Resource Patchiness as a Resolution to the Food Paradox in the Sea.

AbstractAverage concentrations of biota in the ocean are low, presenting a critical problem for ocean consumers. High-resolution sampling, however, demonstrates that the ocean is peppered with narrow hot spots of organism activity. To determine whether these resource aggregations could provide a significant solution to the ocean's food paradox, a conceptual graphical model was developed that facilitates comparisons of the role of patchiness in predator-prey interactions across taxa, size scales, and ecosystems. The model predicts that predators are more reliant on aggregated resources for foraging success when the average concentrations of resources is low, the size discrepancy between predator and prey is great, the predator has a high metabolic rate, and/or the predator's foraging time is limited. Size structure differences between marine and terrestrial food webs and a vast disparity in the overall mean density of their resources lead to the conclusion that high-density aggregations of prey are much more important to the survival of oceanic predators than their terrestrial counterparts, shaping the foraging decisions that are available to an individual and setting the stage on which evolutionary pressures can act. Patches of plenty may be rare, but they play an outsized role in behavioral, ecological, and evolutionary processes, particularly in the sea.

Sex differences in foraging ecology of a zooplanktivorous little auk Alle alle during the pre-laying period: insights from remote sensing and animal-tracking.

BACKGROUND: Energy and time allocation in seabirds differ between consecutive stages of breeding given various requirements of particular phases of the reproductive period. Theses allocations may also be sex-specific considering differential energetic or nutritional requirements of males and females and/or sexual segregation in foraging niches and/or areas. In this study we investigated the foraging ecology of an Arctic, zooplanktivorous seabird, the little auk Alle alle during the pre-laying period using remote sensing of the environment and GPS-TDR loggers deployed on birds. We compared foraging trips range and habitats of birds with other stages of the breeding period and between sexes. RESULTS: We found that little auks during the pre-laying period foraged exclusively in cold sea surface temperature zones (with temperatures < 5 ºC) but in various sea depth zones. They dived to similar depths ranging from -4.0 to -10.9 m, exploring various thermal microhabitats (with mean temperatures values ranging from 2.2 °C in Shelf sea depth zone to 5.9 °C in Deep sea depth zone). The majority of foraging trips and dives characteristics were similar to subsequent phases of breeding. However, home ranges during the pre-laying trips were wider compared to the incubation period. As expected, females exhibited wider foraging niches compared to males (wider range of sea surface temperature and sea depth in foraging locations), which could be explained by sex specific energetic and/or nutritional requirements (females producing an egg). We also delineated local foraging areas important for little auks during their whole breeding season. Protection of these areas is crucial for sustaining the local marine biodiversity. CONCLUSIONS: We found that little auks females during the pre-laying period explored wider foraging niches compared to males. These differences may be attributed to sex-specific nutritional or/and energetical constraints at this stage of breeding. The results of this study also emphasize the importance of shelf Arctic-type water masses as the foraging areas for little auks during successive stages of breeding.

Evolutionary lability of food caching behaviour in mammals.

Food hoarding provides animals access to resources during periods of scarcity. Studies on mammalian caching indicate associations with brain size, seasonality and diet but are biased to a subset of rodents. Whether the behaviour is generalizable at other taxonomic scales and/or is influenced by other ecological factors is less understood. Population density may influence food caching due to food competition or pilferage, but this remains untested in a comparative framework. Using phylogenetic analyses, we assessed the role of morphology (body and brain size), climate, diet breadth and population density on food caching behaviour evolution at multiple taxonomic scales. We also used a long-term dataset on caching behaviour of red squirrels (Tamiasciurus fremonti) to test key factors (climate and population density) on hoarding intensity. Consistent with previous smaller scale studies, we found the mammalian ancestral state for food caching was larderhoarding, and scatterhoarding was derived. Caching strategy was strongly associated with brain size, population density and climate. Mammals with larger brains and hippocampal volumes were more likely to scatterhoard, and species living at higher population densities and in colder climates were more likely to larderhoard. Finer-scale analyses within families, sub-families and tribes indicated that the behaviour is evolutionary labile. Brain size in family Sciuridae and tribe Marmotini was larger in scatterhoarders, but not in other tribes. Scatterhoarding in tribe Marmotini was more likely in species with lower population densities while scatterhoarding in tribe Sciurini was associated with warmer climates. Red squirrel larderhoarding intensity was positively related to population density but not climate, implicating food competition or pilferage as an important mechanism mediating caching behaviour. Our results are consistent with previous smaller-scale studies on food caching and indicate the evolutionary patterns of mammalian food caching are broadly generalizable. Given the lability of caching behaviour as evidenced by the variability of our results at finer phylogenetic scales, comparative analyses must consider taxonomic scale. Applying our results to conservation could prove useful as changes in population density or climate may select for different food caching strategies and thus can inform management of threatened and endangered species and their habitats.

A seabird's eye view: visual fields of some seabirds (Laridae and Procellariidae) from tropical latitudes.

The visual field of a bird defines the amount of information that can be extracted from the environment around it, using the eyes. Previous visual field research has left large phylogenetic gaps, where tropical bird species have been comparatively understudied. Using the ophthalmoscopic technique, we measured the visual fields of seven tropical seabird species, to understand what are the primary determinants of their visual fields. The visual field topographies of the seven seabird species were relatively similar, despite the two groups of Terns (Laridae) and Shearwaters (Procellariidae) being phylogenetically distant. We propose this similarity is due to their largely similar foraging ecology. These findings support previous research that foraging ecology rather than relatedness is the key determining factor behind a bird's visual field topography. Some bird species were identified to have more limited binocular fields, such as Brown Noddies (Anous stolidus) where binocularity onsets lower down within the visual field, resulting in a larger blind area about the head.

Niche partitioning and individual specialisation in resources and space use of sympatric fur seals at their range margin.

Ecological theory predicts niche partitioning between high-level predators living in sympatry as a mechanism to minimise the selective pressure of competition. Accordingly, male Australian fur seals Arctocephalus pusillus doriferus and New Zealand fur seals A. forsteri that live in sympatry should exhibit partitioning in their broad niches (in habitat and trophic dimensions) in order to coexist. However, at the northern end of their distributions in Australia, both are recolonising their historic range after a long absence due to over-exploitation, and their small population sizes suggest competition should be weak and may allow overlap in niche space. We found some niche overlap, yet clear partitioning in diet trophic level (δ15N values from vibrissae), spatial niche space (horizontal and vertical telemetry data) and circadian activity patterns (timing of dives) between males of each species, suggesting competition may remain an active driver of niche partitioning amongst individuals even in small, peripheral populations. Consistent with individual specialisation theory, broad niches of populations were associated with high levels of individual specialisation for both species, despite putative low competition. Specialists in isotopic space were not necessarily specialists in spatial niche space, further emphasising their diverse individual strategies for niche partitioning. Males of each species displayed distinct foraging modes, with Australian fur seals primarily benthic and New Zealand fur seals primarily epipelagic, though unexpectedly high individual specialisation for New Zealand fur seals might suggest marginal populations provide exceptions to the pattern generally observed amongst other fur seals.

A multicomponent approach to studying cultural propensities during foraging in the wild.

Determining the cultural propensities or cultural behaviours of a species during foraging entails an investigation of underlying drivers and motivations. In this article, we propose a multicomponent approach involving behaviour, ecology, and physiology to accelerate the study of cultural propensities in the wild. We propose as the first component the use of field experiments that simulate natural contexts, such as foraging behaviours and tool use opportunities, to explore social learning and cultural tendencies in a variety of species. To further accelerate this component, we discuss and advocate for the use of modern machine learning video analysis tools. In conjunction, we examine non-invasive methods to measure ecological influences on foraging such as phenology, fruit availability, dietary intake; and physiological influences such as stress, protein balance, energetics, and metabolism. We feature non-invasive urine sampling to investigate urea, creatinine, ketone bodies, the thyroid hormone triiodothyronine (T3), cortisol and connecting peptides of insulin. To conclude, we highlight the benefits of combining ecological and physiological conditions with behavioural field experiments. This can be done across wild species, and provides the framework needed to test ecological hypotheses related to cultural behaviour.

Following the feeder: A global synthesis of disturbance-based foraging associations of birds.

Species interactions link animal behaviour to community structure and macroecological patterns of biodiversity. One common type of trophic species interaction is disturbance foraging-the act of obtaining food at a disturbance created by another organism. Disturbance foraging is widespread across the animal kingdom, especially among birds, yet previous research has been largely anecdotal and we still lack a synthetic understanding of how this behaviour varies geographically, phylogenetically and ecologically. To address these gaps, we conducted a comprehensive literature review to test focal hypotheses about disturbance foraging behaviour in birds. We found that avian disturbance foraging was geographically ubiquitous, occurring in both aquatic and terrestrial habitats across six continents and four oceans. Consistent with predictions based on established species diversity gradients in different habitat types, the majority of terrestrial observations occurred at tropical latitudes, whereas aquatic observations took place most frequently in temperate marine waters. Although disturbance foraging was widespread across the avian phylogeny, contrary to our prediction, the behaviour was also conserved phylogenetically (Pagel's λ = 0.7) and clustered within suboscine landbirds in terrestrial environments and seabirds in aquatic environments. Similarly, although disturbers were taxonomically diverse as we predicted, interactions were unexpectedly dominated by swarm-raiding ants in terrestrial environments and cetaceans in aquatic environments. Diet and body mass were also important predictors of disturbance foraging associations: Responders followed disturbers with similar diets and larger body sizes. Overall, our hypothesis-testing framework provides insight into the importance of geography, phylogeny and ecology as predictors of disturbance foraging behaviour. We anticipate that this comprehensive assessment of disturbance foraging will serve to generate additional hypotheses and spark future research and management considerations about this fascinating but poorly studied suite of species interactions, especially as biotic interactions face unprecedented risks in our rapidly changing world.

Feeding morphology and body size shape resource partitioning in an eared seal community.

Body size and feeding morphology influence how animals partition themselves within communities. We tested the relationships among sex, body size, skull morphology and foraging in sympatric otariids (eared seals) from the eastern North Pacific Ocean, the most diverse otariid community in the world. We recorded skull measurements and stable carbon (δ13C) and nitrogen (δ15N) isotope values (proxies for foraging) from museum specimens in four sympatric species: California sea lions (Zalophus californianus), Steller sea lions (Eumetopias jubatus), northern fur seals (Callorhinus ursinus) and Guadalupe fur seals (Arctocephalus townsendi). Species and sexes had statistical differences in size, skull morphology and foraging significantly affecting the δ13C values. Sea lions had higher δ13C values than fur seals, and males of all species had higher values than females. The δ15N values were correlated with species and feeding morphology; individuals with stronger bite forces had higher δ15N values. We also found a significant community-wide correlation between skull length (indicator of body length), and foraging, with larger individuals having nearshore habitat preferences, and consuming higher trophic level prey than smaller individuals. Still, there was no consistent association between these traits at the intraspecific level, indicating that other factors might account for foraging variability.

Ecologically driven differences in individual diet specialization across three populations of Guiana dolphin.

Populations usually considered foraging generalists may include specialized individuals that feed on a restricted subset of the prey spectrum consumed by the population. By analyzing the time series of δ13C and δ15N values in sequential growth layer groups within tooth dentin, we measured population- and individual-level variation in resource use of three populations of Guiana dolphins (Sotalia guianensis)-Caravelas River, Babitonga Bay, and Norte Bay-along a latitudinal gradient in the southwestern Atlantic Ocean. We show that the Guiana dolphin at Caravelas River is a generalist population consisting of individual dietary specialists, likely due to the absence of other resident dolphin populations thus allowing individuals to target prey across a wide range of habitats. The Babitonga Bay population is also composed of individual specialists potentially due to the selective foraging behavior of some individuals on high-quality prey sources within and near the bay. In contrast, the Norte Bay population comprises individual generalists, which likely reflects its distinctive cohesive social organization, coexistence with two other dolphin species, and an opportunistic foraging strategy in response to resource fluctuations inherent to the southern limit of the species distribution. Although the Guiana dolphin is generally considered to be a dietary generalist at the population level, our findings reveal that the total niche width of populations and the degree of individual diet specialization are highly context dependent, suggesting dietary plasticity that may be related to a latitudinal gradient in resource availability and environmental conditions.

Picking pithy plants: Pith selectivity by wild white-faced capuchin monkeys, Cebus imitator.

Understanding diet selectivity is a longstanding goal in primate ecology. Deciphering when and why primates consume different resources can provide insights into their nutritional ecology as well as adaptations to food scarcity. Plant pith, the spongy interior of plant stems, is occasionally eaten by primates, but the context is poorly understood. We examine the ecological, mechanical, chemical, and nutritional basis of plant pith selection by a wild, frugivorous-omnivorous primate (Cebus imitator). We test the hypothesis that pith is a fallback food, that is, consumed when fruit is less abundant, and test for differences between plant species from which pith is eaten versus avoided. We collected 3.5 years of capuchin pith consumption data to document dietary species and analyzed "pith patch visits" in relation to fruit availability, visits to fruit patches, and climatic seasonality. We analyzed dietary and non-dietary species for relative pith quantity, mechanical hardness, odor composition, and macronutrient concentrations. Capuchins ate pith from 11 of  ~300 plant species common in the dry forest, most commonly Bursera simaruba. We find that pith consumption is not directly related to fruit availability or fruit foraging but occurs most frequently (84% of patch visits) during the months of seasonal transition. Relative to common non-dietary species, dietary pith species have relatively higher pith quantity, have softer outer branches and pith, and contain more terpenoids, a class of bioactive compounds notable for their widespread medicinal properties. Our results suggest that greater pith quantity, lower hardness, and a more complex, terpenoid-rich odor profile contribute to species selectivity; further, as pith is likely to be consistently available throughout the year, the seasonality of pith foraging may point to zoopharmacognosy, as seasonal transitions typically introduce new parasites or pathogens. Our study furthers our understanding of how climatic seasonality impacts primate behavior and sheds new light on food choice by an omnivorous primate.

Oceanic giants dance to atmospheric rhythms: Ephemeral wind-driven resource tracking by blue whales.

Trophic transfer of energy through marine food webs is strongly influenced by prey aggregation and its exploitation by predators. Rapid aggregation of some marine fish and crustacean forage species during wind-driven coastal upwelling has recently been discovered, motivating the hypothesis that predators of these forage species track the upwelling circulation in which prey aggregation occurs. We examine this hypothesis in the central California Current Ecosystem using integrative observations of upwelling dynamics, forage species' aggregation, and blue whale movement. Directional origins of blue whale calls repeatedly tracked upwelling plume circulation when wind-driven upwelling intensified and aggregation of forage species was heightened. Our findings illustrate a resource tracking strategy by which blue whales may maximize energy gain amid ephemeral foraging opportunities. These findings have implications for the ecology and conservation of diverse predators that are sustained by forage populations whose behaviour is responsive to episodic environmental dynamics.

Sowerby's beaked whale biosonar and movement strategy indicate deep-sea foraging niche differentiation in mesoplodont whales.

Closely related species are expected to diverge in foraging strategy, reflecting the evolutionary drive to optimize foraging performance. The most speciose cetacean genus, Mesoplodon, comprises beaked whales with little diversity in external morphology or diet, and overlapping distributions. Moreover, the few studied species of beaked whales (Ziphiidae) show very similar foraging styles with slow, energy-conserving movement during long, deep foraging dives. This raises the question of what factors drive their speciation. Using data from animal-attached tags and aerial imagery, we tested the hypothesis that two similar-sized mesoplodonts, Sowerby's (Mesoplodon bidens) and Blainville's (Mesoplodon densirostris) beaked whales, exploit a similar low-energy niche. We show that, compared with the low-energy strategist Blainville's beaked whale, Sowerby's beaked whale lives in the fast lane. While targeting a similar mesopelagic/bathypelagic foraging zone, they consistently swim and hunt faster, perform shorter deep dives, and echolocate at a faster rate with higher frequency clicks. Further, extensive near-surface travel between deep dives challenges the interpretation of beaked whale shallow inter-foraging dives as a management strategy for decompression sickness. The distinctively higher frequency echolocation clicks do not hold apparent foraging benefits. Instead, we argue that a high-speed foraging style influences dive duration and echolocation behaviour, enabling access to a distinct prey population. Our results demonstrate that beaked whales exploit a broader diversity of deep-sea foraging and energetic niches than hitherto suspected. The marked deviation of Sowerby's beaked whales from the typical ziphiid foraging strategy has potential implications for their response to anthropogenic sounds, which appears to be strongly behaviourally driven in other ziphiids.

Integrating plant stoichiometry and feeding experiments: state-dependent forage choice and its implications on body mass.

Intraspecific feeding choices comprise a large portion of herbivore foraging decisions. Plant resource quality is heterogeneously distributed, affected by nutrient availability and growing conditions. Herbivores navigate landscapes, foraging not only according to food qualities, but also energetic and nutritional demands. We test three non-exclusive foraging hypotheses using the snowshoe hare (Lepus americanus): (1) herbivore feeding choices and body conditions respond to intraspecific plant quality variation; (2) high energetic demands mitigate feeding responses; and (3) feeding responses are inflated when nutritional demands are high. We measured black spruce (Picea mariana) nitrogen, phosphorus and terpene compositions, as indicators of quality, within a snowshoe hare trapping grid and found plant growing conditions to explain spruce quality variation (R2 < 0.36). We then offered two qualities of spruce (H1) from the trapping grid to hares in cafeteria-style experiments and measured their feeding and body condition responses (n = 75). We proxied energetic demands (H2) with ambient temperature and coat insulation (% white coat) and nutritional demands (H3) with the spruce quality (nitrogen and phosphorus content) in home ranges. Hares with the strongest preference for high-quality spruce lost on average 2.2% less weight than hares who ate the least high-quality spruce relative to low-quality spruce. The results supported our energetic predictions as follows: hares in colder temperatures and with less-insulative coats (lower % white) consumed more spruce and were less selective towards high-quality spruce. Collectively, we found variation in plant growing conditions within herbivore home ranges substantial enough to affect herbivore body conditions, but energetic stats mediate plant-herbivore interactions.

Innate visual attraction in wood ants is a hardwired behavior seen across different motivational and ecological contexts.

Ants are expert navigators combining innate and learnt navigational strategies. Whereas we know that the ants' feeding state segregates visual-navigational memories in ants navigating along a learnt route, it is an open question if the motivational state also affects the ants' innate visual preferences. Wood ant foragers show an innate attraction to conspicuous visual cues. These foragers inhabit cluttered woodland habitat and feed on honeydew from aphids on trees. Hence, the attraction to 'tree-like' objects might be an ecologically relevant behavior that is tailored to the wood ants' foraging ecology. Foragers from other ant species with different foraging ecologies show very different innate attractions. We investigated here the innate visual response of wood ant foragers with different motivational states, i.e., unfed or fed, as well as males that show no foraging activity. Our results show that ants from all three groups orient toward a prominent visual cue, i.e., this intrinsic visuomotor response is not context-dependent, but a hardwired behavior seen across different motivational and ecological contexts. Supplementary Information: The online version contains supplementary material available at 10.1007/s00040-022-00867-3.

Ontogenetic changes in the tooth morphology of bull sharks (Carcharhinus leucas).

Teeth are an integral component of feeding ecology, with a clear link between tooth morphology and diet, as without suitable dentition prey cannot be captured nor broken down for consumption. Bull sharks, Carcharhinus leucas, undergo an ontogenetic niche shift from freshwater to marine habitats, which raises the question: does tooth morphology change with ontogeny? Tooth shape, surface area and thickness were measured using both morphometrics and elliptic Fourier analysis to determine if morphology varied with position in the jaw and if there was an ontogenetic change concordant with this niche shift. Significant ontogenetic differences in tooth morphology as a function of position in the jaw and shark total length were found, with upper and lower jaws of bull sharks presenting two different tooth morphologies. Tooth shape and thickness fell into two groupings, anterior and posterior, in both the upper and lower jaws. Tooth surface area, however, indicated three groupings, mesial, intermediate and distal, in both the upper and lower jaws. While tooth morphology changed significantly with size, showing an inflection at sharks of 135 cm total length, each morphological aspect retained the same tooth groupings throughout. These ontogenetic differences in tooth morphologies reflect tooth strength, prey handling and heterodonty.

Trophic niche of Australian cownose rays (Rhinoptera neglecta) and whitespotted eagle rays (Aetobatus ocellatus) along the east coast of Australia.

Australian cownose rays (Rhinoptera neglecta) and whitespotted eagle rays (Aetobatus ocellatus) are large myliobatiform rays that co-occur off temperate eastern Australia. Here, we performed stable-isotope analyses (δ13 C, δ15 N and δ34 S) on fin clips of both species to gain insights into their trophic interactions and isotopic niches, and assess the effect of preservation (ethanol-stored versus frozen) on isotopic values of fin-clip tissue of R. neglecta. Linear mixed models identified species as the main factor contributing to variation among δ15 N and δ34 S values, and disc width for δ13 C. Bayesian ecological niche modelling indicated a 57.4% to 74.5% overlap of trophic niches, with the niche of R. neglecta being smaller and more constrained. Because values of δ13 C were similar between species, variation in isotopic niches were due to differences in δ15 N and δ34 S values. Linear mixed models failed to detect differences in isotopic values of ethanol-stored and frozen fin tissue of R. neglecta. This study provides the first examination of the trophic ecology of R. neglecta and the comparison of isotopic niche with A. ocellatus, which will facilitate future research into the trophic interactions of these species and aid better resource management.

Phylogeny and foraging behaviour shape modular morphological variation in bat humeri.

Bats show a remarkable ecological diversity that is reflected both in dietary and foraging guilds (FGs). Cranial ecomorphological adaptations linked to diet have been widely studied in bats, using a variety of anatomical, computational and mathematical approaches. However, foraging-related ecomorphological adaptations and the concordance between cranial and postcranial morphological adaptations remain unexamined in bats and limited to the interpretation of traditional aerodynamic properties of the wing (e.g. wing loading [WL] and aspect ratio [AR]). For this reason, the postcranial ecomorphological diversity in bats and its drivers remain understudied. Using 3D virtual modelling and geometric morphometrics (GMM), we explored the phylogenetic, ecological and biological drivers of humeral morphology in bats, evaluating the presence and magnitude of modularity and integration. To explore decoupled patterns of variation across the bone, we analysed whole-bone shape, diaphyseal and epiphyseal shape. We also tested whether traditional aerodynamic wing traits correlate with humeral shape. By studying 37 species from 20 families (covering all FGs and 85% of dietary guilds), we found similar patterns of variation in whole-bone and diaphyseal shape and unique variation patterns in epiphyseal shape. Phylogeny, diet and FG significantly correlated with shape variation at all levels, whereas size only had a significant effect on epiphyseal morphology. We found a significant phylogenetic signal in all levels of humeral shape. Epiphyseal shape significantly correlated with wing AR. Statistical support for a diaphyseal-epiphyseal modular partition of the humerus suggests a functional partition of shape variability. Our study is the first to show within-structure modular morphological variation in the appendicular skeleton of any living tetrapod. Our results suggest that diaphyseal shape correlates more with phylogeny, whereas epiphyseal shape correlates with diet and FG.

Ant colony nest networks adapt to resource disruption.

Animal social structure is shaped by environmental conditions, such as food availability. This is important as conditions are likely to change in the future and changes to social structure can have cascading ecological effects. Wood ants are a useful taxon for the study of the relationship between social structure and environmental conditions, as some populations form large nest networks and they are ecologically dominant in many northern hemisphere woodlands. Nest networks are formed when a colony inhabits more than one nest, known as polydomy. Polydomous colonies are composed of distinct sub-colonies that inhabit spatially distinct nests and that share resources with each other. In this study, we performed a controlled experiment on 10 polydomous wood ant (Formica lugubris) colonies to test how changing the resource environment affects the social structure of a polydomous colony. We took network maps of all colonies for 5 years before the experiment to assess how the networks changes under natural conditions. After this period, we prevented ants from accessing an important food source for a year in five colonies and left the other five colonies undisturbed. We found that preventing access to an important food source causes polydomous wood ant colony networks to fragment into smaller components and begin foraging on previously unused food sources. These changes were not associated with a reduction in the growth of populations inhabiting individual nests (sub-colonies), foundation of new nests or survival, when compared with control colonies. Colony splitting likely occurred as the availability of food in each nest changed causing sub-colonies to change their inter-nest connections. Consequently, our results demonstrate that polydomous colonies can adjust to environmental changes by altering their social network.

Local prey shortages drive foraging costs and breeding success in a declining seabird, the Atlantic puffin.

As more and more species face anthropogenic threats, understanding the causes of population declines in vulnerable taxa is essential. However, long-term datasets, ideal to identify lasting or indirect effects on fitness measures such as those caused by environmental factors, are not always available. Here we use a single year but multi-population approach on populations with contrasting demographic trends to identify possible drivers and mechanisms of seabird population changes in the north-east Atlantic, using the Atlantic puffin, a declining species, as a model system. We combine miniature GPS trackers with camera traps and DNA metabarcoding techniques on four populations across the puffins' main breeding range to provide the most comprehensive study of the species' foraging ecology to date. We find that puffins use a dual foraging tactic combining short and long foraging trips in all four populations, but declining populations in southern Iceland and north-west Norway have much greater foraging ranges, which require more (costly) flight, as well as lower chick-provisioning frequencies, and a more diverse but likely less energy-dense diet, than stable populations in northern Iceland and Wales. Together, our findings suggest that the poor productivity of declining puffin populations in the north-east Atlantic is driven by breeding adults being forced to forage far from the colony, presumably because of low prey availability near colonies, possibly amplified by intraspecific competition. Our results provide valuable information for the conservation of this and other important North-Atlantic species and highlight the potential of multi-population approaches to answer important questions about the ecological drivers of population trends.

Using GPS tracking and stable multi-isotopes for estimating habitat use and winter range in Palearctic ospreys.

We used both satellite tracking and carbon, nitrogen and sulphur stable isotopic analysis (SIA) to infer wintering ecology and habitat use of the Corsican osprey Pandion haliaetus population. A control sample of feathers from 75 individuals was collected within the osprey's northern hemisphere breeding range, to assess the SIA variability across habitat types. An experimental set of SIA on feathers of 18 Corsican adults was examined to infer wintering ground locations and habitat types used during the non-breeding period. We calibrated the SIA using GPS/GSM tracks of 12 Mediterranean adults' movements as wintering site references. We found 50% of individuals were resident and the other half migrated. Ospreys spent the winter at temperate latitudes and showed a high plasticity in habitat selection spread over the Mediterranean basin (marine bays, coastal lagoons/marshland, inland freshwater sites). Complementary to GPS tracking, SIA is, at a broad geographical scale, a reliable method to determine whether ospreys overwinter in a habitat different from that of their breeding area. This study proved that the integration of SIA and GPS/GSM tracking techniques was effective at overcoming the intrinsic limits of each method and achieving greater information for basic ecological studies of migratory birds in aquatic environments.