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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 δ¹³C 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.

Constraints on life-history traits, with their close links to fitness, are widely invoked as limits to niche expansion at most organizational levels. Theoretically, such constraints can maintain individual specialization by preventing adaptation to all niches available, but empirical evidence of them remains elusive for natural populations. This problem may be compounded by a tendency to seek constraints involving multiple traits, neglecting their added potential to manifest in trait expression across environments (i.e., within reaction norms). By replicating genotypes of a colonial marine invertebrate across successional stages in its local community, and taking a holistic approach to the analysis of ensuing reaction norms for fitness, we show the potential for individual specialization to be maintained by genetic constraints associated with these norms, which limit the potential for fitness at one successional stage to improve without loss of fitness at others. Our study provides new insight into the evolutionary maintenance of individual specialization in natural populations and reinforces the importance of reaction norms for studying this phenomenon.

This article is a Commentary on Lofgren et al., 217: 1203–1212 and Martino et al., 217: 1213–1229.

Niche expansion is attained by adaptations in two generalized phenotypical traits—niche position and niche width. This gives room for a wide range of conceptual ways of niche filling. The niche variation hypothesis reduces the range by predicting that expansion occurs by increasing variation in niche position, which has been debated on empirical and theoretical grounds as also other options seem possible. Here, we propose a general theory of niche expansion. We review empirical data and show with an eco-evolutionary model how resource diversity and a trade-off in resource acquisition steer niche evolution consistent with observations. We show that the range can be reduced to a discrete set of two orthogonal ways of niche filling, through (1) strict phenotypical differentiation in niche position or (2) strict individual generalization. When individual generalization is costly, niche expansion undergoes a shift from (2) to (1) at a point where the resource diversity becomes sufficiently large. Otherwise, niche expansion always follows (2), consistent with earlier results. We show that this either–or response can operate at both evolutionary and short-term time scales. This reduces the principles of niche expansion under environmental change to a notion of orthogonality, dictated by resource diversity and a resource-acquisition trade-off.

Many populations of consumers consist of relatively specialized individuals that eat only a subset of the foods consumed by the population at large. Although the ecological significance of individual-level diet variation is recognized, such variation is difficult to document, and its underlying mechanisms are poorly understood. Optimal foraging theory provides a useful framework for predicting how individuals might select different diets, positing that animals balance the “opportunity cost” of stopping to eat an available food item against the cost of searching for something more nutritious; diet composition should be contingent on the distribution of food, and individual foragers should be more selective when they have greater energy reserves to invest in searching for high-quality foods. We tested these predicted mechanisms of individual niche differentiation by quantifying environmental (resource heterogeneity) and organismal (nutritional condition) determinants of diet in a widespread browsing antelope (bushbuck, Tragelaphus sylvaticus) in an African floodplain-savanna ecosystem. We quantified individuals’ realized dietary niches (taxonomic richness and composition) using DNA metabarcoding of fecal samples collected repeatedly from 15 GPS-collared animals (range 6–14 samples per individual, median 12). Bushbuck diets were structured by spatial heterogeneity and constrained by individual condition. We observed significant individual-level partitioning of food plants by bushbuck both within and between two adjacent habitat types (floodplain and woodland). Individuals with home ranges that were closer together and/or had similar vegetation structure (measured using LiDAR) ate more similar diets, supporting the prediction that heterogeneous resource distribution promotes individual differentiation. Individuals in good nutritional condition had significantly narrower diets (fewer plant taxa), searched their home ranges more intensively (intensity-of-use index), and had higher-quality diets prediction that animals with greater endogenous reserves have narrower realized niches because they can invest more time in searching for nutritious foods. Our results support predictions from optimal foraging theory about the energetic basis of individual-level dietary variation and provide a potentially generalizable framework for understanding how individuals’ realized niche width is governed by animal behavior and physiology in heterogeneous landscapes.

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. Intra‐population variability in resource use by consumer species has been recently associated with landscape heterogeneity. By combining data on isotopic niche with a long‐term data set on reproduction, we documented an effect of the heterogeneous breeding environments on the diet and the reproductive output of a top predator, the peregrine falcon. Within a terrestrial/marine gradient of habitat, peregrine falcons predominantly consumed terrestrial prey, but individuals nesting at the marine end produced fewer young than their conspecifics nesting at the terrestrial end.

1. Megafauna assemblages have declined or disappeared throughout much of the world, and many efforts are underway to restore them. Understanding the trophic ecology of such reassembling systems is necessary for predicting recovery dynamics, guiding management, and testing general theory. Yet, there are few studies of recovering large-mammal communities, and fewer still that have characterized food-web structure with high taxonomic resolution. 2. In Gorongosa National Park, large herbivores have rebounded from near-extirpation following the Mozambican Civil War (1977-1992). However, contemporary community structure differs radically from the prewar baseline: medium-sized ungulates now outnumber larger bodied species, and several apex carnivores remain locally extinct. 3. We used DNA metabarcoding to quantify diet composition of Gorongosa's 14 most abundant large-mammal populations. We tested five hypotheses: (i) the most abundant populations exhibit greatest individual-level dietary variability; (ii) these populations also have the greatest total niche width (dietary diversity); (iii) interspecific niche overlap is high, with the diets of less-abundant species nested within those of more-abundant species; (iv) partitioning of forage species is stronger in more structurally heterogeneous habitats; and (v) selectivity for plant taxa converges within guilds and digestive types, but diverges across them. 4. Abundant (and narrow-mouthed) populations exhibited higher among-individual dietary variation, but not necessarily the greatest dietary diversity. Interspecific dietary overlap was high, especially among grazers and in structurally homogenous habitat, whereas niche separation was more pronounced among browsers and in heterogeneous habitat. Patterns of selectivity were similar for ruminants—grazers and browsers alike--but differed between ruminants and non-ruminants. 5. Synthesis. The structure of this recovering food web was consistent with several hypotheses predicated on competition, habitat complexity, and herbivore traits, but it differed from patterns observed in more intact assemblages. We propose that intraspecific competition in the fastest-recovering populations has promoted individual variation and a more nested food web, wherein rare species use subsets of foods eaten by abundant species, and that this scenario is reinforced by weak predation pressure. Future work should test these conjectures and analyse how the taxonomic dietary niche axis studied here interacts with other mechanisms of diet partitioning to affect community reassembly following wildlife declines.

Intra-and inter-specific resource partitioning within predator communities is a fundamental component of trophic ecology, and one proposed mechanism for how populations partition resources is through individual niche variation. The Niche Variation Hypothesis (NVH) predicts that inter-individual trait variation leads to functional trade-offs in foraging efficiency, resulting in populations composed of individual dietary specialists. The degree to which niche specialization persists within a population is plastic and responsive to fluctuating resource availability. We quantified niche overlap and tested the NVH within an Arctic raptor guild, focusing on three species that employ different foraging strategies: golden eagles (generalists); gyrfalcons (facultative specialists); and rough-legged hawks (specialists). Tundra ecosystems exhibit cyclic populations of arvicoline rodents (lemmings and voles), providing a unique system in which to examine predator diet in response to interannual fluctuations in resource availability. Using blood δ¹³C and δ¹⁵N values from 189 raptor nestlings on Alaska’s Seward Peninsula (2014–2019), we calculated isotopic niche width and used Bayesian stable isotope mixing models (BSIMMs) to characterize individual specialization and test the NVH. Nest-level specialization estimated from stable isotopes was strongly correlated with indices of specialization based on camera trap data. We observed a high degree of isotopic niche overlap between the three species and gyrfalcons displayed a positive relationship between individual specialization and population niche width on an interannual basis consistent with the NVH. Our findings suggest plasticity in niche specialization may reduce intra-and inter-specific resource competition under dynamic ecological conditions.

Individual specialization, when individuals exploit only a subset of resources utilized by the population, is a widespread phenomenon. It provides the basis for evolutionary diversification and can impact population and community dynamics. Both phenotypic traits and environmental conditions are predicted to influence individual specialization; however, its adaptive consequences are poorly understood, particularly among large mammalian carnivores that play an important role in shaping ecosystems. We used observations of 2,960 kills made by 49 leopards Panthera pardus in the Sabi Sand Game Reserve, South Africa, to quantify the magnitude of individual dietary specialization in a solitary large carnivore, and to examine the proximate and ultimate drivers of this behaviour. We found evidence of individual specialization in leopard diet, with respect to both the species and size of prey killed. Males tended to be more specialized than females, likely because they could access a wider range of prey due to larger body size. Similarly, individuals that encountered a greater diversity of prey tended to be more specialized. Our results confirmed that ecological opportunity was a key determinant of individual specialization; however, contrary to predictions, per capita resource availability (and by extension, intraspecific competition) did not affect the degree of specialization exhibited by individuals. Surprisingly, dietary specialization appeared to disadvantage male leopards. Specialist males overlapped with fewer resident females, had fewer cubs born on their home ranges and had fewer cubs survive to independence on their home ranges than generalist males. This may have resulted from the high degree of environmental stochasticity experienced during our study, as dietary specialization is expected to advantage individuals more during periods of resource predictability. In summary, we showed that a species usually considered to be a dietary generalist was in fact a heterogeneous collection of specialist and generalist individuals. Individual specialization is typically assumed to be maintained by disruptive and/or fluctuating selection; hence, the somewhat paradoxical coexistence of both in the same population might be explained by a dynamic evolutionary equilibrium that exists between specialists and generalists, in which each benefit under different conditions. Using leopards as a model species, this study provides novel insight into the circumstances under which individual dietary specialization manifests in mammalian carnivore populations. Importantly, it is the first assessment of the fitness consequences of dietary specialization in a large carnivore (photograph credit: James Tyrrell).

Among-individual variation in predator traits is ubiquitous in nature. However, variation among populations in this trait variation has been seldom considered in trophic dynamics. This has left unexplored (a) to what degree does among-individual variation in predator traits regulate prey populations and (b) to what degree do these effects vary spatially. We address these questions by examining how predator among-individual variation in functional traits shapes communities across habitats of varying structural complexity, in field conditions. We manipulated Chinese mantis (Tenodera sinensis) density (six or twelve individuals) and behavioral trait variability (activity level by movement on an open field) in experimental patches of old fields with varying habitat complexity (density of plant material). Then, we quantified their impacts on lower trophic levels, specifically prey (arthropods > 4 mm) and plant biomass. Predator behavioral variability only altered prey biomass in structurally complex plots, and this effect depended on mantis density. In the plots with the highest habitat complexity and mantis density, behaviorally variable groups decreased prey biomass by 40.3%. In complex plots with low mantis densities, low levels of behavioral variability decreased prey biomass by 32.2%. Behavioral variability and low habitat complexity also changed prey community composition, namely by increasing ant biomass by 881%. Our results demonstrate that among-individual trait variation can shape species-rich prey communities. Moreover, these effects depend on both predator density and habitat complexity. Incorporating this important facet of ecological diversity revealed normally unnoticed effects of functional traits on the structure and function of food webs.