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Divergent migration within lake sturgeon (Acipenser fulvescens) populations: Multiple distinct patterns exist across an unrestricted migration corridor

Population structure, distribution, abundance, and dispersal arguably underpin the entire field of animal ecology, with consequences for regional species persistence, and provision of ecosystem services. Divergent migration behaviours among individuals or among populations is an important aspect of the ecology of highly-mobile animals, allowing populations to exploit spatially- or temporally-distributed food and space resources. This study investigated the spatial ecology of lake sturgeon ( Acipenser fulvescens ) within the barrier free Huron-Erie Corridor (HEC), which connects Lake Huron and Lake Erie of the North American Laurentian Great Lakes. Over six years (2011 – 2016), movements of 268 lake sturgeon in the HEC were continuously monitored across the Great Lakes using acoustic telemetry (10 yr battery life acoustic transmitters). Five distinct migration behaviours were identified with hierarchical cluster analysis, based on the phenology and duration of river and lake use. Lake sturgeon in the HEC were found to contain a high level of intraspecific divergent migration, including partial migration with the existence of residents. Specific behaviours included year-round river residency and multiple lake-migrant behaviours that involved movements between lakes and rivers. Over 85% of individuals were assign to migration behaviours as movements were consistently repeated over the study, which suggested migration behaviours were consistent and persistent in lake sturgeon. Differential use of specific rivers or lakes by acoustic-tagged lake sturgeon further subdivided individuals into 14 “contingents” (spatiotemporally segregated subgroups). Contingents associated with one river (Detroit or St. Clair) were rarely detected in the other river, which confirmed that lake sturgeon in the Detroit and St. Clair represent two semi-independent populations that could require separate management consideration for their conservation. The distribution of migration behaviours did not vary between populations, sexes, body size, or among release locations, which indicated that intrapopulation variability in migratory behaviour is a general feature of the spatial ecology of lake sturgeon in un-fragmented landscapes.

Michigan, Ontario

Elk migration influences the risk of disease spillover in the Greater Yellowstone Ecosystem

Wildlife migrations provide important ecosystem services, but they are declining. Within the Greater Yellowstone Ecosystem (GYE) some elk ( Cervus canadensis ) herds are losing migratory tendencies, which may increase spatiotemporal overlap between elk and livestock (domestic bison [ Bison bison ] and cattle [ Bos taurus ]), potentially exacerbating pathogen transmission risk. We combined disease, movement, demographic, and environmental data from eight elk herds in the GYE to examine the differential risk of brucellosis transmission (through aborted fetuses) from migrant and resident elk to livestock. For both migrants and residents, we found that transmission risk from elk to livestock occurred almost exclusively on private ranchlands as opposed to state or federal grazing allotments. Weather variability affected the estimated distribution of spillover risk from migrant elk to livestock, with a 7‐12% increase in migrant abortions on private ranchlands during years with heavier snowfall. In contrast, weather variability did not affect spillover risk from resident elk. Migrant elk were responsible for the majority (68%) of disease spillover risk to livestock because they occurred in greater numbers than resident elk. On a per‐capita basis, however, our analyses suggested that resident elk disproportionately contributed to spillover risk. In five of seven herds, we estimated that the per‐capita spillover risk was greater from residents than from migrants. Averaged across herds, an individual resident elk was 23% more likely than an individual migrant elk to abort on private ranchlands. Our results demonstrate links between migration behavior, spillover risk, and environmental variability, and highlight the utility of integrating models of pathogen transmission and host movement to generate new insights about the role of migration in disease spillover risk. Further, they add to the accumulating body of evidence across taxa that suggests that migrants and residents should be considered separately during investigations of wildlife disease ecology. Finally, our findings have applied implications for elk and brucellosis in the GYE, and suggest that managers should prioritize actions that maintain spatial separation of elk and livestock on private ranchlands during years when snowpack persists into the risk period.

Wyoming

Group density, disease, and season shape territory size and overlap of social carnivores

1. The spatial organization of a population can influence the spread of information, behaviour, and pathogens. Territory size and territory overlap, components of spatial organization, provide key information as these metrics may be indicators of habitat quality, resource dispersion, contact rates, and environmental risk (e.g., indirectly transmitted pathogens). Furthermore, sociality and behaviour can also shape space use, and subsequently, how space use and habitat quality together impact demography. 2. Our study aims to identify factors shaping the spatial organization of wildlife populations and assess the impact of epizootics on space use. We also use network analysis to describe spatial organization and connectivity of social groups. 3. Here, we assessed the seasonal spatial organization of Serengeti lions and Yellowstone wolves at the group level. We examine the factors predicting mean territory size and mean territory overlap for each population using generalized additive models. We further explore the mechanisms by which disease perturbations could cause changes in spatial organization. 4. We demonstrate that lions and wolves were similar in that group-level factors, such as number of groups, shaped spatial organization more than population-level factors, such as population density. Factors shaping territory size were slightly different than factors shaping territory overlap; for example, wolf pack size was an important predictor of territory overlap, but not territory size. Lion spatial networks were more highly connected, while wolf spatial networks varied seasonally. We found that resource dispersion may be more important for driving territory size and overlap for wolves than for lions. Additionally, canine distemper epizootics may alter lion spatial organization, highlighting the importance of including behavioural and movement ecology in studies of pathogen transmission dynamics. 5. We provide insight about when we might expect to observe the impacts of resource dispersion, disease perturbations, and other ecological factors on spatial organization. Our work highlights the importance of monitoring and managing social carnivore populations at the group level. Future research should elucidate the complex relationships between demographics, social and spatial structure, abiotic and biotic conditions, and pathogen infections.

Journal of Animal Ecology

Optimal population prediction of sandhill crane recruitment based on climate-mediated habitat limitations

Prediction is fundamental to scientific enquiry and application; however, ecologists tend to favour explanatory modelling. We discuss a predictive modelling framework to evaluate ecological hypotheses and to explore novel/unobserved environmental scenarios to assist conservation and management decision-makers. We apply this framework to develop an optimal predictive model for juvenile (<1 year old) sandhill crane Grus canadensis recruitment of the Rocky Mountain Population (RMP). We consider spatial climate predictors motivated by hypotheses of how drought across multiple time-scales and spring/summer weather affects recruitment. Our predictive modelling framework focuses on developing a single model that includes all relevant predictor variables, regardless of collinearity. This model is then optimized for prediction by controlling model complexity using a data-driven approach that marginalizes or removes irrelevant predictors from the model. Specifically, we highlight two approaches of statistical regularization, Bayesian least absolute shrinkage and selection operator (LASSO) and ridge regression. Our optimal predictive Bayesian LASSO and ridge regression models were similar and on average 37% superior in predictive accuracy to an explanatory modelling approach. Our predictive models confirmed a priori hypotheses that drought and cold summers negatively affect juvenile recruitment in the RMP. The effects of long-term drought can be alleviated by short-term wet spring–summer months; however, the alleviation of long-term drought has a much greater positive effect on juvenile recruitment. The number of freezing days and snowpack during the summer months can also negatively affect recruitment, while spring snowpack has a positive effect. Breeding habitat, mediated through climate, is a limiting factor on population growth of sandhill cranes in the RMP, which could become more limiting with a changing climate (i.e. increased drought). These effects are likely not unique to cranes. The alteration of hydrological patterns and water levels by drought may impact many migratory, wetland nesting birds in the Rocky Mountains and beyond. Generalizable predictive models (trained by out-of-sample fit and based on ecological hypotheses) are needed by conservation and management decision-makers. Statistical regularization improves predictions and provides a general framework for fitting models with a large number of predictors, even those with collinearity, to simultaneously identify an optimal predictive model while conducting rigorous Bayesian model selection. Our framework is important for understanding population dynamics under a changing climate and has direct applications for making harvest and habitat management decisions.

Journal of Animal Ecology

Understanding natal emigration in group-living carnivores

Dispersal is a critical ecological and evolutionary process, yet the drivers of individual variation in emigration, particularly under human-modified conditions, remain relatively understudied. We investigate the determinants and patterns of emigration in a managed population of cooperative breeders using genetic samples from harvested individuals. We used a novel genetic approach to identify natal emigration through sibship analyses of DNA from harvested grey wolves ( Canis lupus ), a cooperative-breeding species, and evaluated how prey biomass, wolf density, human-caused mortality and sex influenced natal emigration across Idaho, USA, from 2017 to 2023. From our analysis of 182 sibling groups (representing packs), we evaluated natal emigration status for 224 individuals. Of these, 86 were classified as emigrants and 138 as non-dispersers. We assessed how covariates influenced the probability of emigration and evaluated sex differences in apparent dispersal distance. We found no sex differences in emigration propensity or apparent dispersal distance. Emigrants dispersed an average of 81.8 km (range: 30.0–338.9 km) from their inferred natal territories. Emigration was more likely from territories with higher prey biomass and lower wolf density, while human-caused mortality had no detectable effect. We found that ecological drivers, such as prey availability and population density, have a stronger effect on the propensity to emigrate than harvest intensity. These results highlight the importance of resource availability and population dynamics in shaping dispersal patterns in a cooperative breeder. Our study used a novel genetic method to assess natal emigration in a group-living carnivore and developed a framework broadly applicable to social systems where kin structures can be traced to dispersers. Our findings identify landscape-scale conditions that influence dispersal and can inform the management of wild populations.

Idaho

The early bird gets the shrimp: Confronting assumptions of isotopic equilibrium and homogeneity in a wild bird population

1. Because stable isotope distributions in organic material vary systematically across energy gradients that exist in ecosystems, community and population structures, and in individual physiological systems, isotope values in animal tissues have helped address a broad range of questions in animal ecology. It follows that every tissue sample provides an isotopic profile that can be used to study dietary or movement histories of individual animals. Interpretations of these profiles depend on the assumption that metabolic pools are isotopically well mixed and in equilibrium with dietary resources prior to tissue synthesis, and they extend to the population level by assuming isotope profiles are identically distributed for animals using the same proximal dietary resource. As these assumptions are never fully met, studying structure in the variance of tissue isotope values from wild populations is informative. 2. We studied variation in &delta; 13 C, &delta; 15 N, &delta; 2 H and &delta; 18 O data for feathers from a population of eared grebes (Podiceps nigricollis) that migrate to Great Salt Lake each fall to moult feathers. During this time, they cannot fly and feed almost exclusively on superabundant brine shrimp (Artemia franciscana). The ecological simplicity of this situation minimized the usual spatial and trophic complexities often present in natural studies of feather isotope values. 3. Ranges and variances of isotope values for the feathers were larger than those from previously published studies that report feather isotopic variance, but they were bimodally distributed in all isotope dimensions. Isotope values for proximal dietary resources and local surface water show that some of the feathers we assumed to have been grown locally must have been grown before birds reached isotopic equilibrium with local diet or immediately prior to arrival at Great Salt Lake. 4. Our study provides novel insights about resource use strategies in eared grebes during migration. More generally, it demonstrates the utility of studying variance structures and questioning assumptions implicit in the interpretation of stable isotope data from wild animals.

Utah

Droughts reshape apex predator space use and intraguild overlap

1. Droughts are increasing in frequency and severity globally due to climate change, leading to changes in resource availability that may have cascading effects on animal ecology. Resource availability is a key driver of animal space use, which in turn influences interspecific interactions like intraguild competition. Understanding how climate-induced changes in resource availability influence animal space use, and how species-specific responses scale up to affect intraguild dynamics, is necessary for predicting broader community-level responses to climatic changes. 2. Although several studies have demonstrated the ecological impacts of drought, the behavioral responses of individuals that scale up to these broader-scale effects are not well known, particularly among animals in top trophic levels, such as large carnivores. Furthermore, we currently lack understanding of how the impacts of climate variability on individual carnivore behavior are linked to intraguild dynamics, in part because multi-species datasets collected at timescales relevant to climatic changes are rare. 3. Using 11 years of GPS data from four sympatric large carnivore species in southern Africa – lions ( Panthera leo ), leopards ( Panthera pardus ), African wild dogs ( Lycaon pictus) , and cheetahs ( Acinonyx jubatus ) – spanning 4 severe drought events, we test whether drought conditions impact 1) large carnivore space use, 2) broad-scale intraguild spatial overlap, and 3) fine-scale intraguild interactions. 4. Drought conditions expanded space use across species, with carnivores increasing their monthly home range sizes by 35% (wild dogs) to 66% (leopards). Drought conditions increased the amount of spatial overlap between lions and subordinate felids (cheetahs and leopards) by up to 119%, but only lion-cheetah encounter rates were affected by these changes, declining in response to drought. 5. Our findings reveal that drought has a clear signature on the space use of multiple sympatric large carnivore species, which can alter spatiotemporal partitioning between competing species. Our study thereby illuminates the links between environmental change, animal behavior, and intraguild dynamics. While fine-scale avoidance strategies may facilitate intraguild coexistence during periodic droughts, large carnivore conservation may require considerable expansion of protected areas or revised human-carnivore coexistence strategies to accommodate the likely long-term increased space demands of large carnivores under projected increases in drought intensity.

Okavango Delta

Informed breeding dispersal following stochastic changes to patch quality in a pond-breeding amphibian

The unidirectional movement of animals between breeding patches (i.e. breeding dispersal) has profound implications for the ecological and evolutionary dynamics of spatially structured populations. In spatiotemporally variable environments, individuals are expected to adjust their dispersal decisions according to information gathered on the environmental and/or social cues that reflect the fitness prospects in a given breeding patch (i.e. informed dispersal). A paucity of empirical work limited our understanding of the ability of animals to depart from low-quality breeding patches and settle in high-quality breeding patches. We examined the capacity of individuals to respond to stochastic changes in habitat quality via informed breeding dispersal in a pond-breeding amphibian. We conducted a 5-year (2015–2019) capture–recapture study of boreal toads Anaxyrus boreas boreas ( n = 1,100) that breed in beaver ponds in western Wyoming, USA. During early spring of 2017, an extreme flooding event destroyed several beaver dams and resulted in the loss of breeding habitat. We used multi-state models to investigate how temporal changes in pond characteristics influenced breeding dispersal, and determine whether movement decisions were in accordance with prospects for reproductive fitness. Boreal toads more often departed from low-quality breeding ponds (without successful metamorphosis) and settled in high-quality breeding ponds (with successful metamorphosis). Movement decisions were context-dependent and associated with pond characteristics altered by beaver dam destruction. Individuals were more likely to depart from shallow ponds with high vegetation cover and settle in deep ponds with low vegetation cover. The probability of metamorphosis was related to the same environmental cues, suggesting that boreal toads assess the fitness prospects of a breeding patch and adjust movement decisions accordingly (i.e. informed breeding dispersal). We demonstrated that stochastic variability in environmental conditions and habitat quality can underpin dispersal behaviour in amphibians. Our study highlighted the mechanistic linkages between habitat change, movement behaviour and prospects for reproductive performance, which is critical for understanding how wild animals respond to rapid environmental change.

Wyoming

Interactive range‐limit theory (iRLT): An extension for predicting range shifts

A central theme of range‐limit theory (RLT) posits that abiotic factors form high‐latitude/altitude limits, whereas biotic interactions create lower limits. This hypothesis, often credited to Charles Darwin, is a pattern widely assumed to occur in nature. However, abiotic factors can impose constraints on both limits and there is scant evidence to support the latter prediction. Deviations from these predictions may arise from correlations between abiotic factors and biotic interactions, as a lack of data to evaluate the hypothesis, or be an artifact of scale. Combining two tenets of ecology—niche theory and predator–prey theory—provides an opportunity to understand how biotic interactions influence range limits and how this varies by trophic level. We propose an expansion of RLT, interactive RLT (iRLT), to understand the causes of range limits and predict range shifts. Incorporating the main predictions of Darwin's hypothesis, iRLT hypothesizes that abiotic and biotic factors can interact to impact both limits of a species’ range. We summarize current thinking on range limits and perform an integrative review to evaluate support for iRLT and trophic differences along range margins, surveying the mammal community along the boreal‐temperate and forest‐tundra ecotones of North America. Our review suggests that range‐limit dynamics are more nuanced and interactive than classically predicted by RLT. Many (57 of 70) studies indicate that biotic factors can ameliorate harsh climatic conditions along high‐latitude/altitude limits. Conversely, abiotic factors can also mediate biotic interactions along low‐latitude/altitude limits (44 of 68 studies). Both scenarios facilitate range expansion, contraction or stability depending on the strength and the direction of the abiotic or biotic factors. As predicted, biotic interactions most often occurred along lower limits, yet there were trophic differences. Carnivores were only limited by competitive interactions ( n = 25), whereas herbivores were more influenced by predation and parasitism (77%; 55 of 71 studies). We highlight how these differences may create divergent range patterns along lower limits. We conclude by (a) summarizing iRLT; (b) contrasting how our model system and others fit this hypothesis and (c) suggesting future directions for evaluating iRLT.

Journal of Animal Ecology

Temporal and spatial equivalence in demographic responses of emperor penguins (Aptenodytes forsteri) to environmental change

1. Population ecology and biogeography applications often necessitate the transfer of models across spatial and/or temporal dimensions to make predictions outside the bounds of the data used for model fitting. However, ecological data are often spatiotemporally unbalanced such that the spatial or the temporal dimension tends to contain more data than the other. This unbalance frequently leads model transfers to become substitutions, which are predictions to a different dimension than the predictive model was built on. Despite the prevalence of substitutions in ecology, studies validating their performance and their underlying assumptions are scarce. 2. Here, we present a successful case study demonstrating both space-for-time and time-for-space substitutions using emperor penguins (Aptenodytes forsteri) as the focal species. Using abundance-based species distribution models (aSDM) of adult emperor penguins in attendance during spring across 50 colonies, we predict long-term annual fluctuations in fledgling abundance and breeding success at a single colony, Pointe Géologie. Subsequently, we construct statistical models from time series of extended counts on Pointe Géologie to predict average fledgling abundance across 50 colonies. 3. Our analysis reveals that distance to nearest open water (NOW) exhibits the strongest association with both temporal and spatial data. aSDM’s space-for-time substitution performance, as measured by Pearson correlation coefficient was 0.63 and 0.56 when predicting breeding success and fledgling abundance time series, respectively. Linear regression of fledgling abundance on NOW yields similar time-for-space substitution performance when predicting abundance distribution of emperor penguin colonies with a correlation coefficient of 0.58. 4. We posit that such space-time equivalence arises because: 1) emperor penguins colonies conform to their existing fundamental niche; 2) there is not yet any environmental novelty when comparing the spatial vs temporal variation of distance to nearest open water; and 3) models of more specific components of life histories, such as fledgling abundance, rather than occurrence or total population abundance, are more transferable. Identifying these conditions empirically can enhance the qualitative validation of substitutions in cases where direct validation data are lacking.

Journal of Animal Ecology

Population ecology of house mice in unstable habitats

(1) The relationships between habitat change and house mouse populations were studied by monthly live trapping in a corn-wheat-hay rotation on a small Maryland farm. (2) Population density reached 53.0/ha in a wheat/hay field in October and 25.4/ha in corn in September. Populations increased by immigration as wheat or corn grew and ripened and decreased by emigration as hay became tall and dense. (3) Survival rates were high in winter in the relatively stable habitat of the wheat/hay field; they were low throughout the summer in both fields, and were reduced by corn harvest, less so by wheat harvest. If they were related to population density or increase, or to breeding condition, the relationships were obscured by the overriding influence of habitat change. (4) In the spring, when the population in the hay field `crashed,' essentially the entire population moved from long-established ranges in the hay field to the field of ripening wheat, where new ranges were established. In the new field, fewer than 30% of the old associations between individuals persisted. (5) Individual mice maintained home ranges (88.1 +- 6.1 m in length) in the same general area during their residence in a field. Ranges shifted from month to month, perhaps in response to changes in populations and habitat; exploratory travels and other movements also modified home range behaviour. (6) Minimum life expectancy (residence time) was greater from November (4-5 months) than from June/July (1-2 months). Maximum individual age was 17 months. (7) The demographic pattern fell at the r extreme of the r-K continuum. Mice bred from May to October, matured and produced litters rapidly, produced several litters in a season, and had a high turnover rate. (8) It was concluded that migration was a primary mechanism of population regulation in the cropfield mosaic and that it was driven by habitat change, a system in contrast to those described for house mice in confined conditions.

Journal of Animal Ecology

Long-term fitness consequences of early conditions in the kittiwake

1. The long - term fitness consequences of conditions during development are receiving growing attention: they are at the interface between ecological and evolutionary processes. We addressed the influence of the length of the rearing period and 'rank' on fitness components in a long -lived seabird species with deferred breeding: the kittiwake (Rissa tridactyla). Rank, which depends on hatching order, was used as a surrogate for dominance status in the brood. Rank could be viewed as a random factor affecting individuals regardless of their possible 'intrinsic quality' at birth. The length of the rearing period was used as a surrogate for parental effort. It reflects the interaction between numerous factors such as environmental conditions , parental quality, reproductive decisions and effort, and also offspring decisions and intrinsic quality at birth. 2. There was evidence of an influence of the length of the rearing period on local survival before recruitment. Individuals with shorter rearing periods had lower local survival during the first winter (e.g. the relationship was positive). They may incur higher mortality. In rank 1 prebreeders, this relationship was negative in older age-classes. Longer rearing periods and better condition at independence may be associated with stronger migrating ability, and prebreeders that have not yet made settlement decisions may emigrate permanently to distant locations. Such a complex pattern may reflect age-related changes in the relative contribution of mortality and permanent emigration to local survival. 3. The length of the rearing period had long - term consequences on reproductive performance. The relationship was positive but the rate of increase decreased slightly at higher values of the covariate. 4. There was an unambiguously negative influence of rank on survival before recruitment and recruitment probability, but not on demographic parameters specific to the reproductive stage. Juniors recruited later than elder siblings. The disadvantage of juniors may be expressed mainly in terms of higher mortality and disappearance from the population before recruitment.

Journal of Animal Ecology

Effects of host species and environment on the skin microbiome of Plethodontid salamanders

The amphibian skin microbiome is recognized for its role in defence against pathogens, including the deadly fungal pathogen Batrachochytrium dendrobatidis (Bd). Yet, we have little understanding of evolutionary and ecological processes that structure these communities, especially for salamanders and closely related species. We investigated patterns in the distribution of bacterial communities on Plethodon salamander skin across host species and environments. Quantifying salamander skin microbiome structure contributes to our understanding of how host-associated bacteria are distributed across the landscape, among host species, and their putative relationship with disease. We characterized skin microbiome structure (alpha-diversity, beta-diversity and bacterial operational taxonomic unit [OTU] abundances) using 16S rRNA gene sequencing for co-occurring Plethodon salamander species (35 Plethodon cinereus , 17 Plethodon glutinosus , 10 Plethodon cylindraceus ) at three localities to differentiate the effects of host species from environmental factors on the microbiome. We sampled the microbiome of P. cinereus along an elevational gradient ( n = 50, 700–1,000 m a.s.l.) at one locality to determine whether elevation predicts microbiome structure. Finally, we quantified prevalence and abundance of putatively anti-Bd bacteria to determine if Bd-inhibitory bacteria are dominant microbiome members. Co-occurring salamanders had similar microbiome structure, but among sites salamanders had dissimilar microbiome structure for beta-diversity and abundance of 28 bacterial OTUs. We found that alpha-diversity increased with elevation, beta-diversity and the abundance of 17 bacterial OTUs changed with elevation (16 OTUs decreasing, 1 OTU increasing). We detected 11 putatively anti-Bd bacterial OTUs that were present on 90% of salamanders and made up an average relative abundance of 83% ( SD ± 8.5) per salamander. All salamanders tested negative for Bd. We conclude that environment is more influential in shaping skin microbiome structure than host differences in these congeneric species, and suggest that environmental characteristics that covary with elevation influence microbiome structure. High prevalence and abundance of anti-Bd bacteria may contribute to low Bd levels in these populations of Plethodon salamanders.

Journal of Animal Ecology

Predictive modelling of habitat use by marine predators with respect to the abundance and depth distribution of pelagic prey

Understanding the ecological processes that underpin species distribution patterns is a fundamental goal in spatial ecology. However, developing predictive models of habitat use is challenging for species that forage in marine environments, as both predators and prey are often highly mobile and difficult to monitor. Consequently, few studies have developed resource selection functions for marine predators based directly on the abundance and distribution of their prey. We analysed contemporaneous data on the diving locations of two seabird species, the shallow-diving Peruvian Booby ( Sula variegata ) and deeper diving Guanay Cormorant ( Phalacrocorax bougainvilliorum ), and the abundance and depth distribution of their main prey, Peruvian anchoveta ( Engraulis ringens ). Based on this unique data set, we developed resource selection functions to test the hypothesis that the probability of seabird diving behaviour at a given location is a function of the relative abundance of prey in the upper water column. For both species, we show that the probability of diving behaviour is mostly explained by the distribution of prey at shallow depths. While the probability of diving behaviour increases sharply with prey abundance at relatively low levels of abundance, support for including abundance in addition to the depth distribution of prey is weak, suggesting that prey abundance was not a major factor determining the location of diving behaviour during the study period. The study thus highlights the importance of the depth distribution of prey for two species of seabird with different diving capabilities. The results complement previous research that points towards the importance of oceanographic processes that enhance the accessibility of prey to seabirds. The implications are that locations where prey is predictably found at accessible depths may be more important for surface foragers, such as seabirds, than locations where prey is predictably abundant. Analysis of the relative importance of abundance and accessibility is essential for the design and evaluation of effective management responses to reduced prey availability for seabirds and other top predators in marine systems.

Gulf of Guinea

Weak effects of geolocators on small birds: a meta‐analysis controlled for phylogeny and publication bias

Currently, the deployment of tracking devices is one of the most frequently used approaches to study movement ecology of birds. Recent miniaturisation of light‐level geolocators enabled studying small bird species whose migratory patterns were widely unknown. However, geolocators may reduce vital rates in tagged birds and may bias obtained movement data. There is a need for a thorough assessment of the potential tag effects on small birds, as previous meta‐analyses did not evaluate unpublished data and impact of multiple life‐history traits, focused mainly on large species and the number of published studies tagging small birds has increased substantially. We quantitatively reviewed 549 records extracted from 74 published and 48 unpublished studies on over 7,800 tagged and 17,800 control individuals to examine the effects of geolocator tagging on small bird species (body mass <100 g). We calculated the effect of tagging on apparent survival, condition, phenology and breeding performance and identified the most important predictors of the magnitude of effect sizes. Even though the effects were not statistically significant in phylogenetically controlled models, we found a weak negative impact of geolocators on apparent survival. The negative effect on apparent survival was stronger with increasing relative load of the device and with geolocators attached using elastic harnesses. Moreover, tagging effects were stronger in smaller species. In conclusion, we found a weak effect on apparent survival of tagged birds and managed to pinpoint key aspects and drivers of tagging effects. We provide recommendations for establishing matched control group for proper effect size assessment in future studies and outline various aspects of tagging that need further investigation. Finally, our results encourage further use of geolocators on small bird species but the ethical aspects and scientific benefits should always be considered.

Journal of Animal Ecology

Effect of land cover, habitat fragmentation and ant colonies on the distribution and abundance of shrews in southern California

Because effects of habitat fragmentation and anthropogenic disturbance on native animals have been relatively little studied in arid areas and in insectivores, we investigated the roles of different land covers, habitat fragmentation and ant colonies on the distribution and abundance of shrews, Notiosorex crawfordi and Sorex ornatus , in southern California. Notiosorex crawfordi was the numerically dominant species (trap-success rate 0·52) occurring in 21 of the 22 study sites in 85% of the 286 pitfall arrays used in this study. Sorex ornatus was captured in 14 of the sites, in 52% of the arrays with a total trap-success rate of 0·2. Neither of the species was found in one of the sites. The population dynamics of the two shrew species were relatively synchronous during the 4–5-year study; the peak densities usually occurred during the spring. Precipitation had a significant positive effect, and maximum temperature a significant negative effect on the trap-success rate of S. ornatus . Occurrence and abundance of shrews varied significantly between sites and years but the size of the landscape or the study site had no effect on the abundance of shrews. The amount of urban edge had no significant effect on the captures of shrews but increased edge allows invasion of the Argentine ants, which had a highly significant negative impact on the abundance of N. crawfordi . At the trap array level, the percentage of coastal sage scrub flora had a significant positive, and the percentage of other flora had a significant negative effect on the abundance of N. crawfordi . The mean canopy height and the abundance of N. crawfordi had a significant positive effect on the occurrence of S. ornatus . Our study suggests that the loss of native coastal sage scrub flora and increasing presence of Argentine ant colonies may significantly effect the distribution and abundance of N. crawfordi . The very low overall population densities of both shrew species in most study sites make both species susceptible to extinction in isolated habitat fragments due to environmental stochasticity, and anthropogenic disturbance.

Journal of Animal Ecology

Survival costs of chick rearing in black-legged kittiwakes

1. We tested for costs of chick rearing in the black-legged kittiwake Rissa tridactyla (Linnaeus) by removing entire clutches from 149 of 405 randomly selected nests, in which one or both mates was colour-banded. After the manipulation, we monitored adult nest attendance and body condition at unmanipulated and manipulated nests, and measured the survival and fecundity of these adults the following year. 2. Late in the chick-rearing period, adults from unmanipulated nests (i.e. with chicks) went on significantly longer foraging trips, and were significantly lighter for their size, than adults from manipulated nests (i.e. without chicks). 3. Adults from unmanipulated nests also survived to the following nesting season at a significantly lower rate than those from the manipulated nests (0·898 vs. 0·953), suggesting that attempting to raise chicks can reduce life expectancy by 55%. 4. There was a tendency for adults from nests that were unmanipulated in year one to have lower reproductive success in year two, primarily because of reduced fledging success, and a higher incidence of non-breeding. 5. These findings suggest that mass loss in kittiwakes during chick rearing may not be adaptive. Raising chicks can lead to reproductive costs, and the causal mechanism appears to be a reduction in body condition. 6. We compare our results with previous brood (or clutch) size manipulation experiments that have measured adult body condition, survival and/or future fecundity. Although the empirical evidence suggests that long-lived species are more likely to experience survival costs than short-lived species, we believe the opposite may be true. We suggest that shifting the experimental protocol of cost of reproduction studies from brood enlargements (an approach taken in most prior studies) to brood reductions will provide more accurate quantifications of naturally occurring costs. 7. The cost of reproduction is one mechanism proposed to explain the reduced survival rates reported for kittiwake populations in the North Atlantic relative to those in the North Pacific ocean. Oceanographic data, however, suggest that lower food availability may limit survival of kittiwakes in the North Atlantic where a deeper mixed layer and reduced primary production combine to make conditions less favourable for this seabird during the winter months.

Alaska

Sources of variation in survival and breeding site fidelity in three species of European ducks

1. We used long-term capture-recapture-recovery data and a modelling approach developed by Burnham (1993) to test a priori predictions about sources of variation in annual survival rates and fidelity within a population of individually marked females in three species of European ducks from a breeding ground study site in Latvia. 2. True annual survival was higher for diving ducks (tufted duck 0-72, common pochard 0-65) and lower for northern shoveler (0-52). Survival of female diving ducks was positively correlated with mean winter temperatures at Western European wintering areas, the relationship being much stronger for pochard. 3. We present the first unbiased estimates of breeding fidelity and permanent emigration in European ducks. Estimated fidelity rates were high (0'88-1-0) and emigration rates low (0-0-12) for all three species, and we found strong evidence for age-specific differences in fidelity of pochards. Unusual long-distance (up to 2500 km) breeding dispersal movements that we found in female tufted ducks have not been documented in any other European waterfowl and are most probably a result of saturated nesting habitats. 4. Fidelity was a function of patch reproductive success in the previous year for all three species providing support for the idea that patch success is an important cue influencing fidelity. 5. Fidelity probability increased to 1.0 for shovelers during the last 12 years of study following provision of critical improvements in nesting habitats and suggested that habitat conditions and reproductive success determined site fidelity and settling patterns for shoveler and probably also influenced fidelity of the two other species. In predictable habitats, fidelity is a parameter that reflects the integration of fitness components and is thus a good quantity for assessing the effectiveness of habitat management actions.

Journal of Animal Ecology