Niche Variation Hypothesis Research Articles

The adaptive value of density-dependent habitat specialization and social network centrality

Density dependence is a fundamental ecological process. In particular, animal habitat selection and social behavior often affect fitness in a density-dependent manner. The Ideal Free Distribution (IFD) and niche variation hypothesis (NVH) present distinct predictions associated with Optimal Foraging Theory about how the effect of habitat selection on fitness varies with population density. Using caribou (Rangifer tarandus) in Canada as a model system, we test competing hypotheses about how habitat specialization, social behavior, and annual reproductive success (co)vary across a population density gradient. Within a behavioral reaction norm framework, we estimate repeatability, behavioral plasticity, and covariance among social behavior and habitat selection to investigate the adaptive value of sociality and habitat selection. In support of NVH, but not the IFD, we find that at high density habitat specialists had higher annual reproductive success than generalists, but were also less social than generalists, suggesting the possibility that specialists were less social to avoid competition. Our study supports niche variation as a mechanism for density-dependent habitat specialization.

Variation in dental morphology and dietary breadth in primates and their kin

Abstract Sympatric animals may consume diets of differing breadths as a strategy to reduce competition. Studying patterns of dietary breadth in extinct taxa is difficult because available data are generally limited to morphology. Dental topographic analysis (DTA) is useful for comparing occlusal morphology and allows for examination of dietary adaptations in extinct taxa. What remains unknown is how dental morphology, quantified using DTA, covaries with dietary breadth. The niche variation hypothesis (NVH) posits that taxa with broader ecological niches will be characterized by greater variability in morphology relative to specialized taxa. Therefore, we predict that taxa with greater dietary breadth will have more varied dental morphology compared to specialists as a result of the molar morphology of specialists being under greater genetic control relative to generalists, with specialists requiring teeth specially adapted to efficiently process a smaller range of food sources. We measured curvature, complexity, and relief of the M2 of 3 pairs of closely related euarchontan mammals (primates and treeshrews), with each taxon within a pair categorized as a generalist or specialist. Our results indicate that generalists do not consistently show greater variability in dental morphology compared to specialists among primates, but that atelids and treeshrews do generally follow the predictions of the NVH, with the caveat that our treeshrew sample is small. This suggests that while dietary specialists may be under greater genetic constraint with respect to their molar topography, a link between dietary breadth and dental form is not clear. Our study demonstrates that variation in dental topography does not necessarily reflect dietary breadth and highlights the fact that it is difficult to categorize even the most specialized primates (i.e. bamboo lemurs) as “dietary specialists.”

Evolution of intraspecific floral variation in a generalist-specialist pollination system.

Intraspecific processes impact macroevolutionary patterns through individual variation, selection, and ecological specialisation. According to the niche variation hypothesis, the broader ecological niche of gen- eralist species results in an increased morphological variation among individuals, either because they are constituted of diversified specialised individuals each exploiting a fraction of the species' niche, or because they are constituted of true generalist individuals that experience relaxed selection. To test this hypoth- esis, we surveyed the individual floral morphology of species of Antillean Gesneriaceae, a group that has transitioned between specialisation for hummingbird pollination and generalisation multiple times throughout its evolutionary history. We characterised the profiles of corollas using geometric morpho- metrics and compared the intraspecific shape variance of specialists and generalists in a phylogenetic context. We used three approaches that differently accounted for the high dimensionality of morphologi- cal traits, the ancestral reconstruction of pollination syndromes over time, and the error associated with the estimation of the intraspecific variance. Our findings provide partial support for the niche variation hypothesis. If considering the whole shape in the analysis corroborated this idea, decomposing the shape into principal components indicated that not all aspects of the corolla exhibit the same pattern of vari- ation. Specifically, pollination generalists tend to display greater intraspecific variation than specialists in terms of tubularity, but not of curvature. Accounting for the error in the variance estimation also reduced the support for the hypothesis, suggesting that larger sample sizes may be required to reach stronger conclusions. This study emphasises the reciprocal influence between plants and their pollinators on floral morphology at different biodiversity scales, and suggests that ecological strategies of species can affect patterns of morphological variation at macroevolutionary scales.

DNA metabarcoding, diversity partitioning and null models reveal mechanisms of seasonal trophic specialization in a Mediterranean warbler.

Optimal Foraging Theory (OFT) predicts that a population's trophic niche expansion should occur in periods of food scarcity as individuals begin to opportunistically exploit sub-optimal food items. However, the Niche Variation Hypothesis (NVH) posits that niche widening may result from increased among-individual differentiation due to food partitioning to avoid competition. We tested these hypotheses through a DNA metabarcoding study of the Sardinian Warbler (Curruca melanocephala) diet over a year. We used null models and the decomposition of beta diversity on among-individual dietary differentiation to infer the mechanisms driving the population's niche variation. Warblers fed frequently on berries, with a peak in late summer and, to a lesser extent, in autumn. Their diet also included a wide range of arthropods, with their prevalence varying among seasons. Consistent with OFT, the population's niche width was narrower in spring/summer when the population was strongly specialized in berries. In winter, the population's niche expanded, possibly reflecting seasonal declines in food abundance. As predicted by NVH, among-individual differentiation tended to be higher in winter, but this was mainly due to increased differences in dietary richness rather than to the partitioning of resources. Overall, our results suggest that within-individual niche does not increase in lean periods, and instead, individuals adopt either a more opportunistic or more specialized foraging strategy. Increased competition in periods of scarcity may help explain such patterns, but instead of showing increased food partitioning as expected from NVH, it may reflect OFT mechanisms on individuals with differential competitive ability to access better food resources.

Deer dietary responses to wildfire: Optimal foraging, individual specialization, or opportunism?

Increasing impacts of wildfire on arid regions of the world fuelled by climate change highlight the need to better understand how natural communities respond to fire. We took advantage of a large (1660-km2 ) wildfire that erupted in northern California during an in-progress study of black-tailed deer (Odocoileus hemionus columbianus) to investigate deer use of and diets within burned and unburned habitats before and after the fire. We compared deer diet breadth to predictions of optimal foraging theory, the niche variation hypothesis, and opportunistic (i.e., generalist) foraging expectations under the assumption that overall availability and diversity of forage in burned areas declined immediately after the fire and increased as the plant community recovered in the next 3 years after the fire. We used faecal pellet counts to document space use and metabarcoding to study diet during pre-fire, post-fire, and recovery periods. Pellet counts supported predictions that deer increased use of unburned sites and reduced use of burn sites after the fire and began to return to burned sites in subsequent sampling years. Diet diversity did not differ significantly between control and burn sites before the fire, but was lower in burn than control sites post-fire (p < .001), when and where diet was dominated by oak (Quercus spp). In contrast, during subsequent years, diet diversity was higher (including more herbaceous plants) in burn than control sites (p < .05). In contrast to predictions of optimal foraging and niche variation hypotheses, individual deer foraged as generalists for which changes in dietary niche breadth paralleled fire-induced changes in diversity of the plant community.

Individual contribution to niche expansion in amphibians: a test of the niche variation hypothesis

The niche variation hypothesis (NVH) suggests that populations released from competition are able to expand their realized ecological niche. This increase in total niche width (TNW) can arise by 1) increasing heterogeneity among individuals' niche through individual specialization, i.e. inter‐individual variation in resource use, (IS) occurring when different individuals of a population use a subset of the population's resource pool, or 2) by expanding the niche of all individuals within the population. Although several morphological and phenotypical studies have confuted the NVH, there is generally strong support for this hypothesis when behavioural or ecological traits are considered, and in particular when the trophic niche is measured at the individual level. In this study our primary aim is to corroborate the NVH, providing robust evidence for a significant and positive relationship between population TNW and IS in amphibian populations, at a global scale.

Seasonal variation of population and individual dietary niche in the avivorous bat, Ia io

The variation in niche breadth can affect how species respond to environmental and resource changes. However, there is still no clear understanding of how seasonal variability in food resources impacts the variation of individual dietary diversity, thereby affecting the dynamics of a population's dietary niche breadth. Optimal foraging theory (OFT) and the niche variation hypothesis (NVH) predict that when food resources are limited, the population niche breadth will widen or narrow due to increased within-individual dietary diversity and individual specialization or reduced within-individual dietary diversity, respectively. Here, we used DNA metabarcoding to examine the composition and seasonality of diets of the avivorous bat Ia io. Furthermore, we investigated how the dietary niches changed among seasons and how the population niche breadth changed when the availability of insect resources was reduced in autumn. We found that there was differentiation in dietary niches among seasons and a low degree of overlap, and the decrease of insect resource availability and the emergence of ecological opportunities of nocturnal migratory birds might drive dietary niche shifts toward birds in I. io. However, the population's dietary niche breadth did not broaden by increasing the within-individual dietary diversity or individual specialization, but rather became narrower by reducing dietary diversity via predation on bird resources that served as an ecological opportunity when insect resources were scarce in autumn. Our findings were consistent with the predictions of OFT, because birds as prey for bats provided extremely different resources from those of insects in size and nutritional value. Our work highlights the importance of size and quality of prey resources along with other factors (i.e., physiological, behavioral, and life-history traits) in dietary niche variation.

Diet and disturbance: Seaside Sparrow resource use driven by oiling and Hurricane Isaac

The 2010 Deepwater Horizon (DWH) disaster led to extensive oil deposition in Louisiana’s coastal wetlands, followed two years later by Hurricane Isaac. These disturbances led to changes in saltmarsh invertebrate communities, potentially affecting higher-level predators and the saltmarsh food web. Seaside Sparrows (Ammospiza maritima) are ubiquitous, year-round residents of the coastal saltmarsh affected by the DWH spill, where they consume diverse invertebrates and may be considered an indicator species for the ecosystem’s integrity. We used DNA metabarcoding to evaluate prey consumed by Seaside Sparrows to understand how sparrows responded to residual contamination from the DWH oil spill and ecosystem disturbance caused by Hurricane Isaac. To do so, we evaluated metrics of diet (prey richness, diversity, overall diet composition) and resource use (total niche width, individual specialization) from 2011 to 2017 on oiled, unoiled, and reference sites. We found that while diet composition varied across years and site type, Hurricane Isaac had an even greater effect on the richness and diversity of prey consumed. Resource use—as measured by the total niche width of the populations and degree of individual specialization—was most stable on unoiled sites compared to unoiled and reference sites. Finally, we analyzed resource use for each combination of site type and year (i.e.: “2014 oiled sites”), which indicated a strong correlation between individual specialization and total niche width: as total niche width increased, individuals became more specialized, following the predictions of the Niche Variation Hypothesis.

Testing the niche variation hypothesis in pinnipeds

Abstract Individual trophic specialisation (ITS) has many important consequences for ecological and evolutionary processes. An old hypothesis highlighting the relevance of ITS is the niche variation hypothesis (NVH), which proposes that populations composed of trophically specialised individuals have a wider population niche than populations composed of more generalist individuals. Pinnipeds are a good model to test the NVH because they are mostly generalist species, living in dense colonies, and exhibiting variation in trophic niche width among populations. We tested the NVH in pinnipeds using longitudinal isotopic data: published δ13C and δ15N data obtained from sequential sections of whiskers (vibrissae) from individuals belonging to 14 populations. To the best of our knowledge, this is the first study evaluating the NVH by using longitudinal isotopic data. We compiled studies that published raw data on δ13C and δ15N for pinniped whisker segments to estimate the within‐individual and the between‐individuals components of variance, the total niche width (TNW), and ITS. One‐dimensional (i.e. δ13C or δ15N) and multi‐dimensional (i.e. δ13C and δ15N) analyses were used. Also, we tested whether an evolutionary component would affect the degree of ITS among pinniped species with different levels of shared ancestry. Our results indicate that, in line with the NVH, pinniped populations composed of more specialised individuals tend to show wider trophic niches. When analysing each sex separately, the hypothesis is supported for females but not for males. We believe that physiological and behavioural differences between sexes may explain this result. In females, high TNW is mainly related to greater differentiation among individuals, while in males, it is mainly related to high diversity of resources consumed by all individuals.

Population niche width is driven by within-individual niche expansion and individual specialization in introduced brook trout in mountain lakes

The width of a population's resource-use niche is determined by individual diet breadth (“within-individual component”) and the degree of niche partitioning between individuals (“between-individual component”). The balance between these two factors affects ecological stability and evolutionary trajectories, and may shift as ecological opportunity permits broader population niches. Lakes in California’s Sierra Nevada Mountains vary in resource diversity for introduced brook trout (Salvelinus fontinalis) due to elevation, lake morphometry, and watershed features. We compared the relative contributions of within- and between-individual niche components to two measures of the dietary niches of thirteen populations of brook trout: prey taxonomic composition and prey size distribution. For both taxonomic and size diversity of fish diets, population niche width was positively related to both the within- and between-individual components. For taxonomic diversity, the two components increased in parallel, while for size diversity, the between-individual component became more important relative to the within-individual component in populations with the greatest niche widths. Our results support the Niche Variation Hypothesis that populations with broader niches are more heterogeneous among individuals and show that individual niche width and individual specialization can operate in parallel to expand the population niche.

"Jack-of-all-trades" is parthenogenetic.

Sex is evolutionarily more costly than parthenogenesis, evolutionary ecologists therefore wonder why sex is much more frequent than parthenogenesis in the majority of animal lineages. Intriguingly, parthenogenetic individuals and species are as common as or even more common than sexuals in some major and putative ancient animal lineages such as oribatid mites and rotifers. Here, we analyzed oribatid mites (Acari: Oribatida) as a model group because these mites are ancient (early Paleozoic), widely distributed around the globe, and include a high number of parthenogenetic species, which often co‐exist with sexual oribatid mite species. There is evidence that the reproductive mode is phylogenetically conserved in oribatid mites, which makes them an ideal model to test hypotheses on the relationship between reproductive mode and species' ecological strategies. We used oribatid mites to test the frozen niche variation hypothesis; we hypothesized that parthenogenetic oribatid mites occupy narrow specialized ecological niches. We used the geographic range of species as a proxy for specialization as specialized species typically do have narrower geographic ranges than generalistic species. After correcting for phylogenetic signal in reproductive mode and demonstrating that geographic range size has no phylogenetic signal, we found that parthenogenetic lineages have a higher probability to have broader geographic ranges than sexual species arguing against the frozen niche variation hypothesis. Rather, the results suggest that parthenogenetic oribatid mite species are more generalistic than sexual species supporting the general‐purpose genotype hypothesis. The reason why parthenogenetic oribatid mite species are generalists with wide geographic range sizes might be that they are of ancient origin reflecting that they adapted to varying environmental conditions during evolutionary history. Overall, our findings indicate that parthenogenetic oribatid mite species possess a widely adapted general‐purpose genotype and therefore might be viewed as “Jack‐of‐all‐trades.”

Feeding ecology of the endangered Apennine Yellow-bellied Toad (Bombina pachypus) in central Italy

Anurans are generalist opportunistic consumers, but many species show some degree of diet specialization. Within populations, individuals may specialize in specific dietary resources (individual specialization). Indeed, generalist populations may be composed of individuals using the whole range of available resources or of diverse individuals exploiting specific food items. Based on the niche variation hypothesis (NHV), an increase in population niche width may be related to a higher incidence of individual specialization. We analyzed the feeding ecology of the endangered Apennine Yellow-bellied Toad ( Bombina pachypus (Bonaparte, 1838)) in central Italy. This species showed a generalist feeding strategy, wide trophic niche, high individual generalization, and a predominant use of terrestrial resources. The study species inhabits seasonal habitats determining variation in food availability and diversity, and this may have driven toads to be unselective and consume almost any encountered palatable item. The absence of sexual size dimorphism may explain the absence of trophic differentiation between sexes. The analysis of individual feeding traits revealed a week's degree of individual specialization positively correlated to niche width, partly supporting the NHV. The obtained information can be used for the implementation of effective conservation strategies of locally imperilled species, especially those breeding in temporary ponds.

Isotopic niche partitioning and individual specialization in an Arctic raptor guild

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.

Dentin hardness differences across various mammalian taxa.

Differences in dentin microstructure have been used as a tool for dietary reconstruction; however, the extent that diet is associated with this aspect of dental morphology has yet to be empirically tested. We conducted microhardness tests of mammalian dentin sections, hypothesizing that species with adaptations to particularly hard diets would have softer dentin, owing to a higher proportion of soft intertubular dentin. Species adapted to abrasive diets, in contrast, should have harder dentin, resulting from a higher proportion of hypermineralized peritubular dentin. We examined molar dentin hardness in ten mammalian taxa with durophagous diets, abrasive diets, and a comparative "control" group of mechanical generalists. Samples included six primate taxa and four non-primate species representing various dietary regimes. Our results reveal significant variation among taxa in overall hardness, but the data do not distinguish between hard and abrasive diets. Several taxa with generalized (i.e., mechanically diverse) diets resemble each other in exhibiting large variance in hardness measurements and comparably soft dentin. The high variation in these species appears to be either a functional signal supporting the niche variation hypothesis or indicate the absence of sustained unidirectional selective pressure. A possible phylogenetic signal of dentin hardness in the data also holds promise for future systematic investigations.

Relationships Among Trophic Niche Width, Morphological Variation, and Genetic Diversity of Hemiculter leucisculus in China

According to the niche variation hypothesis (NVH), the populations with wider niches are phenotypically more variable than those with narrow niches. Giller expanded the NVH, suggesting that the niche width, morphological variation, and genetic diversity are all positively correlated. However, the hypothesis has been a subject of debate and discussion. In the present study, the NVH was tested by analyzing the relationships among trophic niche width, morphological variation, and genetic diversity of Hemiculter leucisculus, a widespread cyprinid fish. The fish samples were collected from six sites across Haihe, the Yellow, and the Yangtze River basins in China. The relationships among trophic niche width, morphological variation, and genetic diversity were analyzed using Pearson correlation at the inter-population level. Our analysis indicated that trophic niche width is significantly positively correlated with morphological variation, which corroborates the NVH. Morphological variation was significantly correlated to genetic diversity. However, no relationship was observed between trophic niche width and genetic diversity. We inferred that the dietary niche of H. leucisculus might change due to the plastic response toward environmental changes rather than due to the genetic variation. We also suggest that the effects of environment and heredity on the niche of the freshwater fish should be quantified separately in further studies.

Adaptive foraging in the Anthropocene: can individual diet specialization compensate for biotic homogenization?

Biotic homogenization is a ubiquitous consequence of human disturbance. Through a mix of local extinctions and invasions, diverse communities of specialists are often replaced by or inundated with generalist species, resulting in uncertain consequences for ecological functions. While concern about biotic homogenization is growing, intraspecific variation and individual diet specialization (IS) have also emerged as key drivers of ecological functions. The niche variation hypothesis predicts that when a population is released from interspecific competition (ie “ecological release”), intraspecific competition will promote resource niche expansion and IS. It then follows that if biotic homogenization reduces taxonomic diversity and interspecific competition, intraspecific competition within the remaining taxa should expand population resource niches and increase the prevalence of IS. Here, we integrate the rapidly growing fields of biotic homogenization and intraspecific variation and assess the potential for generalist species to compensate for multiple forms of biotic homogenization via adaptive foraging, resource niche expansion, and IS.

Niche-trait relationships at individual and population level in three co-occurring passerine species.

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

How to Become a Generalist Species? Individual Niche Variation Across Habitat Transformation Gradients

Species in transformed habitats, frequently labeled as environmental generalists, tend to show broader niches than species in natural habitats. However, how population niche expansion translates into changes in the niches of individual organisms remains unclear, particularly in the context of habitat transformation. Niche expansion could be a product of individuals having broader niches, greater distances among individuals’ niches, or a combination of both processes. This would challenge the traditional conceptions on niche dynamics, which emphasize the role played by individual specialization (IS). Here, using stable isotopes, we computed total niche width (TNW), its within- and between-individual components (WIC and BIC), and IS (the ratio WIC/TNW), in 13 populations of 6 bird species and 8 populations of 3 frog species in natural and transformed habitats. We confirmed that species had broader niche width in transformed than in natural habitats, yet population niche expansion across habitats was mainly a product of increased distance between individuals. Within each habitat type, increases in TNW were linked to increases in WIC for all habitat types, while relationships between TNW and BIC were found in transformed but not in natural habitats. Hence, both increased individual niche width and increased distance among individuals were apparent within habitats, particularly in transformed ones, where increases in WIC dominated. Neither across or within habitats was niche expansion associated with increasing IS. Therefore, our results overturn traditional conceptions associated with the niche variation hypothesis and illustrate that niche expansion is not invariably associated with increased IS, because the distance between individual’s niches (BIC) can increase, as well as the breadth of those niches (WIC).

Bioenergy landscapes drive trophic shifts in generalist ants

Changes in trophic niche-the pathways through which an organism obtains energy and nutrients-are a fundamental way in which organisms respond to environmental conditions. But the capacity for species to alter their trophic niches in response to global change, and the ways they do so when able, remain largely unknown. Here we examine food webs in three long-term and large-scale experiments to test how resource availability and nutritional requirements interact to determine an organism's trophic niche in the context of one of the largest global trends in land use-the rise in bioenergy production. We use carbon and nitrogen stable isotope analyses to characterize arthropod food webs across three biofuel crops representing a gradient in plant resource richness (corn monocultures, fields dominated by native switchgrass and restored prairie), and to quantify changes in the trophic niche of a widespread generalist ant species across habitats. In doing so, we measure the effects of basal resource richness on food chain length, niche breadth and trophic position. We frame our results in the context of two hypotheses that explain variation in trophic niche-the niche variation hypothesis which emphasizes the importance of resource diversity and ecological opportunity, and the optimal diet hypothesis which emphasizes dietary constraints and the availability of optimal resources. Increasing plant richness lengthened food chains by 10%-20% compared to monocultures. Niche breadths of generalist ants did not vary with resource richness, suggesting they were limited by optimal diet requirements and constraints rather than by ecological opportunity. The ants instead responded to changes in plant richness by shifting their estimated trophic position. In resource-poor monocultures, the ants were top predators, sharing a trophic position with predatory spiders. In resource-rich environments, in contrast, the ants were omnivores, relying on a mix of animal prey and plant-based resources. In addition to highlighting novel ecosystem impacts of alternate bioenergy landscapes, our results suggest that niche breadth and trophic diversification depend more on the presence of optimal resources than on ecological opportunity alone.

A test of the Niche Variation Hypothesis in a ruminant herbivore.

Despite the shared prediction that the width of a population's dietary niche expands as food becomes limiting, the Niche Variation Hypothesis (NVH) and Optimal Foraging Theory (OFT) offer contrasting views about how individuals alter diet selection when food is limited. Classical OFT predicts that dietary preferences do not change as food becomes limiting, so individuals expand their diets as they compensate for a lack of preferred foods. In contrast, the NVH predicts that among-individual variation in cognition, physiology or morphology create functional trade-offs in foraging efficiency, thereby causing individuals to specialize on different subsets of food as food becomes limiting. To evaluate (a) the predictions of the NVH and OFT and (b) evidence for physiological and cognitive-based functional trade-offs, we used DNA microsatellites and metabarcoding to quantify the diet, microbiome and genetic relatedness (a proxy for social learning) of 218 moose Alces alces across six populations that varied in their degree of food limitation. Consistent with both the NVH and OFT, dietary niche breadth increased with food limitation. Increased diet breadth of individuals-rather than increased diet specialization-was strongly correlated with both food limitation and dietary niche breadth of populations, indicating that moose foraged in accordance with OFT. Diets were not constrained by inheritance of the microbiome or inheritance of diet selection, offering support for the little-tested hypothesis that functional trade-offs in food use (or lack thereof) determine whether populations adhere to the predictions of the NVH or OFT. Our results indicate that both the absence of strong functional trade-offs and the digestive physiology of ruminants provide contexts under which populations should forage in accordance with OFT rather than the NVH. Also, because dietary niche width increased with increased food limitation, OFT and the NVH provide theoretical support for the notion that plant-herbivore interaction networks are plastic rather than static, which has important implications for understanding interspecific niche partitioning. Lastly, because population-level dietary niche breadth and calf recruitment are correlated, and because calf recruitment can be a proxy for food limitation, our work demonstrates how diet data can be employed to understand a populations' proximity to carrying capacity.