg and clutch mass within and among diverse geographic regions and explore hypotheses related to allometry, clutch size, nest predation, adult mortality, and parental care as correlates and possible explanations of variation. We studied 74 species of Passeriformes at four latitudes on three continents: the north temperate United States, tropical Venezuela, subtropical Argentina, and south temperate South Africa. Egg and clutch mass increased with adult body mass in all locations, but differed among locations for the same body mass, demonstrating that egg and clutch mass have evolved to some extent independent of body mass among regions. A major portion of egg mass variation was explained by an inverse relationship with clutch size within and among regions, as predicted by life-history theory. However, clutch size did not explain all geographic differences in egg mass; eggs were smallest in South Africa despite small clutch sizes. These small eggs might be explained by high nest predation rates in South Africa; life-history theory predicts reduced reproductive effort under high risk of offspring mortality. This prediction was supported for clutch mass, which was inversely related to nest predation but not for egg mass. Nevertheless, clutch mass variation was not fully explained by nest predation, possibly reflecting interacting effects of adult mortality. Tests of the possible effects of nest predation on egg mass were compromised by limited power and by counterposing direct and indirect effects. Finally, components of parental investment, defined as effort per offspring, might be expected to positively coevolve. Indeed, egg mass, but not clutch mass, was greater in species that shared incubation by males and females compared with species in which only females incubate eggs. However, egg and clutch mass were not related to effort of parental care as measured by incubation attentiveness. Ecological and life-history correlates of egg and clutch mass variation found here follow from theory, but possible evolutionary causes deserve further study.     

Bias in ring‐recovery studies: causes of mortality of little owls Athene noctua and implications for population assessment

Recoveries of marked animals hold long‐term, large‐scale information on survival and causes of mortality, but are prone to bias towards dead recoveries and casualties in the range of presence of potential finders. Thus, accounting for circumstance‐related recovery probabilities is crucial in statistical approaches. For the little owl, a species of conservation concern in central Europe, raw ring recoveries suggested a strong human‐related impact on survival. We analysed the proportions of the main causes of death using a large sample of radio‐tracked birds as a reference. We compared ring recoveries in southern Germany collected 1950–2012 (n = 465 dead recoveries of 2007 recoveries of 30 623 ringed birds) with data from a radio‐tracking study in the same region 2009–2012 (n = 177 dead recoveries of 377 tagged individuals). Two assumptions of multi‐state ring recovery modelling were unrealistic. First, not all dispatched rings remained available to potential finders. Instead, 34% of tracked birds were displaced to sites where rings were irretrievable, resulting in biased estimates of recovery probability. Second, the proportions of irretrievable rings were disproportional, with 48% in predated birds and 5% in human‐induced mortality. Consequently, the sample of rings from which recoveries were drawn differed from the sample of dispatched rings. Accounting for these biases in a multi‐state model, we estimated the frequencies of main causes of mortality to 45% for predation, 20% for casualties due to traffic and at buildings and 34% for all other causes. In radio‐tracked birds, predation was even more dominant (76%). Integrating mark–recapture data and telemetry observations allowed detecting and quantifying so far unknown recovery bias and resulted in improved estimates of key population parameters. The demography of little owls likely depends mainly on predator–prey relationships rather than on human‐induced deaths.     

Integrating high‐speed videos in capture‐mark‐recapture studies of insects

Capture–mark–recapture (CMR) studies have been used extensively in ecology and evolution. While it is feasible to apply CMR in some animals, it is considerably more challenging in small fast‐moving species such as insects. In these groups, low recapture rates can bias estimates of demographic parameters, thereby handicapping effective analysis and management of wild populations. Here, we use high‐speed videos (HSV) to capture two large dragonfly species, Anax junius and Rhionaeschna multicolor, that rarely land and, thus, are particularly challenging for CMR studies. We test whether HSV, compared to conventional “eye” observations, increases the “resighting” rates and, consequently, improves estimates of both survival rates and the effects of demographic covariates on survival. We show that the use of HSV increases the number of resights by 64% in A. junius and 48% in R. multicolor. HSV improved our estimates of resighting and survival probability which were either under‐ or overestimated with the conventional observations. Including HSV improved credible intervals for resighting rate and survival probability by 190% and 130% in A. junius and R. multicolor, respectively. Hence, it has the potential to open the door to a wide range of research possibilities on species that are traditionally difficult to monitor with distance sampling, including within insects and birds.     

Climate change leads to differential shifts in the timing of annual cycle stages in a migratory bird

Shifts in reproductive phenology due to climate change have been well documented in many species but how, within the same species, other annual cycle stages (e.g. moult, migration) shift relative to the timing of breeding has rarely been studied. When stages shift at different rates, the interval between stages may change resulting in overlaps, and as each stage is energetically demanding, these overlaps may have negative fitness consequences. We used long‐term data of a population of European pied flycatchers (Ficedula hypoleuca) to investigate phenological shifts in three annual cycle stages: spring migration (arrival dates), breeding (egg‐laying and hatching dates) and the onset of postbreeding moult. We found different advancements in the timing of breeding compared with moult (moult advances faster) and no advancement in arrival dates. To understand these differential shifts, we explored which temperatures best explain the year‐to‐year variation in the timing of these stages, and show that they respond differently to temperature increases in the Netherlands, causing the intervals between arrival and breeding and between breeding and moult to decrease. Next, we tested the fitness consequences of these shortened intervals. We found no effect on clutch size, but the probability of a fledged chick to recruit increased with a shorter arrival‐breeding interval (earlier breeding). Finally, mark–recapture analyses did not detect an effect of shortened intervals on adult survival. Our results suggest that the advancement of breeding allows more time for fledgling development, increasing their probability to recruit. This may incur costs to other parts of the annual cycle, but, despite the shorter intervals, there was no effect on adult survival. Our results show that to fully understand the consequences of climate change, it is necessary to look carefully at different annual cycle stages, especially for organisms with complex cycles, such as migratory birds.     

Allometric scaling and Bayesian priors for annual survival of birds and mammals

Allometric theory predicts that instantaneous mortality rates scale with body mass with a negative quarter power. Such a relationship would mean that the survival rate of one species is partly predictable from the survival rate of other species. We develop allometric regression models for annual adult survival of birds and mammals, using data collected from the literature. These models conform to the predictions of the allometric theory; the value of negative one-quarter for the scaling parameter is within the 95% credible interval, which is [-0.31, -0.10] for birds and [-0.35, -0.15] for mammals. The predictions are very well supported when evaluated using an independent set of data. The regression models can be used to provide objective and informative Bayesian priors for annual adult survival rates of birds and mammals or to act as a point of comparison in new studies.     

Annual survival of adult Atlantic Puffins Fratercula arctica is positively correlated with Herring Clupea harengus availability

Atlantic Herring is a keystone species in several marine ecosystems, supporting intensive fisheries as well as many predators including seabirds. Biomass of this stock in eastern North America has declined considerably in recent years, potentially putting at risk populations of its predators. Although adult survival in seabirds is considered robust to moderate changes in food availability, it is also the life‐history component most critical to sustaining populations of long‐lived birds. To investigate the possibility that Atlantic Puffin survival has been affected by reduced abundance of its main prey, we analysed the encounter histories of 2999 Atlantic Puffins ringed on Machias Seal Island to estimate annual adult survival for the years 1999–2011 and assess trends in survival and the effects of several biological and environmental covariates. Features of Puffin biology and resighting procedures likely to introduce heterogeneity into our resighting probabilities were accounted for and models of survival were assessed using standard methods. We used the variance components procedure in Program MARK and survival estimates from a time‐varying model to estimate the process variance (biological variation in survival) accounted for by suspected covariates of survival. Two proxies of food availability each explained more than half of the variation in annual survival: fishery landings of Atlantic Herring (52%) and per cent (by mass) of 1‐group Herring in the diet of Puffin chicks (51%). In addition to these proxies, May sea‐surface temperature accounted for 37% of variance in survival, but winter values of North Atlantic Oscillation showed no effect. Of those parameters of Puffin biology examined, chick growth rate explained 19% of the process variance in annual survival; laying date, fledging condition and fledging date all explained no variance. A decline in fishery landings of Herring since the early 1990s, and a concurrent decline in adult Puffin survival, reinforces concern for the health of the population of Herring, a keystone forage fish in this region, and of the community of marine predators in the Gulf of Maine that rely on Herring for their survival and reproduction.