Morphological distance and inter-nest distance account for intra-specific prey overlap in digger wasps (Hymenoptera: Crabronidae)

Although inter-individual diet variation is common in predatory wasp populations, the factors accounting for such variation are still largely unknown. Here, we asked if paired diet dissimilarity in three species of digger wasps correlates with morphological distance and inter-nest distance, two factors previously linked to diet partitioning in vertebrates. Results sharply differed among species and generations. All sampled populations showed significant inter-individual diet variation for prey taxa, but only in half of the cases for prey size. In one generation of two species [Bembix zonata Klug and Stizus continuus (Klug)], similar-sized wasps had similar prey taxonomic spectra (and for S. continuus also similar prey size spectra), a phenomenon which probably reduces intra-specific competition. In addition, B. zonata females nesting closer to each other had more similar prey taxonomic spectra, suggesting that distant females probably hunt on different patches that harbour different prey species. For the females of a further species (Bembix merceti Parker), pairwise size difference and inter-nest distance did not affect prey dissimilarity. Both morphological distance and inter-nest distance are potentially important in shaping the overlap of individual resource use in wasps, though probably only in certain conditions such as a highly clumped distribution of nests and size-related constraints on prey selection.     

Generalist niche, specialist strategy: the diet of an Australian percichthyid

Dietary analysis revealed that an impoundment population of Australian bass Macquaria novemaculeata holds a generalist niche, but one arising from persistent individual specialization and interindividual variation. This 'individual specialist' strategy appeared adaptive, but the strength of individual specialization was largely independent of variation in diet composition, except during blooms of Daphnia sp. Diet composition and dietary overlap showed only moderate ontogenetic variation, and niche breadth showed no relationship with ontogeny. Macquaria novemaculeata showed an asymmetric predator and prey size distribution, consistent with many aquatic predators, with positive relationships between fish size and average, maximum and minimum prey size. There was no asymmetry in the relative size-based niche breadths of individuals, however, which indicates that the niche is a fixed 'window' of relative prey sizes. The difference in the dietary niche and prey-size relationships of M. novemaculeata at the population and individual levels highlights the necessity of assessing the niche at both these levels.     

Specialisation in prey capture drives coexistence among sympatric spider‐hunting wasps

1. Sister taxa that coexist in the same space and time often face competition due to the use of similar resources. However, some closely related species can adopt fine‐grained specialisation in resource use to coexist. This study investigated niche overlap between three sympatric spider‐hunting wasp species of the genus Trypoxylon (Hymenoptera: Crabronidae) known to nest in three of the habitats found in the study area. 2. First, the co‐occurrence of these wasp species in the three habitats was estimated, as a proxy for potential competition. Then, the following hypotheses were tested: (i) niche partitioning is seen more often between species that co‐occur in a habitat, whereas there is niche overlap between species nesting in distinct habitats (prey specialisation hypothesis); and (ii) wasp species capture prey according to their size (physical constraint hypothesis). 3. Two pairs of wasp species were found consistently nesting in the same habitat. Niche partitioning based on prey taxa occurred regardless of the habitat preference. It was also found that differences in the size of wasps reflected distinctions in the size of their prey. 4. These findings were consistent over the years, showing that the significance of specialisation in foraging activities and physical constraints during prey capture can play key roles in the coexistence of sympatric species. The distinctions in the foraging strategies of these wasps are discussed, as well as potential mechanisms driving the evolution in prey specialisation, with insights for future studies.     

Food niche overlap and ecological separation in a multi-species community of shrews in the Siberian taiga

An investigation of the feeding habits and prey availability in a community of seven species of shrew (Insectivora: Soricidae) inhabiting the taiga of Central Siberia was carried out with the aim of quantifying levels of niche overlap and elucidating modes of ecological separation amongst these coexisting species. All species took a wide range of invertebrate prey, and overlap in the numbers of shared prey taxa was high, but differences in dietary composition of certain taxa reduced overlap between most species. Small species fed almost exclusively on small arthropods, mostly Araneae, Chilopoda and Coleoptera, while medium and large-sized species took high proportions of oligochaetes. Prey were mostly taken in proportions approximately equal to their availability, although certain prey appeared to be selected. All shrews took prey in a range of sizes, and the high dietary occurrence of small invertebrates reflected their availability and high encounter rate in field samples. Dietary occurrence of small prey was negatively correlated, and large prey positively correlated, with body size of shrew. Smaller shrews were predominantly ground-surface foragers while larger species were more subterranean, with body size and dietary occurrence of soil prey being positively correlated. Differences in prey size and foraging mode reduced niche overlap between shrew species of widely differing sizes. Each shrew species did not occupy a separate, well-defined food niche. Instead, the community was sub-divided into three functional groups: large and small species which tended towards specialization with relatively low levels of overlap, and intermediate, generalist species with higher levels of overlap.     

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.     

Does prey mobility affect niche width and individual specialization in hunting wasps? A network‐based analysis

Niche width and inter‐individual diet variation in predator populations are known to be affected by intrinsic factors such as body size, age, cognitive constraints, and by intra‐ and interspecific competition. By contrast, how variation in prey biological traits may affect niche width and partitioning is still a poorly explored topic. One of these candidate traits is prey mobility, which can affect the predators’ niche because acting on the rate of encounter and, assuming mobility as a proxy for escape capability, on the success of predator attacks. Here we analysed 20 wasp populations and their prey as individual‐based food‐webs to test if prey mobility may explain niche width (here defined by the Shannon entropy‐based index ) and patterns of inter‐individual diet variation (here defined by the interaction‐exclusiveness index H₂′ and the interaction evenness index E₂). Niche width was very variable among populations and overall network specialization (H₂′) was always higher than the expected by null models. In case of high‐speed flying prey (e.g. flies, bees), wasps showed wider niches () and lower specialization (H₂′) than in case of non‐flying or slow‐flying prey (e.g. spiders, beetles). Evenness (E₂), on the other side, did not vary with prey mobility. Altogether, these results suggest that highly elusive prey may lead to wider predators’ niche but somehow limit their individual niche specialization.     

Taxon‐biased diet preference in the ‘generalist’ beetle‐hunting wasp Cerceris rubida provides insights on the evolution of prey specialization in apoid wasps

Opportunism and specialization appear to be widespread in apoid wasps, although the factors affecting the diet preference (and thus explaining the degree of specialization) are still largely unknown. Four hypotheses that stressed the importance of the size, sex, habitat, and taxonomic identity of prey of the beetle‐hunting digger wasp, Cerceris rubida, were formulated and tested. The wasp population hunted for phytophagous beetles belonging to abundant families around the wasp nesting site. In practice, the prey appeared to be hunted only in two cultivated fields, thus habitat accounted for a majority of the observed diet. The size of wasps was furthermore correlated with the size of their prey, and thus this also accounted for the frequencies of hunted prey and the strong individual specialization for both taxa and size. However, in the exploited habitat, some species were significantly over‐hunted than expected and some other significantly avoided by the wasps, causing an unexpected major role of prey taxon on the probability of being hunted, over the other explanatory variables (body size, body shape, sex, availability). This contrasts to that found in other wasp species, which appear to select prey basing essentially on their ecology and size or their relative abundance (opportunism). The results obtained in the present study show that even an apparent ‘generalist’ predator may turn out to be taxonomically specialized. Together with a re‐evaluation of previous studies, our results further suggest that the effect of size constraints and the developmental plan of prey (holometaboulous versus hemimetabolous) may have promoted either taxonomic opportunism or specialization in different lineages of apoid wasps. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 544–558.     

Feeding ecology of the endangered Valencia toothcarp,Valencia hispanica (Actinopterygii: Valenciidae)

Patterns of food resource utilisation by Valencia toothcarp were studied in a small coastal lagoon in the Iberian Peninsula, its most northern population. Emphasis was placed on feeding activity, diet composition, seasonal and intraspecific variation in diet and feeding strategy. Important features of the overall feeding pattern of the Valencia toothcarp are decreased feeding activity during winter, concentration on benthic invertebrates and narrow dietary breadth, with a focus on gammarid amphipods. The feeding of V. hispanica was independent of seasonal variation in food availability, because its main prey were much the same throughout the year. There were no differences in seasonal niche width and niche overlap between seasons was high. There are also no differences in diets between sexes, although diet composition, niche width and niche overlap differ between juvenile and adult fish. The Valencia toothcarp population from Santes Creus lagoon was formed by specialist individuals, with narrow niche widths, that fed on four preferential prey types, but also consumed some occasional prey.     

The influence of habitat accessibility on the dietary and morphological specialisation of an aquatic predator

Individual diet and habitat specialisation are widespread in animal taxa and often related to levels of predation and competition. Mobile consumers such as predatory fish can stabilise lake food webs by ranging over a larger area than their prey, thereby switching between habitats. Although, this switching assumes that the predator has equal preference for the available prey, individual diet specialisation and morphological adaptations to different habitats could potentially prevent individuals from switching between habitats. In this study, we assessed the niche width and individual specialisation in Eurasian perch Perca fluviatilis in response to a shift in habitat use by manipulating the ability for this top predator to couple habitats. We ran an eight weeks pond experiment, to test the effect of habitat switching on diet and morphological specialisations. We show that habitat coupling influenced individual diet specialisation and niche use in expected directions where specialisation increased with decreasing habitat switching. In contrast to expectations, the morphological variation decreased with increasing diet specialisation. Our results expand on previous work and suggest that individual specialisation and niche width can impact the ability of mobile predators to couple habitats. Furthermore, it shows the importance of individual specialisations in relation to habitat coupling.     

Dietary niche breadth in a local community of passerine birds: an analysis using phylogenetic contrasts

The analysis of a local community of forest passerines (13 species) using phylogenetic contrasts shows a correlation between body size of bird species and mean prey size, minimum prey size, maximum prey size and the size range of dietary items. This suggests that larger birds drop small prey taxa from their prey list, because of the difficulty of capturing very small prey, for energetic reasons or because of microhabitat usage. We find some support for the third hypothesis. Dietary niche breadth calculated across prey taxa is not related to body size. Dietary niche breadth, however, is correlated with size-corrected measurements of the bill and locomotor apparatus. Long and slender bills increase the dietary niche breadth. Thus subtle differences constrain foraging and the techniques of extracting certain prey taxa form crevices. Dietary niche breadth and foraging diversity are positively correlated with population density: at least locally dietary generalists occur at higher breeding densities than specialists.     

Prey use of wetland benthivorous sunfishes: ontogenetic, interspecific and seasonal variation

The intensity of competitive interactions between fishes is partly determined by prey use and ontogenetic niche shifts. In a wetland where distinct habitat shifts are missing we compared prey use of three generalist benthivorous sunfishes to look for evidence of ontogenetic, interspecific, and “seasonal” variation in prey composition. Diet analysis revealed evidence of diet ontogeny in warmouth (Lepomis gulosus, 30–152 mm standard length, SL), but not in bluespotted sunfish (Enneacanthus gloriosus, 30–47 mm SL) or dollar sunfish (Lepomis marginatus, 30–60 mm SL). Bluespotted and dollar sunfishes consumed small dipteran and amphipod prey and had similar diets in both seasons suggesting a potential for strong interspecific competition. In the dry season, warmouth shifted from using smaller insect prey to larger decapod and fish prey with increasing size. This shift to prey types that were little used by the other species reduced dietary niche overlap with the other sunfishes. After drought and re-flooding (in the wet season), decapods and small fish were less abundant in the wetland and the warmouth ontogenetic shift was less distinct. When matched for gape width, prey composition differed between warmouth and both dollar and bluespotted sunfishes in the wet season, suggesting differences in sunfish foraging modes, but prey use differences were less clear in the dry season when prey were abundant. Both warmouth ontogenetic diet shifts and seasonal variation in prey use (probably mediated by prey abundance) had strong influences on diet overlap and therefore the potential for intra- and interspecific competition between sunfishes in this wetland ecosystem.     

The effects of morphology,phylogeny and prey availability on trophic resource partitioning in an anuran community

Several factors influence the partitioning of trophic resources in ecological communities, such as morphology, evolutionary history, and resource availability. Although the effects of morphology, phylogeny, and resource availability on trophic ecology have long been explored by theoretical studies, little has been done to empirically test these relationships. Here, we tested whether phylogenetic and morphological distances correlate with trophic niche overlap using a path analysis of multiple partial regression of distance matrices. Also, we tested whether niche breadth is influenced by body size using Phylogenetic Generalized Least Squares analysis. Trophic niche overlap was better explained by morphology per se than by the phylogenetic distance. We also found that predator's body size influences niche breadth calculated considering prey traits and availability, but not when we do not include these availability data. Additionally, trophic niche breadth was usually smaller when we considered prey traits and availability, differently from niche overlap, whose values increased when we did not consider these data. Our findings show that the interpretation of trophic niche in communities changes if we consider availability data, affecting inferences about coexistence and trophic specialization. Our study contributes to understanding trophic specialization and emphasizes the importance of incorporating prey availability and their traits into diet analysis.     

Dietary overlap and seasonality in three species of mormoopid bats from a tropical dry forest

Competing hypotheses explaining species’ use of resources have been advanced. Resource limitations in habitat and/or food are factors that affect assemblages of species. These limitations could drive the evolution of morphological and/or behavioural specialization, permitting the coexistence of closely related species through resource partitioning and niche differentiation. Alternatively, when resources are unlimited, fluctuations in resources availability will cause concomitant shifts in resource use regardless of species identity. Here, we used next‐generation sequencing to test these hypotheses and characterize the diversity, overlap and seasonal variation in the diet of three species of insectivorous bats of the genus Pteronotus. We identified 465 prey (MOTUs) in the guano of 192 individuals. Lepidoptera and Diptera represented the most consumed insect orders. Diet of bats exhibited a moderate level of overlap, with the highest value between Pteronotus parnellii and Pteronotus personatus in the wet season. We found higher dietary overlap between species during the same seasons than within any single species across seasons. This suggests that diets of the three species are driven more by prey availability than by any particular predator‐specific characteristic. P. davyi and P. personatus increased their dietary breadth during the dry season, whereas P. parnellii diet was broader and had the highest effective number of prey species in all seasons. This supports the existence of dietary flexibility in generalist bats and dietary niche overlapping among groups of closely related species in highly seasonal ecosystems. Moreover, the abundance and availability of insect prey may drive the diet of insectivores.     

Intra‐ and inter‐individual variation show distinct trends as drivers of seasonal changes in the resource use of a neotropical marsupial

Individual specialization can influence important ecological and evolutionary traits and both inter‐ and intra‐individual variation in resource use can drive niche shifts in natural populations. We evaluated the predominance of these two factors for determining seasonal differences in the trophic niche of the didelphid marsupial Gracilinanus agilis (Burmeister, 1854) in the highly seasonal Brazilian savanna. In the three sampled sites, the population of G. agilis increased its dietary niche width in the warm–wet season, when food resources are more abundant, and there were no differences between sexes and no interaction between season and sex. However, the evaluation of intra‐individual variation indicated that females reduce the number of items consumed during the warm–wet season, whereas males show no seasonal differences. Inter‐individual variation nonetheless followed the overall population pattern because both sexes increased their spread with respect to food‐item consumption in the warm–wet season. Additionally, we found positive relationships between body length and diet only in the warm–wet season, when larger animals fed more on invertebrates and less on fruits than the small ones. Our results show a previously unknown pattern for mammals, in which the trophic niche is wider during the high‐resource season as a result of inter‐individual variation along the body‐size axis. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 737–747.     

Patterns of coexistence in sexual and asexual species of Cnemidophorus lizards

Summary The lizard genus Cnemidophorus (family Teiidae) contains sexual as well as parthenogenetic species. The theoretical two-fold fitness advantage of asexuality does not translate into any obvious distributional or numerical superiority of the parthenogenic species in the southwestern US and northern Mexico where their ranges overlap. I tested the prediction that the genetically diverse sexual species should have a higher between-individual niche width than a similar sympatric asexual species by studying the prey in stomach contents of sympatric and allopatric populations of C. tigris (sexual) and C. sonorae (asexual) in southern Arizona. The expectation proved true for niche breadths based on both prey length and prey taxa categories. The within-individual component of niche breadth was not different between species. Meaningful comparisons between species in sympatry and allopatry are confounded by the uncontrolled differences in the availability and diversity of food items between sites. Before the generality of these results can be assessed the study should be repeated in other areas where sexual and asexual species are syntopic and of similar body size.     

The ecology of individuals: incidence and implications of individual specialization

Most empirical and theoretical studies of resource use and population dynamics treat conspecific individuals as ecologically equivalent. This simplification is only justified if interindividual niche variation is rare, weak, or has a trivial effect on ecological processes. This article reviews the incidence, degree, causes, and implications of individual-level niche variation to challenge these simplifications. Evidence for individual specialization is available for 93 species distributed across a broad range of taxonomic groups. Although few studies have quantified the degree to which individuals are specialized relative to their population, between-individual variation can sometimes comprise the majority of the population's niche width. The degree of individual specialization varies widely among species and among populations, reflecting a diverse array of physiological, behavioral, and ecological mechanisms that can generate intrapopulation variation. Finally, individual specialization has potentially important ecological, evolutionary, and conservation implications. Theory suggests that niche variation facilitates frequency-dependent interactions that can profoundly affect the population's stability, the amount of intraspecific competition, fitness-function shapes, and the population's capacity to diversify and speciate rapidly. Our collection of case studies suggests that individual specialization is a widespread but underappreciated phenomenon that poses many important but unanswered questions.     

Seasonal variation in terrestrial resource subsidies influences trophic niche width and overlap in two aquatic snake species: a stable isotope approach

Quantifying diet is essential for understanding the functional role of species with regard to energy processing, transfer, and storage within ecosystems. Recently, variance structure in the stable isotope composition of consumer tissues has been touted as a robust tool for quantifying trophic niche width, a task that has previously proven difficult due to bias in direct dietary analyses and difficulties in integrating diet composition over time. We used carbon and nitrogen stable isotope analyses to examine trophic niche width of two sympatric aquatic snakes, banded watersnakes Nerodia fasciata and black swamp snakes Seminatrix pygaea inhabiting an isolated wetland where seasonal migrations of amphibian prey cause dramatic shifts in resource availability. Specifically, we characterized snake and prey isotope compositions through time, space, and ontogeny and examined isotope values in relation to prey availability and snake diets assessed by gut content analysis. We determined that prey cluster into functional groups based on similarity of isotopic composition and seasonal availability. Isotope variance structure indicated that the trophic niche width of the banded watersnake was broader (more generalist) than that of the black swamp snake. Banded watersnakes also exhibited seasonal variation in isotope composition, suggesting seasonal diet shifts that track amphibian prey availability. Conversely, black swamp snakes exhibited little seasonal variation but displayed strong ontogenetic shifts in carbon and nitrogen isotope composition that closely paralleled ontogenetic shifts in their primary prey, paedomorphic mole salamanders Ambystoma talpoideum. Although niche dimensions are often treated as static, our results demonstrate that seasonal shifts in niche dimensions can lead to changes in niche overlap between sympatric species. Such short‐term fluctuations in niche overlap can influence competitive interactions and consequently the composition and dynamics of communities and ecosystems.     

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.     

Diet niche relationships among North American grassland and shrubsteppe birds

Summary We consider the dietary relationships of the numerically dominant breeding bird species in four North American grassland/shrubsteppe habitats, sampled over 2–3 consecutive years. Overall, the diets of these species contained primarily insects: orthopterans comprised 29% of the diet biomass, coleopterans 24%, and lepidopteran larvae 23%, while seeds contributed 15% of the average diet. These diets varied substantially, however, and we evaluated several aspects of this variation. Intersexual differences in diets within a species were few, despite the occurrence of significant sexual size dimorphism in several species. For many species, however, there were substantial shifts in dietary composition between years at a given location; overall, the average between-year similarity of species' dietary composition was 70%. Different species exhibited rather different diet patterns. Horned Larks were relatively omnivorous, had broad diet composition niches, and varied considerably in diets between different locations. Meadowlarks were also broad-niched and geographically variable in their diets, but were the most highly carnivorous of the species we considered. Dietary niche breadths of Grasshopper Sparrows were intermediate, but diet composition was rather stable, both between years and between locations. Chestnut-collared Longspurs exhibited narrow diet niches, but substantial annual variation: each year this species apparently exploited a different but limited set of prey types rather heavily. Larger avian predators generally consumed a broader array of functional groups of prey, but did not differ in the taxonomic variety of their diets from small birds. Variation in diet composition between individuals within local populations was considerable; in most species, an individual contained on the average 30–40% of the prey taxa represented in entire population smaples.Patterns of dietary overlap among species were quite inconsistent from year to year at most locations, although at the shrubsteppe site overlap among all species present was consistently quite high. Relatively few cooccurring species pairs exhibited low diet overlap. The degree of diet niche overlap was unrelated to body size differences of the birds, despite as much as six-fold differences in weight among some coexisting species. Relationships of the bird species on another dimension of the trophic niche, prey size, also differed substantially between sites and years. The ranking of co-occurring species by the mean sizes of the prey they consumed generally did not parallel their rankings by body sizes, and in some cases the smallest and the largest species present ate prey of similar sizes. At the shrubsteppe site, all the breeding species exhibited quite similar frequency distributions of prey sizes in their diets.As species number and diversity increased in the breeding avifaunas, diet niche breadths generally decreased, species packing by prey size decreased, and diet composition niche overlap remained relatively unchanged. These trends are in at least partial agreement with predictions of diffuse competition theory, but the patterns were derived from broad inter-site comparisons of overall site averages, and the relationships generally did not hold within local assemblages of species. In general, our attempts to match values of dietary niche features with site characteristics failed to demonstrate close agreement with the predictions of prevailing ecological theory based upon assumptions of resource limitation and competition. Instead, our findings seem generally most consistent with the suggestion that food is not normally limiting to bird populations in these systems, and individuals and populations are exploiting the food resources in an opportunistic fashion, which leads to considerable individual, between-year, and between-location variation in diet compositions and interspecific overlaps.Our attempts to discern clear relationships that accord with theoretical expectations in these avian assemblages are thwarted by our lack of detailed information on the resource base and by the lack of clear tests that will separate alternative hypotheses of community organization and structuring. We suggest that these complications may compromise the findings of many community studies.     

Individual specialization in diet by a generalist marine predator reflects specialization in foraging behaviour

1. We studied chick diet in a known-age, sexed population of a long-lived seabird, the Brünnich's guillemot (Uria lomvia), over 15 years (N = 136; 1993-2007) and attached time-depth-temperature recorders to examine foraging behaviour in multiple years (N = 36; 2004-07). 2. Adults showed specialization in prey fed to offspring, described by multiple indices calculated over 15 years: 27% of diet diversity was attributable to among-individual variation (within-individual component of total niche width = 0.73); average similarity of an individual's diet to the overall diet was 65% (mean proportional similarity between individuals and population = 0.65); diet was significantly more specialized than expected for 70% of individuals (mean likelihood = 0.53). These indices suggest higher specialization than the average for an across-taxa comparison of 49 taxa. 3. Foraging behaviour varied along three axes: flight time, dive depth and dive shape. Individuals showed specialized individual foraging behaviour along each axis. These foraging strategies were reflected in the prey type delivered to their offspring and were maintained over scales of hours to years. 4. Specialization in foraging behaviour and diet was greater over short time spans (hours, days) than over long time spans (years). Regardless of sex or age, the main component of variation in foraging behaviour and chick diet was between individuals. 5. Plasma stable isotope values were similar across years, within a given individual, and variance was low relative to that expected from prey isotope values, suggesting adult diet specialized across years. Stable isotope values were similar among individuals that fed their nestlings similar prey items and there was no difference in trophic level between adults and chicks. We suggest that guillemots specialize on a single foraging strategy regardless of whether chick-provisioning and self-feeding. With little individual difference in body mass and physiology, specialization likely represents learning and memorizing optimal feeding locations and behaviours. 6. There was no difference in survival or reproductive success between specialists and generalists, suggesting these are largely equivalent strategies in terms of evolutionary fitness, presumably because different strategies were advantageous at different levels of prey abundance or predictability. The development of individual specialization may be an important precursor to diversification among seabirds.