Directness of resource use metrics affects predictions of bear body fat gain

Many North American ursids rely on an annual hyperphagic period to obtain fat reserves necessary for winter survival and reproduction. Identifying causes of variation in body fat gain may improve understanding of how bear resource use affects body condition. We used data from southcentral Alaska to model changes in percentage body fat of adult female American black bears (Ursus americanus) in 1998 and 2000 and brown bears (Ursus arctos) in 2000. We used year, proportion of radio locations in different habitats, distance to streams containing salmon (Onchorynchus spp.), and degree of radio location clustering as predictors for black bears and elevation, distance to streams containing salmon, and degree of radio location clustering as predictors for brown bears. Degree of location clustering was the only predictor variable supported by parameter coefficients in black bear models, supporting our hypothesis that metrics of energetics perform better as predictors of body condition than habitat use. With every unit increase in location clustering black bear body fat increased 2 %. No predictor variables influenced variation in brown bear change in body fat. Some variables previously found useful for predicting bear presence (e.g., habitat) were not useful in predicting changes in body fat, an important biological outcome for these species. Rather than assuming fitness benefits of habitat-level selection, we recommend including metrics of energetics that might more directly influence biological outcomes.     

Feeding habits may explain the morphological uniqueness of brown bears on Etorofu Island,Southern Kuril Islands in East Asia

The brown bear (Ursus arctos) population on Etorofu Island, Southern Kuril Islands, has several unique morphological features: (1) the presence of white‐pelage bears within the population and (2) a larger body size than bears on a larger neighbour island, Hokkaido Island. Nevertheless, little ecological information is available about Etorofu brown bears. In the present study, we reveal the unique feeding habits of Etorofu brown bears and suggest that their unique morphological features and diet are related. The feeding habits of brown bears on Etorofu Island were assessed using carbon, nitrogen, and sulfur stable isotope analysis, and their feeding habits were compared with those of bears on the eastern side of Hokkaido Island. According to the stable isotope analysis, the dependence on salmon is great for bears on Etorofu but only slight for bears on Hokkaido. Our results suggest that the feeding habits of Etorofu brown bears may explain their unique morphological features because a white pelage colour confers an advantage when catching salmon, and a carnivorous diet can make their body size larger. The variation in feeding habits can be an important driver of the speciation and evolution of animals.     

Individual diet specialisation in sparrows is driven by phenotypic plasticity in traits related to trade‐offs in animal performance

Individual diet specialisation (IS) is frequent in many animal taxa and affects population and community dynamics. The niche variation hypothesis (NVH) predicts that broader population niches should exhibit greater IS than populations with narrower niches, and most studies that examine the ecological factors driving IS focus on intraspecific competition. We show that phenotypic plasticity of traits associated with functional trade‐offs is an important, but unrecognised mechanism that promotes and maintains IS. We measured nitrogen isotope (δ¹⁵N) and digestive enzyme plasticity in four populations of sparrows (Zonotrichia capensis) to explore the relationship between IS and digestive plasticity. Our results show that phenotypic plasticity associated with functional trade‐offs is related in a nonlinear fashion with the degree of IS and positively with population niche width. These findings are opposite to the NVH and suggest that among individual differences in diet can be maintained via acclimatisation and not necessarily require a genetic component.     

Stress and Reproductive Hormones in Grizzly Bears Reflect Nutritional Benefits and Social Consequences of a Salmon Foraging Niche

Physiological indicators of social and nutritional stress can provide insight into the responses of species to changes in food availability. In coastal British Columbia, Canada, grizzly bears evolved with spawning salmon as an abundant but spatially and temporally constrained food source. Recent and dramatic declines in salmon might have negative consequences on bear health and ultimately fitness. To examine broadly the chronic endocrine effects of a salmon niche, we compared cortisol, progesterone, and testosterone levels in hair from salmon-eating bears from coastal BC (n = 75) with the levels in a reference population from interior BC lacking access to salmon (n = 42). As predicted, testosterone was higher in coastal bears of both sexes relative to interior bears, possibly reflecting higher social density on the coast mediated by salmon availability. We also investigated associations between the amount of salmon individual bears consumed (as measured by stable isotope analysis) and cortisol and testosterone in hair. Also as predicted, cortisol decreased with increasing dietary salmon and was higher after a year of low dietary salmon than after a year of high dietary salmon. These findings at two spatial scales suggest that coastal bears might experience nutritional or social stress in response to on-going salmon declines, providing novel insights into the effects of resource availability on fitness-related physiology.     

Beyond ecological opportunity: Prey diversity rather than abundance shapes predator niche variation

1. Ecological opportunity (i.e. the diversity of available resources) has a pivotal role in shaping niche variation and trophic specialisation of animals. However, ecological opportunity can be described with regard to both diversity and abundance of resources. The degree to which these two components contribute to niche variation remains unexplored.
2. To address this, we used an extensive dataset on fish diet and benthic invertebrate diversity and density from 73 sampling events in three Norwegian rivers in order to explore realised trophic niches and the response of dietary niche variation along gradients of resource diversity (potential trophic niches), resource density (as a proxy of resource abundance) and fish density (as a proxy of inter‐ and intra‐specific competition) in a freshwater top predator (the brown trout, Salmo trutta L.).
3. Linear models indicated that individual and population niche variation increased with increasing ecological opportunity in terms of prey diversity. However, no simple cause‐and‐effect associations between niche indices and prey abundance were found. Our multiple regression analyses indicated that the abundance of certain resources (e.g. Chironomidae) can interact with prey diversity to determine individual and population realised trophic niches. Niche variation (within‐individual component and inter‐individual diet variation) decreased with increasing inter‐ and intra‐specific competition.
4. This study extends prevailing trophic ecology theory by identifying diversity, rather than density, of available prey resources as a primary driver of niche variation in fish of temperate riverine systems with no extensive resource limitation. The study also shows that ecological opportunity may mask the direction of the effect (compression or expansion) of competition on niche variation when food resources are diverse.
5. Our study provides novel empirical insight to the driving forces behind niche variation and reveals that diversity, rather than density, of available prey resources may be a primary driver of niche variation in freshwater fish. Our study supports the view that a broader potential trophic niche promotes broader realised trophic niche variation by individuals, which leads to individual niche diversification by opening access to alternatives resources, resulting in a concomitant rise in the realised trophic niche width of the population.

     

Niche–trait relationships at individual and population level in three co‐occurring passerine species

The niche variation hypothesis (NVH) predicts that populations with wider niches exhibit greater morphological variation through increased interindividual differences in both niche and morphology. In this study, we examined niche–trait relationships in three passerine species (Cyanoderma ruficeps, Sinosuthora webbiana, and Zosterops simplex). A total of 289 C. ruficeps from 7 sites, 259 S. webbiana from 8 sites, and 144 Z. simplex from 6 sites were sampled along an elevation gradient (0–2,700 m) in Taiwan from 2009 to 2017. We measured bill traits (length, width, and depth of bill) and body size traits (length of head, tarsus, and wing) of the birds, which were reduced to four principal components (bill PC1, bill PC2, body size PC1, and body size PC2). We collected feather tissues for stable carbon and nitrogen isotope analyses to quantify their isotope niche. We quantified interindividual differences in isotope space and trait space with four diversity metrics (divergence, dispersion, evenness, and uniqueness) and tested whether interindividual differences in isotope space and trait space are positively associated. We quantified population isotope niche width by Bayesian ellipse area and population morphological variation by variances of the PCs. The results showed that individual uniqueness in isotope niche and bill morphology (average closeness of individuals within the population isotope/trait space) were positively associated across three species. Furthermore, isotope niche width and bill PC1 (reflecting the size of bill) variation at population level were also positively associated across the three species, supporting the NVH. Of the three species, C. ruficeps and S. webbiana showed stronger support for the NVH than Z. simplex, possibly due to the latter having narrower elevational distribution and a more specialized, plant‐based diet. The diversity metrics represented different aspects of interindividual differences in niche/trait space, and for the passerines, individual uniqueness appeared to play an important role in their niche–trait dynamics.     

Community structure affects trophic ontogeny in a predatory fish

While most studies have focused on the timing and nature of ontogenetic niche shifts, information is scarce about the effects of community structure on trophic ontogeny of top predators. We investigated how community structure affects ontogenetic niche shifts (i.e., relationships between body length, trophic position, and individual dietary specialization) of a predatory fish, brown trout (Salmo trutta). We used stable isotope and stomach content analyses to test how functional characteristics of lake fish community compositions (competition and prey availability) modulate niche shifts in terms of (i) piscivorous behavior, (ii) trophic position, and (iii) individual dietary specialization. Northern Scandinavian freshwater fish communities were used as a study system, including nine subarctic lakes with contrasting fish community configurations: (i) trout‐only systems, (ii) two‐species systems (brown trout and Arctic charr [Salvelinus alpinus] coexisting), and (iii) three‐species systems (brown trout, Arctic charr, and three‐spined sticklebacks [Gasterosteus aculeatus] coexisting). We expected that the presence of profitable small prey (stickleback) and mixed competitor–prey fish species (charr) supports early piscivory and high individual dietary specialization among trout in multispecies communities, whereas minor ontogenetic shifts were expected in trout‐only systems. From logistic regression models, the presence of a suitable prey fish species (stickleback) emerged as the principal variable determining the size at ontogenetic niche shifts. Generalized additive mixed models indicated that fish community structure shaped ontogenetic niche shifts in trout, with the strongest positive relationships between body length, trophic position, and individual dietary specialization being observed in three‐species communities. Our findings revealed that the presence of a small‐sized prey fish species (stickleback) rather than a mixed competitor–prey fish species (charr) was an important factor affecting the ontogenetic niche‐shift processes of trout. The study demonstrates that community structure may modulate the ontogenetic diet trajectories of and individual niche specialization within a top predator.     

Alaskan brown bears (Ursus arctos) aggregate and display fidelity to foraging neighborhoods while preying on Pacific salmon along small streams

The interaction between brown bears (Ursus arctos) and Pacific salmon (Oncorhynchus spp.) is important to the population dynamics of both species and a celebrated example of consumer‐mediated nutrient transport. Yet, much of the site‐specific information we have about the bears in this relationship comes from observations at a few highly visible but unrepresentative locations and a small number of radio‐telemetry studies. Consequently, our understanding of brown bear abundance and behavior at more cryptic locations where they commonly feed on salmon, including small spawning streams, remains limited. We employed a noninvasive genetic approach (barbed wire hair snares) over four summers (2012–2015) to document patterns of brown bear abundance and movement among six spawning streams for sockeye salmon, O. nerka, in southwestern Alaska. The streams were grouped into two trios on opposite sides of Lake Aleknagik. Thus, we predicted that most bears would forage within only one trio during the spawning season because of the energetic costs associated with swimming between them or traveling around the lake and show fidelity to particular trios across years because of the benefits of familiarity with local salmon dynamics and stream characteristics. Huggins closed‐capture models based on encounter histories from genotyped hair samples revealed that as many as 41 individuals visited single streams during the annual 6‐week sampling season. Bears also moved freely among trios of streams but rarely moved between these putative foraging neighborhoods, either during or between years. By implication, even small salmon spawning streams can serve as important resources for brown bears, and consistent use of stream neighborhoods by certain bears may play an important role in spatially structuring coastal bear populations. Our findings also underscore the efficacy of noninvasive hair snagging and genetic analysis for examining bear abundance and movements at relatively fine spatial and temporal scales.     

Pulsed resource availability changes dietary niche breadth and partitioning between generalist rodent consumers

Identifying the mechanisms that structure niche breadth and overlap between species is important for determining how species interact and assessing their functional role in an ecosystem. Without manipulative experiments, assessing the role of foraging ecology and interspecific competition in structuring diet is challenging. Systems with regular pulses of resources act as a natural experiment to investigate the factors that influence the dietary niches of consumers. We used natural pulses of mast‐fruiting of American beech (Fagus grandifolia) to test whether optimal foraging or competition structure the dietary niche breadth and overlap between two congener rodent species (Peromyscus leucopus and P. maniculatus), both of which are generalist consumers. We reconstructed diets seasonally over a 2‐year period using stable isotope analysis (δ¹³C, δ¹⁵N) of hair and of potential dietary items and measured niche dynamics using standard ellipse area calculated within a Bayesian framework. Changes in niche breadth were generally consistent with predictions of optimal foraging theory, with both species consuming more beechnuts (a high‐quality food resource) and having a narrower niche breadth during masting seasons compared to nonmasting seasons when dietary niches expanded and more fungi (a low‐quality food source) were consumed. In contrast, changes in dietary niche overlap were consistent with competition theory, with higher diet overlap during masting seasons than during nonmasting seasons. Overall, dietary niche dynamics were closely tied to beech masting, underscoring that food availability influences competition. Diet plasticity and niche partitioning between the two Peromyscus species may reflect differences in foraging strategies, thereby reducing competition when food availability is low. Such dietary shifts may have important implications for changes in ecosystem function, including the dispersal of fungal spores.     

Functional macronutritional generalism in a large omnivore,the brown bear

We combine a recently developed framework for describing dietary generalism with compositional data analysis to examine patterns of omnivory in a large widely distributed mammal. Using the brown bear (Ursus arctos) as a model species, we collected and analyzed data from the literature to estimate the proportions of macronutrients (protein, carbohydrate, and lipid) in the diets of bear populations. Across their range, bears consumed a diversity of foods that resulted in annual population diets that varied in macronutrient proportions, suggesting a wide fundamental macronutrient niche. The variance matrix of pairwise macronutrient log‐ratios indicated that the most variable macronutrient among diets was carbohydrate, while protein and lipid were more proportional or codependent (i.e., relatively more constant log‐ratios). Populations that consumed anthropogenic foods, such agricultural crops and supplementary feed (e.g., corn), had a higher geometric mean proportion of carbohydrate, and lower proportion of protein, in annual diets. Seasonally, mean diets were lower in protein and higher in carbohydrate, during autumn compared to spring. Populations with anthropogenic subsidies, however, had higher mean proportions of carbohydrate and lower protein, across seasons compared to populations with natural diets. Proportions of macronutrients similar to those selected in experiments by captive brown bears, and which optimized primarily fat mass gain, were observed among hyperphagic prehibernation autumn diets. However, the majority of these were from populations consuming anthropogenic foods, while diets of natural populations were more variable and typically higher in protein. Some anthropogenic diets were close to the proportions selected by captive bears during summer. Our results suggest that omnivory in brown bears is a functional adaptation enabling them to occupy a diverse range of habitats and tolerate variation in the nutritional composition and availability of food resources. Furthermore, we show that populations consuming human‐sourced foods have different dietary macronutrient proportions relative to populations with natural diets.     

Ontogenetic and individual diet variation in amphibian larvae across an environmental gradient

1. Variation among individuals within size or age classes can have profound effects on community dynamics and food‐web structure. We investigated the potential influence of habitat disturbance on intrapopulation niche variation. 2. Amphibians occupy a range of lentic habitats from short‐hydroperiod intermittent ponds to long‐hydroperiod permanent ponds. We quantified ontogenetic diet variation and individual specialisation in wood frog tadpoles (Lithobates sylvaticus) and blue‐spotted salamander larvae (Ambystoma laterale) to investigate the influence of hydroperiod on population niche width across a natural hydroperiod gradient using stable isotope and gut content analyses. In one of the few tests using larval forms, we tested the niche variation hypothesis, which predicts that populations with larger niche widths also have increased individual variation. 3. Our results support the niche variation hypothesis, indicating that more generalised populations exhibit higher within‐individual diet variation. We report gradual changes in the relative importance of diet items, decreased dietary overlap and increased trophic position in L. sylvaticus throughout development. A. laterale became more enriched in δ¹³C and increased in δ¹⁵N throughout its larval period. We did not find a relationship between hydroperiod and niche parameters, indicating that niches are conserved across heterogeneous habitats. In contrast to most documented cases, we estimated low levels of individual specialisation in amphibian larvae. 4. Amphibians are an important link between aquatic and terrestrial ecosystems, whereby diet shifts can influence food‐web structure by altering energy flow pathways and the trophic position of higher consumers, ultimately changing food‐chain length.     

Latitudinal variation in ecological opportunity and intraspecific competition indicates differences in niche variability and diet specialization of Arctic marine predators

Individual specialization (IS), where individuals within populations irrespective of age, sex, and body size are either specialized or generalized in terms of resource use, has implications on ecological niches and food web structure. Niche size and degree of IS of near‐top trophic‐level marine predators have been little studied in polar regions or with latitude. We quantified the large‐scale latitudinal variation of population‐ and individual‐level niche size and IS in ringed seals (Pusa hispida) and beluga whales (Delphinapterus leucas) using stable carbon and nitrogen isotope analysis on 379 paired ringed seal liver and muscle samples and 124 paired beluga skin and muscle samples from eight locations ranging from the low to high Arctic. We characterized both within‐ and between‐individual variation in predator niche size at each location as well as accounting for spatial differences in the isotopic ranges of potential prey. Total isotopic niche width (TINW) for populations of ringed seals and beluga decreased with increasing latitude. Higher TINW values were associated with greater ecological opportunity (i.e., prey diversity) in the prey fish community which mainly consists of Capelin (Mallotus villosus) and Sand lance (Ammodytes sp.) at lower latitudes and Arctic cod (Boreogadus saida) at high latitudes. In beluga, their dietary consistency between tissues also known as the within‐individual component (WIC) increased in a near 1:1 ratio with TINW (slope = 0.84), suggesting dietary generalization, whereas the slope (0.18) of WIC relative to TINW in ringed seals indicated a high degree of individual specialization in ringed seal populations with higher TINWs. Our findings highlight the differences in TINW and level of IS for ringed seals and beluga relative to latitude as a likely response to large‐scale spatial variation in ecological opportunity, suggesting species‐specific variation in dietary plasticity to spatial differences in prey resources and environmental conditions in a rapidly changing ecosystem.     

Environmental stability increases relative individual specialisation across populations of an aquatic top predator

The concept of the niche has long been a central pillar in ecological theory, with a traditional focus on quantifying niches at the species or population level. However, the importance of individual‐level niche variation is increasingly being recognised, with a strong focus on individual specialisation. While examples illustrating the contribution of the individual niche to whole population niche structure are accumulating rapidly, surprisingly little is known about the conditions that shape the differences between these two potentially divergent components. Though theory predicts that stability should influence the extent of such intra‐specific specialisation, we know of no previous study that has investigated its role in individual specialisation, and the differentiation between individual‐ and population niches. Here, we studied the diet of individuals from multiple populations of an aquatic top‐predator, Salmo trutta, inhabiting contrasting stable, groundwater fed and unstable, surface water fed pre‐alpine streams. Based on stomach content analysis, we found that individuals living in stable environments displayed a higher degree of specialisation than those in unstable environments, with the between‐individual component of niche width being approximately twice as high in the former. We subsequently validated these results by evidence gained from stable isotope analysis of muscle tissue. As such, we reveal that environmental stability can significantly influence individual niches within populations, leading to increased specialisation.     

Dining from the coast to the summit: Salmon and pine nuts determine the summer body condition of female brown bears on the Shiretoko Peninsula

Body condition in mammals fluctuates depending on energy intake and expenditure. For brown bears (Ursus arctos), high‐protein foods facilitate efficient mass gain, while lipids and carbohydrates play important roles in adjusting dietary protein content to optimal levels to maximize energy intake. On the Shiretoko Peninsula, Hokkaido, Japan, brown bears have seasonal access to high‐lipid pine nuts and high‐protein salmon. To assess seasonal and annual fluctuation in the body condition of adult female brown bears in relation to diet and reproductive status, we conducted a longitudinal study in a special wildlife protection area on the Shiretoko Peninsula during 2012–2018. First, analyses of 2,079 bear scats revealed that pine nuts accounted for 39.8% of energy intake in August and salmon accounted for 46.1% in September and that their consumption by bears varied annually. Second, we calculated the ratio of torso height to torso length as an index of body condition from 1,226 photographs of 12 adult females. Results indicated that body condition continued to decline until late August and started to increase in September when salmon consumption increased. In addition, body condition began to recover earlier in years when consumption of both pine nuts and salmon was high. Furthermore, females with offspring had poorer body condition than solitary females, in particular in late August in years with low salmon consumption. Our findings suggest that coastal and subalpine foods, which are unique to the Shiretoko Peninsula, determine the summer body condition of female brown bears, as well as their survival and reproductive success.     

Population-level resource selection by sympatric brown and American black bears in Alaska

Distribution theory predicts that for two species living in sympatry, the subordinate species would be constrained from using the most suitable resources (e.g., habitat), resulting in its use of less suitable habitat and spatial segregation between species. We used negative binomial generalized linear mixed models with fixed effects to estimate seasonal population-level resource selection at two spatial resolutions for female brown bears (Ursus arctos) and female American black bears (U. americanus) in southcentral Alaska during May–September 2000. Black bears selected areas occupied by brown bears during spring which may be related to spatially restricted (i.e., restricted to low elevations) but dispersed or patchy availability of food. In contrast, black bears avoided areas occupied by brown bears during summer. Brown bears selected areas near salmon streams during summer, presumably to access spawning salmon. Use of areas with high berry production by black bears during summer appeared in response to avoidance of areas containing brown bears. Berries likely provided black bears a less nutritious, but adequate food source. We suggest that during summer, black bears were displaced by brown bears, which supports distribution theory in that black bears appeared to be partially constrained from areas containing salmon, resulting in their use of areas containing less nutritious forage. Spatial segregation of brown and American black bears apparently occurs when high-quality resources are spatially restricted and alternate resources are available to the subordinate species. This and previous work suggest that individual interactions between species can result in seasonal population-level responses.     

Nutritional Consequences of Experimentally Introduced Tourism in Brown Bears

Abstract: Although numerous studies have documented behavioral effects of nature-based tourism on wildlife populations, few studies have determined whether behavioral changes translate to effects on individual condition and population health. This issue is currently a concern for wildlife managers in Alaska, USA, and Canada where bear viewing is a rapidly growing industry expanding into previously undisturbed bear habitats. Rather than record observations at long established tourism sites, we experimentally introduced bear viewing into 2 relatively undisturbed brown bear (Ursus arctos) populations in south-central Alaska. We examined the nutritional consequences of behavioral changes induced by the presence and activity of bear viewers for bears feeding on early summer vegetation and late-summer salmon (Oncorhynchus kisutch and O. nerka). We used Global Positioning System collars, monitored food resource availability, and quantified individual resource use and condition for a year prior to and during the introduction of bear viewing. Though bear viewing altered spatiotemporal resource use in all treatments, total resource use declined only when we exposed bears to 24-hour daily human activity. Energy expenditure, indexed as daily travel distances, was significantly higher when bears responded by altering spatial rather than temporal resource use. However, body weight and composition were unaffected by all treatments as bears shifted their foraging to other locations or times. Managers can minimize nutritional impacts of bear-viewing programs by avoiding spatial displacement and providing predictable time periods when bears can access food resources free of human activity. Bears in this study exhibited a high degree of behavioral plasticity, which may be an important factor in identifying flagship species for sustainable ecotourism programs.     

Community structure of a Neotropical bat fauna as revealed by stable isotope analysis: Not all species fit neatly into predicted guilds

Neotropical bat communities are among the most diverse mammal communities in the world, and a better understanding of these assemblages may permit inferences about how so many species coexist. While broad trophic guilds (e.g., frugivore, insectivore) of bats are recognized, details of diet and similarities among species remain largely unknown. We used stable isotope ratios of carbon (δ¹³C) and nitrogen (δ¹⁵N) to characterize the community structure of a diverse Neotropical bat fauna from Belize to test predictions of niche theory and the competitive exclusion principle. We predicted that (1) interspecific variation in isotopic overlap would be greater within guilds than between guilds, and (2) no two sympatric populations would have isotopic niches that overlap completely, unless there is variation along some other axis (e.g., temporal, spatial). We additionally tested body size as an explanatory metric of potential overlap and predicted that larger‐bodied animals would have greater niche breadths. Results suggest that while guild‐level characterizations of communities are at least somewhat informative, there are multiple examples of intra‐ and inter‐guild species pairs with significantly overlapping isotopic niches, suggesting that, counter to predictions, they may compete for resources. Understanding the trophic structure of animal communities is fundamental to conservation and management of endangered species and ecosystems and important for evolutionary studies, and stable isotope analyses can provide key insights as well as informing hypotheses of the diet of species that are not well known. Abstract in Spanish is available with online material.     

Feeding strategies,resource utilisation and potential mechanisms for competitive coexistence of Atlantic salmon and alpine bullhead in a sub-Arctic river

We scrutinised the seasonal food-niche utilisation of river dwelling Atlantic salmon parr and alpine bullhead in order to examine potential mechanisms that may facilitate coexistence of species with similar niches. Fish were sampled monthly during the ice-free season, and diet composition and feeding strategy of the two species were compared by analyses of stomach contents. The dietary niches and feeding strategy of salmon parr and bullheads were highly similar both at the individual and population levels, with high within-phenotype contributions to niche width and pronounced generalisation observed during time periods with severe resource limitations. Our findings suggest that competitive coexistence with similar niches may be facilitated by a generalisation of niche width as predicted by optimal foraging theory, rather than the specialised niche width predicted by classic niche theory as a response to interspecific competition. Competitive coexistence may be particularly widespread in spatially and temporally dynamic habitats such as northern lotic systems, which thus may select for generalisation and convergence of ecological niches.     

Overlap and partitioning of the ecological and isotopic niches

Recently, it was proposed that stable isotope patterns can be used to quantify the width of the ecological niche of animals. However, the potential effects of habitat use on isotopic patterns of consumers have not been fully explored and consequently isotopic patterns may yield deceptive estimates of niche width. Here, we simulated four different scenarios of a consumer foraging across an isotopically heterogeneous landscape to test the combined effects of habitat and diet selection on the widths of the isotopic niche. We then modeled the actions of a naïve researcher who randomly sampled consumers from the simulated populations, and used these results to assess the overlap and partitioning of the isotopic and the ecological niches when habitat‐derived differences among isotope signatures are not considered. Our results suggest that populations of dietary specialists exhibited broader isotopic niches than populations composed of dietary generalists, and habitat generalists exhibited narrower isotopic niche widths compared with populations of individuals that foraged in specific habitats. The conversion of isotopic niches to ecological niches without knowledge of foraging behavior and habitat‐derived isotopic differences transformed an informative δ‐space into ‘a blurry p‐space’. Therefore, knowledge of habitat‐derived differences in stable isotope values and understanding of habitat use and individual foraging behavior are critical for the correct quantification of the ecological niche.     

Landscape heterogeneity drives intra‐population niche variation and reproduction in an arctic top predator

While intra‐population variability in resource use is ubiquitous, little is known of how this measure of niche diversity varies in space and its role in population dynamics. Here we examined how heterogeneous breeding environments can structure intra‐population niche variation in both resource use and reproductive output. We investigated intra‐population niche variation in the Arctic tundra ecosystem, studying peregrine falcon (Falco peregrinus tundrius, White) breeding within a terrestrial‐marine gradient near Rankin Inlet, Nunavut, Canada. Using stable isotope analysis, we found that intra‐population niches varied at the individual level; we examined within‐nest and among‐nest variation, though only the latter varied along the terrestrial‐marine gradient (i.e., increased among‐nest variability among birds nesting within the marine environment, indicating higher degree of specialization). Terrestrial prey species (small herbivores and insectivores) were consumed by virtually all falcons. Falcons nesting within the marine environment made use of marine prey (sea birds), but depended heavily on terrestrial prey (up to 90% of the diet). Using 28‐years of peregrine falcon nesting data, we found a positive relationship between the proportion of terrestrial habitat surrounding nest sites and annual nestling production, but no relationship with the likelihood of successfully rearing at least one nestling reaching 25 days old. Annually, successful inland breeders raised 0.47 more young on average compared to offshore breeders, which yields potential fitness consequences for this long‐living species. The analyses of niche and reproductive success suggest a potential breeding cost for accessing distant terrestrial prey, perhaps due to additional traveling costs, for those individuals with marine nest site locations. Our study indicates how landscape heterogeneity can generate proximate (niche variation) and ultimate (reproduction) consequences on a population of generalist predator. We also show that within‐individual and among‐individual variation are not mutually exclusive, but can simultaneously arise and structure intra‐population niche variation.