Linking pollen limitation and seed dispersal effectiveness.

Seed production and dispersal are crucial ecological processes impacting plant demography, species distributions and community assembly. Plant-animal interactions commonly mediate both seed production and seed dispersal, but current research often examines pollination and seed dispersal separately, which hinders our understanding of how pollination services affect downstream dispersal services. To fill this gap, we propose a conceptual framework exploring how pollen limitation can impact the effectiveness of seed dispersal for endozoochorous and myrmecochorous plant species. We summarize the quantitative and qualitative effects of pollen limitation on plant reproduction and use Optimal Foraging Theory to predict its impact on the foraging behaviour of seed dispersers. In doing so, we offer a new framework that poses numerous hypotheses and empirical tests to investigate links between pollen limitation and seed dispersal effectiveness and, consequently, post-dispersal ecological processes occurring at different levels of biological organization. Finally, considering the importance of pollination and seed dispersal outcomes to plant eco-evolutionary dynamics, we discussed the implications of our framework for future studies exploring the demographic and evolutionary impacts of pollen limitation for animal-dispersed plants.

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.

Diacamma ants adjust liquid foraging strategies in response to biophysical constraints.

Ant foragers provide food to the rest of the colony, often requiring transport over long distances. Foraging for liquid is challenging because it is difficult to transport and share. Many social insects store liquids inside the crop to transport them to the nest, and then regurgitate to distribute to nest-mates through a behaviour called trophallaxis. Some ants instead transport fluids with a riskier behaviour called pseudotrophallaxis-holding a drop of liquid between the mandibles through surface tension. Ants share this droplet with nest-mates without ingestion or regurgitation. We hypothesised that ants optimize their liquid-collection approach depending on viscosity. Using an ant that employs both trophallaxis and pseudotrophallaxis, we investigated the conditions where each liquid-collection behaviour is favoured by measuring biophysical properties, collection time and reaction to food quality for typical and viscosity-altered sucrose solutions. We found that ants collected more liquid per unit time by mandibular grabbing than by drinking. At high viscosities ants switched liquid collection method to mandibular grabbing in response to viscosity and not to sweetness. Our results demonstrate that ants change transport and sharing methods according to viscosity-a natural proxy for sugar concentration-thus increasing the mass of sugar returned to the nest per trip.

Temporal change in floral availability leads to periods of resource limitation and affects diet specificity in a generalist pollinator.

Generalist species are core components of ecological networks and crucial for the maintenance of biodiversity. Generalist species and networks are expected to be more resilient, and therefore understanding the dynamics of specialization and generalization in ecological networks is a key focus in a time of rapid global change. Whilst diet generalization is frequently studied, our understanding of how it changes over time is limited. Here we explore temporal variation in diet specificity in the honeybee (Apis mellifera), using pollen DNA metabarcoding of honey samples, through the foraging season, over two years. We find that, overall, honeybees are generalists that visit a wide range of plants, but there is temporal variation in the degree of specialization. Temporal specialization of honeybee colonies corresponds to periods of resource limitation, identified as a lack of honey stores. Honeybees experience a lack of preferred resources in June when switching from flowering trees in spring to shrubs and herbs in summer. Investigating temporal patterns in specialization can identify periods of resource limitation that may lead to species and network vulnerability. Diet specificity must therefore be explored at different temporal scales in order to fully understand species and network stability in the face of ecological change.

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.

Early-life diet specificity is associated with long-lasting differences in apparent survival in a generalist predator.

Early-life conditions can have long-term fitness consequences. However, it is still unclear what optimal rearing conditions are, especially for long-lived carnivores. A more diverse diet ('balanced diet') might optimize nutrient availability and allow young to make experiences with a larger diversity of prey, whereas a narrow diet breadth ('specialized diet') might result in overall higher energy net gain. A diet that is dominated by a specific prey type (i.e. fish, 'prey type hypothesis') might be beneficial or detrimental, depending for example, on its toxicity or contaminant load. Generalist predators such as the white-tailed eagle Haliaeetus albicilla provide an interesting possibility to examine the relationship between early life diet and long-term offspring survival. In the Åland Islands, an archipelago in the Baltic Sea, white-tailed eagles live in various coastal habitats and feed on highly variable proportions of birds and fish. We use data from 21,116 prey individuals that were collected from 120 territories during the annual surveys, to examine how early-life diet is associated with apparent annual survival of 574 ringed and molecular-sexed eaglets. We supplement this analysis by assessing the relationships between diet, reproductive performance and nestling physical condition, to consider whether they are confounding with possible long-term associations. We find that early-life diet is associated with long-term fitness: Nestlings that are fed a diverse diet are in lower physical condition but have higher survival rates. Eagles that are fed more fish as nestlings have lower survival as breeding-age adults, but territories associated with fish-rich diets have higher breeding success. Our results show that young carnivores benefit from a high diversity of prey in their natal territory, either through a nutritional or learning benefit, explaining the higher survival rates. The strong relationship between early-life diet and adult survival suggests that early life shapes adult foraging decisions and that eating fish is associated with high costs. This could be due to high levels of contaminants or high competition for fish-rich territories. Long-lasting consequences of early-life diet are likely not only limited to individual-level consequences but have the potential to drive eco-evolutionary dynamics in this population.

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.

Biological Intelligence Inspired Trajectory Design for Energy Harvesting UAV Networks.

In this paper, the problem of trajectory design for energy harvesting unmanned aerial vehicles (UAVs) is studied. In the considered model, the UAV acts as a moving base station to serve the ground users, while collecting energy from the charging stations located at the center of a user group. For this purpose, the UAV must be examined and repaired regularly. In consequence, it is necessary to optimize the trajectory design of the UAV while jointly considering the maintenance costs, the reward of serving users, the energy management, and the user service time. To capture the relationship among these factors, we first model the completion of service and the harvested energy as the reward, and the energy consumption during the deployment as the cost. Then, the deployment profitability is defined as the ratio of the reward to the cost of the UAV trajectory. Based on this definition, the trajectory design problem is formulated as an optimization problem whose goal is to maximize the deployment profitability of the UAV. To solve this problem, a foraging-based algorithm is proposed to find the optimal trajectory so as to maximize the deployment profitability and minimize the average user service time. The proposed algorithm can find the optimal trajectory for the UAV with low time complexity at the level of polynomial. Fundamental analysis shows that the proposed algorithm achieves the maximal deployment profitability. Simulation results show that, compared to Q-learning algorithm, the proposed algorithm effectively reduces the operation time and the average user service time while achieving the maximal deployment profitability.

Using Gaze for Behavioural Biometrics.

A principled approach to the analysis of eye movements for behavioural biometrics is laid down. The approach grounds in foraging theory, which provides a sound basis to capture the uniqueness of individual eye movement behaviour. We propose a composite Ornstein-Uhlenbeck process for quantifying the exploration/exploitation signature characterising the foraging eye behaviour. The relevant parameters of the composite model, inferred from eye-tracking data via Bayesian analysis, are shown to yield a suitable feature set for biometric identification; the latter is eventually accomplished via a classical classification technique. A proof of concept of the method is provided by measuring its identification performance on a publicly available dataset. Data and code for reproducing the analyses are made available. Overall, we argue that the approach offers a fresh view on either the analyses of eye-tracking data and prospective applications in this field.

Dietary abundance distributions: Dominance and diversity in vertebrate diets.

Diet composition is among the most important yet least understood dimensions of animal ecology. Inspired by the study of species abundance distributions (SADs), we tested for generalities in the structure of vertebrate diets by characterising them as dietary abundance distributions (DADs). We compiled data on 1167 population-level diets, representing >500 species from six vertebrate classes, spanning all continents and oceans. DADs near-universally (92.5%) followed a hollow-curve shape, with scant support for other plausible rank-abundance-distribution shapes. This strong generality is inherently related to, yet incompletely explained by, the SADs of available food taxa. By quantifying dietary generalisation as the half-saturation point of the cumulative distribution of dietary abundance (sp50, minimum number of foods required to account for 50% of diet), we found that vertebrate populations are surprisingly specialised: in most populations, fewer than three foods accounted for at least half the diet. Variation in sp50 was strongly associated with consumer type, with carnivores being more specialised than herbivores or omnivores. Other methodological (sampling method and effort, taxonomic resolution), biological (body mass, frugivory) and biogeographic (latitude) factors influenced sp50 to varying degrees. Future challenges include identifying the mechanisms underpinning the hollow-curve DAD, its generality beyond vertebrates, and the biological determinants of dietary generalisation.

Investigating foundations for hominin fire exploitation: Savanna-dwelling chimpanzees (Pan troglodytes verus) in fire-altered landscapes.

Humans' extensive use of fire is one behavior that sets us apart from all other animals. However, our ancestors' reliance on controlled forms of fire-i.e., for cooking-was likely preceded by a long familiarity with fire beginning with passive exploitation of naturally burned landscapes and followed by intermediate steps including active ecological modification via intentional burning. Here we explore our pyrophilic beginnings using observational data from savanna-dwelling chimpanzees. These data highlight the extent to which anthropogenic burning impacts the behavior and ecology of sympatric primates and provides an opportunity to study the ways in which apes living in a fire-altered world exploit opportunities presented by burning. Using monthly burn scar data and daily range use data we quantify the impact of burning episodes on chimpanzee habitat. Over the course of one dry season, approximately 74% of the total estimated range of the Fongoli community of savanna-dwelling chimpanzees (Pan troglodytes verus) was impacted by fire. We combine fire occurrences with behavioral data to test for relationships between burning and rate of encounter with food items and duration of subsequent patch residence time. Results show more frequent encounters and shorter patch residence times in burned areas. These data can be leveraged as a frame of reference for conceptualizing our extinct relatives' behavior around fire.

Individual specialization and generalization in predator-prey dynamics: The determinant role of predation efficiency and prey reproductive rates.

Individual specialization and generalization refer to the breadth of prey types consumed by predators among all available prey. The ecological factors mechanistically determining individual differences and the coexistence of foraging strategies remain to be clarified. Formal quantitative models can elucidate the complex nonlinear mechanisms underlying predator-prey interactions. In this study, we built a dynamical model with multiple prey, with different nutritional values and reproductive rates, which are consumed by specialist and generalist predators. We then analyzed the viability of individual foraging strategies in all possible scenarios. Relative prey fertility and relative efficiency of predation, rather than prey nutritional value, determined the success of specialists and generalists. Less reproducing prey and the specialists relying on them face the highest danger of extinction, while generalists cannot thrive where specialists are sufficiently efficient in relation to the number of available prey. Our study provides new perspectives for empirical studies on individual specialization.

Human spatial memory is biased towards high-calorie foods: a cross-cultural online experiment.

BACKGROUND: Human memory appears to prioritise locations of high-calorie foods, likely as an adaptation for foraging within fluctuating ancestral food environments. Importantly, this "high-calorie bias" in human spatial memory seems to yield consequences for individual eating behaviour in modern food-abundant settings. However, as studies have mainly been conducted in European (Dutch) populations to date, we investigated whether the existence of the cognitive bias can be reasonably generalised across countries that vary on culturally-relevant domains, such as that of the USA and Japan. Furthermore, we investigated whether sociodemographic factors moderate the expression of the high-calorie spatial memory bias in different populations. METHODS: In a cross-cultural online experiment, we measured the food location memory of diverse participants from the USA (N = 72; 44.4% Male; 54 ± 15.99 years) and Japan (N = 74; 56.8% Male; 50.85 ± 17.32 years), using a validated computer-based spatial memory task with standardised images of high-calorie and low-calorie foods. To directly compare the magnitude of the high-calorie spatial memory bias in a broader cultural scope, we also included data from a previous online experiment that identically tested the food spatial memory of a Dutch sample (N = 405; 56.7% Male; 47.57 ± 17.48 years). RESULTS: In the US sample, individuals more accurately recalled (i.e. had lower pointing errors for) locations of high-calorie foods versus that of low-calorie alternatives (Mean difference = -99.23 pixels, 95% CI = [-197.19, -1.28]) - regardless of one's hedonic preferences, familiarity with foods, and encoding times. Likewise, individuals in the Japanese sample displayed an enhanced memory for locations of high-calorie (savoury-tasting) foods (Mean difference = -40.41 pixels, 95% CI = [-76.14, -4.68]), while controlling for the same set of potential confounders. The magnitude of the high-calorie bias in spatial memory was similar across populations (i.e. the USA, Japan, and the Netherlands), as well as across diverse sociodemographic groups within a population. CONCLUSIONS: Our results demonstrate that the high-calorie bias in spatial memory transcends sociocultural boundaries. Since the cognitive bias may negatively impact on our dietary decisions, it would be wise to invest in strategies that intervene on our seemingly universal ability to efficiently locate calorie-rich foods.

Energy-based step selection analysis: Modelling the energetic drivers of animal movement and habitat use.

The energetic gains from foraging and costs of movement are expected to be key drivers of animal decision-making, as their balance is a large determinant of body condition and survival. This fundamental perspective is often missing from habitat selection studies, which mainly describe correlations between space use and environmental features, rather than the mechanisms behind these correlations. To address this gap, we present a novel parameterisation of step selection functions (SSFs), that we term the energy selection function (ESF). In this model, the likelihood of an animal selecting a movement step depends directly on the corresponding energetic gains and costs, and we can therefore assess how moving animals choose habitat based on energetic considerations. The ESF retains the mathematical convenience and practicality of other SSFs and can be quickly fitted using standard software. In this article, we outline a workflow, from data gathering to statistical analysis, and use a case study of polar bears Ursus maritimus to demonstrate application of the model. We explain how defining gains and costs at the scale of the movement step allows us to include information about resource distribution, landscape resistance and movement patterns. We further demonstrate this process with a case study of polar bears and show how the parameters can be interpreted in terms of selection for energetic gains and against energetic costs. The ESF is a flexible framework that combines the energetic consequences of both movement and resource selection, thus incorporating a key mechanism into habitat selection analysis. Further, because it is based on familiar habitat selection models, the ESF is widely applicable to any study system where energetic gains and costs can be derived, and has immense potential for methodological extensions.

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.

Feather pecking and foraging uncorrelated - the redirection hypothesis revisited.

1. Feather pecking (FP) is said to be a redirection of food-related foraging pecks to feathers. The effects of three foraging enrichments on the pecking behaviours of layer pullets in pens and as hens in cages (Gallus gallus domesticus) were studied.2. Treatments included no added enrichment (Control), addition of a mix of wheat and sorghum grains (Grain), a mix of wheat and sorghum grains with lucerne and oaten chaff (Chaff), and lucerne hay (Hay). In pens, Hay was provided in racks, while Grain and Chaff were scattered on the floor. In cages, all treatments were provided in racks. It was predicted that enrichments that encouraged more pecking activity would be most successful at reducing FP.3. Overall pecking activity was similar between all treatments. The Chaff groups tended to FP less than Grain groups, and Hay groups had the least FP. In cages, both Chaff and Hay groups had significantly less FP than either the Grain or Control groups. FP did not reflect stimulus peck rates. Although Chaff groups pecked at their rack significantly less than either Grain or Hay groups, their FP was on a par with the birds in the Hay groups. Feather quality followed these trends.4. The fact that these results were inconsistent with the original hypothesis led to a proposal for a modified hypothesis. Rather than the appetitive component of food searching motivation being redirected to feathers, it is the appetitive component of exploratory motivation, and the ability to reach the goal to update information about the environment is what feeds back to the motivation which leads to FP. This modified hypothesis is consistent with the results of this and many other studies and with modern concepts of motivation, foraging, exploration and food selection.

Using niche construction theory to generate testable foraging hypotheses at Liang Bua.

Niche construction theory (NCT) has emerged as a promising theoretical tool for interpreting zooarchaeological material. However, its juxtaposition against more established frameworks like optimal foraging theory (OFT) has raised important criticism around the testability of NCT for interpreting hominin foraging behavior. Here, we present an optimization foraging model with NCT features designed to consider the destructive realities of the archaeological record after providing a brief review of OFT and NCT. Our model was designed to consider a foragers decision to exploit an environment given predation risk, mortality, and payoff ratios between different ecologies, like more-open or more-forested environments. We then discuss how the model can be used with zooarchaeological data for inferring environmental exploitation by a primitive hominin, Homo floresiensis, from the island of Flores in Southeast Asia. Our example demonstrates that NCT can be used in combination with OFT principles to generate testable foraging hypotheses suitable for zooarchaeological research.

Human behavioral ecology and niche construction.

We examine the relationship between niche construction theory (NCT) and human behavioral ecology (HBE), two branches of evolutionary science that are important sources of theory in archeology. We distinguish between formal models of niche construction as an evolutionary process, and uses of niche construction to refer to a kind of human behavior. Formal models from NCT examine how environmental modification can change the selection pressures that organisms face. In contrast, formal models from HBE predict behavior assuming people behave adaptively in their local setting, and can be used to predict when and why people engage in niche construction. We emphasize that HBE as a field is much broader than foraging theory and can incorporate social and cultural influences on decision-making. We demonstrate how these approaches can be formally incorporated in a multi-inheritance framework for evolutionary research, and argue that archeologists can best contribute to evolutionary theory by building and testing models that flexibly incorporate HBE and NCT elements.

The Ecology of Individual Differences Empirically Applied to Space-Use and Movement Tactics.

Movement provides a link between individual behavioral ecology and the spatial and temporal variation in an individual's landscape. Individual variation in movement traits is an important axis of animal personality, particularly in the context of foraging ecology. We tested whether individual caribou (Rangifer tarandus) displayed plasticity in movement and space-use behavior across a gradient of resource aggregation. We quantified first-passage time and range-use ratio as proxies for movement-related foraging behavior and examined how these traits varied at the individual level across a foraging resource gradient. Our results suggest that individuals adjusted first-passage time but not range-use ratio to maximize access to high-quality foraging resources. First-passage time was repeatable, and intercepts for first-passage time and range-use ratio were negatively correlated. Individuals matched first-passage time but not range-use ratio to the expectations of our patch-use model that maximized access to foraging resources, a result that suggests that individuals acclimated their movement patterns to accommodate both intra- and interannual variation in foraging resources on the landscape. Collectively, we highlight repeatable movement and space-use tactics and provide insight into how individual plasticity in movement interacts with landscape processes to affect the distribution of behavioral phenotypes and potentially fitness and population dynamics.

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.