Interindividual Diet Variation Research Articles

Diet and between-tissue isotope comparisons reveal different foraging strategies for age and sex of a Saffron Finch (Sicalis flaveola Linnaeus, 1766) population.

Measuring stable isotopes in different tissues offers the opportunity to provide insight into the foraging ecology of a species. This study aimed to assess how diet varies between yellow females, yellow males, and dull individuals of a Saffron Finch (Sicalis flaveola) population. We measured δ13C and δ15N in blood over a year, and in different feathers, to estimate seasonal consistency of resource use for each category. We conducted this study in a private farm in the Central Brazilian savannas. We sampled 195 individuals in seven field samplings between January 2017 and March 2018. The mean blood δ13C values were similar among yellow females, yellow males and dull individuals, indicating that this population of Saffron Finch predominantly accesses similar resources throughout the year, with a predominant C4 signal. Although Saffron Finch is considered a granivorous species, the mean δ15N values found indicate that both adults and juveniles also incorporate in their tissues some invertebrate. The slight isotope-tissue difference between feathers and blood is similar to the reported in previous studies and may reflect tissue-to-tissue discrimination. The isotopic space of yellow males was greater than that of yellow females and dull individuals, indicating greater dietary diversity due to greater inter-individual variation in diet. In Saffron Finch, which delays plumage maturation, competition-driven partitioning of food resources seems essential in driving carotenoid-based plumage coloration between age classes and sexes.

Food web of lizard species in a land-bridge island from Western Mexico

Island ecosystems provide valuable opportunities to explore some aspects associated with resource partitioning. While some ecological studies have suggested that island populations could keep a narrow dietary niche due to limited food sources, recent studies challenge this long-held belief for insular lizards. Herein, we evaluated the trophic web of the lizard community from María Cleofas Island in the Gulf of California, Western Mexico. We quantified the inter-individual diet variation for each lizard species, compared prey diversity between species, and analyzed any potential variations across different years. Our research revealed a wide variety of arthropods within the stomach contents of the lizards, regardless of their foraging strategy and habitat use. Our results also showed the inclusion of prey items that have not been previously reported in studies focusing on mainland-island populations. For example, we observed Anolis nebulosus predating cockroaches, hemipterans, and isopods. We observed differences in prey groups among lizard species, highlighting the broader niche of Anolis nebulosus, Phyllodactylus cleofasensis, and Aspidoscelis communis, as well as the narrowest trophic niche of Ctenosaura pectinata. Our research on the lizard community of María Cleofas Island has not only demonstrated the wide dietary diversity among species but has also expanded our understanding of trophic relationships in island ecosystems, with important implications for ecological studies.

Rare and Hungry: Feeding Ecology of the Golden Alpine Salamander, an Endangered Amphibian in the Alps.

Amphibians are considered critical species in the nutrient flow within and across ecosystems, and knowledge on their trophic ecology and niches is crucial for their conservation. For the first time we studied the trophic ecology of the rare and endemic Salamandra atra aurorae in a mixed temperate forest in northern Italy. We aimed to define the realized trophic niche, investigate the prey selectivity and explore possible levels of individual specialization. In summer 2022 we obtained stomach contents from 53 salamanders by stomach flushing and prey availability using pitfall traps. We used the Costello graphical method to analyse the realized trophic niche, and the relativized electivity index to study prey selectivity. Our results show that the Golden Alpine salamander adopts a generalist feeding strategy with positive selection of few prey categories (e.g., Myriapoda, Hymenoptera except Formicidae). Food preference seems to be driven by size, movement ability and chitinization of the prey. A high degree of inter-individual diet variation, modularity and clustering was found, describing a scenario that can be framed in a Distinct Preference model framework. This study gives new insights on the trophic ecology of the Alpine salamander complex, whose subspecies appear to adopt similar feeding strategies.

Differences in individual bowhead whale Balaena mysticetus habitat use, foraging dive depth and diet during the peak feeding season

Shifts in zooplankton quantity and quality caused by climate change could challenge the ability of bowhead whales to meet their energetic requirements. When facing such selection pressure, intra-population variation dampens the negative effects and provides population-level resilience. Previous studies observed inter-individual diet variation in bowhead whales, but the mechanism responsible for the variation was undetermined. We investigated foraging variability in Eastern Canada-West Greenland bowhead whales using dietary biomarkers (stable isotopes, fatty acids) and movement data (satellite telemetry with time-depth recorders) from the same individuals. We found that bowhead whale individuals using distinct summer and fall foraging habitats displayed differences in horizontal movements, foraging dive depth, and diet. For individuals using the Canadian Arctic Archipelago habitat (Foxe Basin, Gulf of Boothia, Prince Regent Inlet, Lancaster Sound and Admiralty Inlet, Nunavut), they performed long distance movements across regions, and their foraging dive depth was generally shallow, but increased from July to November. These whales displayed higher δ13C and δ15N values and ratios of C16:1n7/C16:0. Individuals using the West Baffin Bay habitat (Cumberland Sound, Baffin Bay, Davis Strait) were more localized in their horizontal movements and consistent over time in their foraging dive depth, which was generally deeper. These whales displayed lower δ13C and δ15N values and ratios of C16:1n7/C16:0. Overall, this inter-individual variation in diet and foraging behaviour could indicate some niche variation which would be beneficial for the population under changing habitats and prey availability.

Inter-individual variation in the diet within a group of Japanese macaques and its relationship with social structure investigated by stable isotope and DNA analyses.

We investigated individual variation in diet in relation to age-sex class and kin relationship in 28 of 40 members of a small group of wild Japanese macaques (Macaca fuscata). We used stable isotope ratios from hair as an index of individual dietary profiles, genetic relatedness as an index of kin relationship, and mitochondrial DNA (mtDNA) haplotype as a marker of being an immigrant or native member of the group. The range of carbon and nitrogen stable isotope ratios from hair of individual macaques (δ13C: -24.1‰ to -22.6‰, δ15N:3.8-5.5‰), which reflected their diet over a period of ~ 6months, implied small individual variation in diet. The results of PERMANOVA implied that there were no significant effects of age class, sex, or mtDNA haplotype on hair stable isotope ratios between individuals, or on the variation in individual diet. However, the isotope values of males with mtDNA haplotypes that differed from those of the native females appeared to differ from those of other group members, which implies that immigrant males might have had a different diet profile from that of native group members. Furthermore, there was a weak correlation trend between genetic relatedness and differences in stable isotope ratios between pairs of individuals. Differences in stable isotope values were more marked in pairs with a more distant genetic relationship. This implies that within the group, closely related kin tended to forage together to avoid competing for food. However, this effect might have been weak because the size of the group was small relative to the size of the food patches, thereby reducing competition.

Patterns of diet composition of a whiptail lizard species conforms to the Shared Preferences Model of interindividual variation in prey consumption

Abstract We analysed the interindividual variation in diet, patterns of food resource use and topology of the trophic networks of juvenile, male, and female Ameivula ocellifera (Spix, 1825) during one year, covering wet and dry seasons. Using a network-based approach, we evaluated the degree of variation in the diet of individuals, and the degree of nestedness and clustering to identify patterns of resource use and the topology of individuals food webs. We found a high degree of interindividual variation in the diet of juveniles, males, and females during the study period, which seems to be related to nested patterns of resource use of males and females. These results also suggest that males and females seem to have both generalist and specialist individuals, of which the diets of specialists are nested within the diets of generalists. For juveniles, we found evidence of interindividual variation in diet in both seasons, although patterns of food resource partition that may cause it could not be identified. Insect larvae of several groups were the main food resource in all three lizard groups during the wet season, while termites were the dominant food item during the dry season. Our results conform to the Shared Preferences Model where individuals should consume the same top-ranked prey when they are abundant in the environment, replacing it with suboptimal items as preferred ones become scarce.

A Midsummer Night’s Diet: Snapshot on Trophic Strategy of the Alpine Salamander, Salamandra atra

Information on the trophic ecology of the Alpine salamander, Salamandra atra, is scattered and anecdotal. We studied for the first time the trophic niche and prey availability of a population from an area located in Italian Dolomites during the first half of August. Considering that S. atra is a typical nocturnal species, we collected food availability separately for diurnal and nocturnal hours. Our aims were: (i) to obtain information on the realized trophic niche; (ii) to provide a direct comparison between trophic strategy considering only nocturnal preys or considering all preys; (iii) to study trophic strategy of this species at the individual level. In two samplings nights we obtained prey from 50 individuals using stomach flushing technique. Trophic strategy was determined using the graphical Costello method and selectivity using the relativized electivity index. During the short timeframe of our sample, this salamander showed a generalized trophic strategy. The total trophic availability differed significantly from nocturnal availability. Interindividual diet variation is discussed in the light of the optimal diet theory. Finally, we highlighted that considering or not the activity time of the studied taxon and its preys may lead to a conflicting interpretation of the trophic strategies.

Individual–resource networks reveal distinct fruit preferences of selective individuals from a generalist population of the Helmeted Manakin

Ecological studies traditionally assume that generalist populations are homogeneous in the use of food resources, but empirical evidence supports that intraspecific differences in morphology, physiology and behaviour affect foraging decisions and promote diet variation among individuals. Furthermore, the temporal availability of resources may shape the dynamics of population trophic niche, which ultimately depends on individual niches. In this study, we investigated the seasonal changes in individual‐based networks between the Helmeted Manakin Antilophia galeata, a generalist frugivorous bird, and fruiting plants, following theoretical models of interindividual diet variation based on the Optimal Diet Theory. Selective individuals were the majority of the generalist population of the Helmeted Manakin. Our results suggest that the structure of the individual‐resource networks varied seasonally. We found that modularity was higher than expected by chance in the wet season, when fruit availability was also higher. In the dry season, modules disappeared and the network became more nested. These findings are consistent with the Distinct Preference Model of diet variation. We suggest that downscaling ecological networks to the individual level may reveal emergent properties that, albeit existent, are not evident in species–resources networks.

Food habits and ontogenetic dietary partitioning of American crocodiles in a tropical Pacific Island in Central America

AbstractStudies on food habits are fundamental to understanding the ecology of a species and its interactions with the community to which it belongs. Among crocodylians, diet affects a variety of biological, physiological, and behavioral characteristics. However, despite having one of the largest distributions across the Americas, some aspects of Crocodylus acutus’ natural history remain poorly studied, particularly in insular areas. We characterized American crocodiles’ food habits in Coiba Island, Panama, assessing ontogenetic dietary variation and dietary overlap by age group and size. We captured and collected stomach content samples from 49 individuals from four transects from March to December 2013. From these samples, we could taxonomically identify three phyla, four subphyla, eight classes, 11 orders, 17 families, 14 genera, and 12 species as prey items. However, not all samples could be identified to the lowest taxon (species), having most of them identified only to family level. Large juveniles had the largest proportion of prey items and subadults the largest proportion of gastroliths and vegetal content. Percent occurrence per major categories (insects, arachnids, crustaceans, fish, reptiles, birds, and mammals) showed crustaceans and insects as the most prominent groups of prey items on this island. Overlapping group analysis showed a reduction in the consumption of invertebrates (crustaceans and insects) as individuals aged. However, these items were the most common throughout all American crocodiles sampled. Dietary overlap analyses showed a likely ontogenetic dietary partitioning with high overlap (>60%) between small and large juveniles and low overlap (<30%) among small juveniles, subadults, and adults. To date, 71 species have been reported as prey items for American crocodiles. However, relying on prey items identified only to genus, we had at least 97 prey items. Thus, C. acutus can be defined as generalist with a broad spectrum of prey inhabiting all types of habitats and having all types of consumption classifications. Overall, American crocodiles inhabiting coastal areas present some differences in both dietary composition and structure with those dwelling inland and freshwater habitats as well as an interindividual diet variation, which reflects the plasticity and adaptability of C. acutus to a variety of conditions.

Generalisation within specialization: inter-individual diet variation in the only specialized salamander in the world.

Specialization is typically inferred at population and species level but in the last decade many authors highlighted this trait at the individual level, finding that generalist populations can be composed by both generalist and specialist individual. Despite hundreds of reported cases of individual specialization there is a complete lack of information on inter-individual diet variation in specialist species. We studied the diet of the Italian endemic Spectacled Salamander (Salamandrina perspicillata), in a temperate forest ecosystem, to disclose the realised trophic niche, prey selection strategy in function of phenotypic variation and inter-individual diet variation. Our results showed that Salamandrina is highly specialized on Collembola and the more specialized individuals are the better performing ones. Analyses of inter-individual diet variation showed that a subset of animals exhibited a broader trophic niche, adopting different foraging strategies. Our findings reflects the optimal foraging theory both at population and individual level, since animals in better physiological conditions are able to exploit the most profitable prey, suggesting that the two coexisting strategies are not equivalent. At last this species, feeding on decomposers of litter detritus, could play a key role determining litter retention rate, nutrient cycle and carbon sequestration.

From video recordings to whisker stable isotopes: a critical evaluation of timescale in assessing individual foraging specialisation in Australian fur seals.

Estimating the degree of individual specialisation is likely to be sensitive to the methods used, as they record individuals' resource use over different time-periods. We combined animal-borne video cameras, GPS/TDR loggers and stable isotope values of plasma, red cells and sub-sampled whiskers to investigate individual foraging specialisation in female Australian fur seals (Arctocephalus pusillus doriferus) over various timescales. Combining these methods enabled us to (1) provide quantitative information on individuals' diet, allowing the identification of prey, (2) infer the temporal consistency of individual specialisation, and (3) assess how different methods and timescales affect our estimation of the degree of specialisation. Short-term inter-individual variation in diet was observed in the video data (mean pairwise overlap = 0.60), with the sampled population being composed of both generalist and specialist individuals (nested network). However, the brevity of the temporal window is likely to artificially increase the level of specialisation by not recording the entire diet of seals. Indeed, the correlation in isotopic values was tighter between the red cells and whiskers (mid- to long-term foraging ecology) than between plasma and red cells (short- to mid-term) (R(2) = 0.93-0.73 vs. 0.55-0.41). δ(13)C and δ(15)N values of whiskers confirmed the temporal consistency of individual specialisation. Variation in isotopic niche was consistent across seasons and years, indicating long-term habitat (WIC/TNW = 0.28) and dietary (WIC/TNW = 0.39) specialisation. The results also highlight time-averaging issues (under-estimation of the degree of specialisation) when calculating individual specialisation indices over long time-periods, so that no single timescale may provide a complete and accurate picture, emphasising the benefits of using complementary methods.

Individual variation in anthropogenic resource use in an urban carnivore

With increasing urbanization, some animals are adapting to human-dominated systems, offering unique opportunities to study individual adaptation to novel environments. One hypothesis for why some wildlife succeed in urban areas is that they are subsidized with anthropogenic food. Here, we combine individual-level movement patterns with diet composition based on stable isotope analysis to assess the degree to which a rapidly growing population of coyotes (Canis latrans) in Chicago consumes anthropogenic resources. We used telemetry to classify coyotes into three groups based on social class and home range composition: (1) residents with home ranges in urban nature preserves; (2) residents with home ranges that had a high proportion of urban land; and (3) transients that had relatively large home ranges and variable use of urban land. We found that natural and anthropogenic resources in this system can be reliably partitioned with carbon isotopes. Mixing models revealed that resident coyotes associated with most urban nature preserves consumed trace to minimal amounts of anthropogenic resources, while coyotes that live in the urban matrix consume moderate (30-50%) to high (>50%) proportions of anthropogenic resources. Lastly, we found evidence of prey switching between natural and anthropogenic resources and a high degree of inter-individual variation in diet among coyotes. In contrast to the expectation that urban adaptation may dampen ecological variation, our results suggest individuality in movement and diet exemplifies the successful establishment of coyotes in urban Chicago. Our study also suggests that direct anthropogenic food subsidization is not a prerequisite for successful adaptation to urban environments.

Niche expansion in a resilient endemic species following introduction of a novel top predator

Summary Introduced species can have profound direct ecological impacts on native species, yet their potential indirect effects remain relatively unexplored. For instance, introduced predators may directly affect some native species via predation, which may in turn have indirect consequences for other species that are released from competition. We explore this possibility in East Africa's Lake Victoria basin, where the introduction of the predatory Nile perch, Lates niloticus, in the 1950s and 60s contributed to the overall or local extinction of hundreds of native fish by the 1980s. We ask whether this dramatic change in assemblage composition has led to competitive release and niche expansion in Rastrineobola argentea, a resilient native cyprinid that has thrived in this highly perturbed ecosystem. To address this question, we compare the trophic ecology of R. argentea before (1966) and after (2011) the introduction of the Nile perch in Lake Victoria; and across eight satellite lakes that differ in their history of Nile perch invasion. Using this combination of spatial and temporal comparisons, we test for increases in dietary niche breadth (niche expansion) and changes in the level of individual specialisation of R. argentea in invaded versus uninvaded contexts. In our historical comparison, we find good evidence for dietary niche expansion and an increase in interindividual diet variation in R. argentea over time. Across lakes, however, strong bottom‐up effects (i.e. variation in prey availability) appear to obscure any potential top‐down effects of the Nile perch introduction on the trophic ecology of R. argentea. Overall, we find substantial temporal and spatial variation in the diet and niche breadth of R. argentea, but the underlying drivers remain uncertain, given the complexity of both anthropogenic and natural ecological changes in Lake Victoria over the past century. Understanding both the direct and indirect impacts of introduced species is challenging, but important for successful long‐term management of human‐altered ecosystems.

Ecosystem fragmentation drives increased diet variation in an endemic livebearing fish of the Bahamas.

One consequence of human-driven habitat degradation in general, and habitat fragmentation in particular, is loss of biodiversity. An often-underappreciated aspect of habitat fragmentation relates to changes in the ecology of species that persist in altered habitats. In Bahamian wetlands, ecosystem fragmentation causes disruption of hydrological connectivity between inland fragmented wetlands and adjacent marine areas, with the consequent loss of marine piscivores from fragmented sections. We took advantage of this environmental gradient to investigate effects of ecosystem fragmentation on patterns of resource use in the livebearing fish Gambusia hubbsi (Family Poeciliidae), using both population- and individual-level perspectives. We show that fragmentation-induced release from predation led to increased G. hubbsi population densities, which consequently led to lower mean growth rates, likely as a result of higher intraspecific competition for food. This was accompanied by a broadening of dietary niches via increased interindividual diet variation, suggesting a negative effect of predation and a positive effect of intraspecific competition on the degree of diet variation in natural populations. Our results therefore indicate that habitat fragmentation can greatly impact the ecology of resilient populations, with potentially important ecological and evolutionary implications.

Complex-to-predict generational shift between nested and clustered organization of individual prey networks in digger wasps.

Although diet has traditionally been considered to be a property of the species or populations as a whole, there is nowadays extensive knowledge that individual specialization is widespread among animal populations. Nevertheless, the factors determining the shape of interactions within food webs remain largely undiscovered, especially in predatory insects. We used an aggregation of the digger wasp Bembix merceti to 1) analyse patterns of individual prey use across three flying seasons in a network–based context; and 2) test the effect of four potential factors that might explain network topologies (wasp mass, nest spatial distribution, simultaneous nest-provisioning, prey availability). Inter-individual diet variation was found in all three years, under different predator-prey network topologies: Individuals arranged in dietary clusters and displayed a checkerboard pattern in 2009, but showed nestedness in 2008 and 2010. Network topologies were not fully explained by the tested factors. Larger females consumed a higher proportion of the total number of prey species captured by the population as a whole, in such a way that nested patterns may arise from mass-dependent prey spectrum width. Conversely, individuals with similar body mass didn’t form clusters. Nested patterns seemed to be associated with a greater availability of the main prey species (a proxy for reduced intra-specific competition). Thus, according with theory, clusters seemed to appear when competition increased. On the other hand, the nests of the individuals belonging to a given cluster were not more closely located, and neither did individuals within a cluster provision their nests simultaneously. Thus, a female-female copying behaviour during foraging was unlikely. In conclusion, wasp populations can maintain a considerable individual variation across years under different food web organizations. The tested factors only partially accounted for the shift in network properties, and new analyses should be carried out to elucidate how diet network topologies arise in wasp populations.

Multi-scale effects of nestling diet on breeding performance in a terrestrial top predator inferred from stable isotope analysis.

Inter-individual diet variation within populations is likely to have important ecological and evolutionary implications. The diet-fitness relationships at the individual level and the emerging population processes are, however, poorly understood for most avian predators inhabiting complex terrestrial ecosystems. In this study, we use an isotopic approach to assess the trophic ecology of nestlings in a long-lived raptor, the Bonelli’s eagle Aquila fasciata, and investigate whether nestling dietary breath and main prey consumption can affect the species’ reproductive performance at two spatial scales: territories within populations and populations over a large geographic area. At the territory level, those breeding pairs whose nestlings consumed similar diets to the overall population (i.e. moderate consumption of preferred prey, but complemented by alternative prey categories) or those disproportionally consuming preferred prey were more likely to fledge two chicks. An increase in the diet diversity, however, related negatively with productivity. The age and replacements of breeding pair members had also an influence on productivity, with more fledglings associated to adult pairs with few replacements, as expected in long-lived species. At the population level, mean productivity was higher in those population-years with lower dietary breadth and higher diet similarity among territories, which was related to an overall higher consumption of preferred prey. Thus, we revealed a correspondence in diet-fitness relationships at two spatial scales: territories and populations. We suggest that stable isotope analyses may be a powerful tool to monitor the diet of terrestrial avian predators on large spatio-temporal scales, which could serve to detect potential changes in the availability of those prey on which predators depend for breeding. We encourage ecologists and evolutionary and conservation biologists concerned with the multi-scale fitness consequences of inter-individual variation in resource use to employ similar stable isotope-based approaches, which can be successfully applied to complex ecosystems such as the Mediterranean.

The marine side of a terrestrial carnivore: intra-population variation in use of allochthonous resources by arctic foxes.

Inter-individual variation in diet within generalist animal populations is thought to be a widespread phenomenon but its potential causes are poorly known. Inter-individual variation can be amplified by the availability and use of allochthonous resources, i.e., resources coming from spatially distinct ecosystems. Using a wild population of arctic fox as a study model, we tested hypotheses that could explain variation in both population and individual isotopic niches, used here as proxy for the trophic niche. The arctic fox is an opportunistic forager, dwelling in terrestrial and marine environments characterized by strong spatial (arctic-nesting birds) and temporal (cyclic lemmings) fluctuations in resource abundance. First, we tested the hypothesis that generalist foraging habits, in association with temporal variation in prey accessibility, should induce temporal changes in isotopic niche width and diet. Second, we investigated whether within-population variation in the isotopic niche could be explained by individual characteristics (sex and breeding status) and environmental factors (spatiotemporal variation in prey availability). We addressed these questions using isotopic analysis and Bayesian mixing models in conjunction with linear mixed-effects models. We found that: i) arctic fox populations can simultaneously undergo short-term (i.e., within a few months) reduction in both isotopic niche width and inter-individual variability in isotopic ratios, ii) individual isotopic ratios were higher and more representative of a marine-based diet for non-breeding than breeding foxes early in spring, and iii) lemming population cycles did not appear to directly influence the diet of individual foxes after taking their breeding status into account. However, lemming abundance was correlated to proportion of breeding foxes, and could thus indirectly affect the diet at the population scale.

The nested assembly of individual-resource networks

1. Much of the current understanding of ecological systems is based on theory that does not explicitly take into account individual variation within natural populations. However, individuals may show substantial variation in resource use. This variation in turn may be translated into topological properties of networks that depict interactions among individuals and the food resources they consume (individual-resource networks). 2. Different models derived from optimal diet theory (ODT) predict highly distinct patterns of trophic interactions at the individual level that should translate into distinct network topologies. As a consequence, individual-resource networks can be useful tools in revealing the incidence of different patterns of resource use by individuals and suggesting their mechanistic basis. 3. In the present study, using data from several dietary studies, we assembled individual-resource networks of 10 vertebrate species, previously reported to show interindividual diet variation, and used a network-based approach to investigate their structure. 4. We found significant nestedness, but no modularity, in all empirical networks, indicating that (i) these populations are composed of both opportunistic and selective individuals and (ii) the diets of the latter are ordered as predictable subsets of the diets of the more opportunistic individuals. 5. Nested patterns are a common feature of species networks, and our results extend its generality to trophic interactions at the individual level. This pattern is consistent with a recently proposed ODT model, in which individuals show similar rank preferences but differ in their acceptance rate for alternative resources. Our findings therefore suggest a common mechanism underlying interindividual variation in resource use in disparate taxa.

Individual‐level diet variation in four species of Brazilian frogs

1. Many natural populations exploiting a wide range of resources are actually composed of relatively specialized individuals. 2. This interindividual variation is thought to be a consequence of the invasion of 'empty' niches in depauperate communities, generally in temperate regions. If individual niches are constrained by functional trade-offs, the expansion of the population niche is only achieved by an increase in interindividual variation, consistent with the 'niche variation hypothesis'. 3. According to this hypothesis, we should not expect interindividual variation in species belonging to highly diverse, packed communities. 4. In the present study, we measured the degree of interindividual diet variation in four species of frogs of the highly diverse Brazilian Cerrado, using both gut contents and delta(13)C stable isotopes. 5. We found evidence of significant diet variation in the four species, indicating that this phenomenon is not restricted to depauperate communities in temperate regions. 6. The lack of correlations between the frogs' morphology and diet indicate that trade-offs do not depend on the morphological characters measured here and are probably not biomechanical. The nature of the trade-offs remains unknown, but are likely to be cognitive or physiological. 7. Finally, we found a positive correlation between the population niche width and the degree of diet variation, but a null model showed that this correlation can be generated by individuals sampling randomly from a common set of resources. Therefore, albeit consistent with, our results cannot be taken as evidence in favour of the niche variation hypothesis.

Intrapopulation Diet Variation in Four Frogs (Leptodactylidae) of the Brazilian Savannah

Age and sex-based as well as individual-level diet variation are known to occur in many natural populations, and may have important ecological and evolutionary implications. In the case of individual-level diet variation, most examples come from species-poor, temperate communities, and it is currently believed that it results from population niche expansion following interspecific competitive release. We investigated and measured the intrapopulation diet variation in four species of frogs, Leptodactylus (=Adenomera) sp., Eleutherodactylus cf. juipoca, L. fuscus, and Proceratophrys sp., that are part of species-rich frog communities of the Brazilian Cerrado. Specifically, we investigated age and sex-related, as well as individual-level, diet variation. We measured individual-level diet variation with the IS index of individual specialization, which is a measure of the degree of overlap between individual niches and the population niches. We found no ontogenetic shifts or sex-related differences in the types of prey consumed. However, we found evidence of individual-level diet variation in the four studied species (IS ~ 0.2–0.5). There was a negative correlation between IS and the population niche width (r = –0.980; P < 0.0001), indicating that interindividual diet variation is more pronounced in more generalized populations. This pattern suggests that individual niche widths remain constrained even when population niche breadth is wide, consistent with the presence of functional trade-offs. We found no evidence that these trade-offs arise from morphology, since there was no diet–morphology correlation. We hypothesize that trade-offs have a behavioral or physiological basis, which needs further investigation. This is the first documented case of individual-level diet variation in a diverse tropical community, indicating that this phenomenon is not restricted to competitive release-driven niche expansion in temperate, depauperate communities.