Ontogenetic and individual diet variation in amphibian larvae across an environmental gradient

1. Variation among individuals within size or age classes can have profound effects on community dynamics and food‐web structure. We investigated the potential influence of habitat disturbance on intrapopulation niche variation. 2. Amphibians occupy a range of lentic habitats from short‐hydroperiod intermittent ponds to long‐hydroperiod permanent ponds. We quantified ontogenetic diet variation and individual specialisation in wood frog tadpoles (Lithobates sylvaticus) and blue‐spotted salamander larvae (Ambystoma laterale) to investigate the influence of hydroperiod on population niche width across a natural hydroperiod gradient using stable isotope and gut content analyses. In one of the few tests using larval forms, we tested the niche variation hypothesis, which predicts that populations with larger niche widths also have increased individual variation. 3. Our results support the niche variation hypothesis, indicating that more generalised populations exhibit higher within‐individual diet variation. We report gradual changes in the relative importance of diet items, decreased dietary overlap and increased trophic position in L. sylvaticus throughout development. A. laterale became more enriched in δ¹³C and increased in δ¹⁵N throughout its larval period. We did not find a relationship between hydroperiod and niche parameters, indicating that niches are conserved across heterogeneous habitats. In contrast to most documented cases, we estimated low levels of individual specialisation in amphibian larvae. 4. Amphibians are an important link between aquatic and terrestrial ecosystems, whereby diet shifts can influence food‐web structure by altering energy flow pathways and the trophic position of higher consumers, ultimately changing food‐chain length.     

Niche segregation of coexisting Arctic charr (Salvelinus alpinus) and brown trout (Salmo trutta) constrains food web coupling in subarctic lakes

1. Generalist fish species are recognised as important couplers of benthic and pelagic food‐web compartments in lakes. However, interspecific niche segregation and individual specialisation may limit the potential for generalistic feeding behaviour. 2. We studied summer habitat use, stomach contents and stable isotopic compositions of the generalist feeder Arctic charr coexisting with its common resource competitor brown trout in five subarctic lakes in northern Norway to reveal population‐level and individual‐level niche plasticity. 3. Charr and trout showed partial niche segregation in all five lakes. Charr used all habitat types and a wide variety of invertebrate prey including zooplankton, whereas trout fed mainly on insects in the littoral zone. Hence, charr showed a higher potential to promote habitat and food‐web coupling compared to littoral‐dwelling trout. 4. The level of niche segregation between charr and trout and between pelagic‐caught and littoral‐caught charr depended on the prevailing patterns of interspecific and intraspecific resource competition. The two fish species had partially overlapping trophic niches in one lake where charr numerically dominated the fish community, whereas the most segregated niches occurred in lakes where trout were more abundant. 5. In general, pelagic‐caught charr had substantially narrower dietary and isotopic niches and relied less on littoral carbon sources compared to littoral‐caught conspecifics that included generalist as well as specialised benthivorous and planktivorous individuals. Despite the partially specialised planktivorous niche and thus reduced potential of pelagic‐dwelling charr to promote benthic–pelagic coupling, the isotopic compositions of both charr subpopulations suggested a significant reliance on both littoral and pelagic carbon sources in all five study lakes. 6. Our study demonstrates that both interspecific niche segregation between and individual trophic specialisation within generalist fish species can constrain food‐web coupling and alter energy mobilisation to top consumers in subarctic lakes. Nevertheless, pelagic and littoral habitats and food‐web compartments may still be highly integrated due to the potentially plastic foraging behaviour of top consumers.     

The ecological causes of individual specialisation

Many generalist populations are composed of specialised individuals, whose niches are small subsets of the population niche. This 'individual specialisation' is a widespread phenomenon in natural populations, but until recently few studies quantified the magnitude of individual specialisation and how this magnitude varies among populations or contexts. Such quantitative approaches are necessary for us to understand how ecological interactions influence the amount of among-individual variation, and how the amount of variation might affect ecological dynamics. Herein, we review recent studies of individual specialisation, emphasising the novel insights arising from quantitative measures of diet variation. Experimental and comparative studies have confirmed long-standing theoretical expectations that the magnitude of among-individual diet variation depends on the level of intra and interspecific competition, ecological opportunity and predation. In contrast, there is little empirical information as to how individual specialisation affects community dynamics. We discuss some emerging methodological issues as guidelines for researchers studying individual specialisation, and make specific recommendations regarding avenues for future research.     

Individual foraging specialisation in a social mammal: the European badger (Meles meles)

Individual specialisation has been identified in an increasing number of animal species and populations. However, in some groups, such as terrestrial mammals, it is difficult to disentangle individual niche variation from spatial variation in resource availability. In the present study, we investigate individual variation in the foraging niche of the European badger (Meles meles), a social carnivore that lives in a shared group territory, but forages predominantly alone. Using stable isotope analysis, we distinguish the extent to which foraging variation in badgers is determined by social and spatial constraints and by individual differences within groups. We found a tendency for individual badgers within groups to differ markedly and consistently in their isotope values, suggesting that individuals living with access to the same resources occupied distinctive foraging niches. Although sex had a significant effect on isotope values, substantial variation within groups occurred independently of age and sex. Individual differences were consistent over a period of several months and in some instances were highly consistent across the two years of the study, suggesting long-term individual foraging specialisations. Individual specialisation in foraging may, therefore, persist in populations of territorial species not solely as a result of spatial variation in resources, but also arising from individuals selecting differently from the same available resources. Although the exact cause of this behaviour is unknown, we suggest that specialisation may occur due to learning trade-offs which may limit individual niche widths. However, ecological factors at the group level, such as competition, may also influence the degree of specialisation.     

Individual diet specialisation in sparrows is driven by phenotypic plasticity in traits related to trade‐offs in animal performance

Individual diet specialisation (IS) is frequent in many animal taxa and affects population and community dynamics. The niche variation hypothesis (NVH) predicts that broader population niches should exhibit greater IS than populations with narrower niches, and most studies that examine the ecological factors driving IS focus on intraspecific competition. We show that phenotypic plasticity of traits associated with functional trade‐offs is an important, but unrecognised mechanism that promotes and maintains IS. We measured nitrogen isotope (δ¹⁵N) and digestive enzyme plasticity in four populations of sparrows (Zonotrichia capensis) to explore the relationship between IS and digestive plasticity. Our results show that phenotypic plasticity associated with functional trade‐offs is related in a nonlinear fashion with the degree of IS and positively with population niche width. These findings are opposite to the NVH and suggest that among individual differences in diet can be maintained via acclimatisation and not necessarily require a genetic component.     

Searching for the fundamental niche using individual‐based habitat selection modelling across populations

Strictly speaking, fundamental niches are inestimable. Nevertheless, ecologists attempt approximating them to understand species’ distribution and plasticity to environmental changes, with invaluable repercussions on both theoretical and applied ecology. So far, individual‐based habitat selection models only characterized realized niches of populations delimited by physical (e.g. fences), historical (colonization) and biotic (competition) barriers constraining access to a subset of resources available to the species. As populations with different realized niches share the same fundamental niche, we developed a novel framework to scale‐up response curves from population‐scale habitat selection models to approximate the species’ optimal habitat choices, unbiased by barriers constraining accessibility. We used GPS‐locations from 147 wild mountain reindeer Rangifer t. tarandus, belonging to 7 of the remaining populations scattered throughout the subspecies’ range. We linked individual choices to accessible habitat features using conditional‐logistic regression with log‐link function in a use‐available design. Focal variables were modeled using 2nd degree polynomials on log‐scale, which correspond to a Gaussian curve used to approximate the fundamental niche optimum (curve mean) and breadth (variance). Using both real and simulated data we demonstrate that robust approximations of a fundamental niche optimum and breadth can be estimated using a relatively small number of representative populations with relatively few individuals. While each classical realized niche model had strong predictive power for the focal population but poorly predicted across populations, the approximation of the fundamental niche allowed for robust inter‐population comparisons in habitat quality. The proposed approach brings individual‐based habitat selection models forward along the continuum from investigating the realized niche of a population towards investigating a species’ fundamental niche, and allows us to quantify empirically the relationship between realized and fundamental niches. This allows improving the understanding of differences in fitness among populations, the prediction of species’ distributions and plasticity to environmental changes, and suggestions for mitigation priorities.     

Intragroup competition predicts individual foraging specialisation in a group‐living mammal

Individual foraging specialisation has important ecological implications, but its causes in group‐living species are unclear. One of the major consequences of group living is increased intragroup competition for resources. Foraging theory predicts that with increased competition, individuals should add new prey items to their diet, widening their foraging niche (‘optimal foraging hypothesis’). However, classic competition theory suggests the opposite: that increased competition leads to niche partitioning and greater individual foraging specialisation (‘niche partitioning hypothesis’). We tested these opposing predictions in wild, group‐living banded mongooses (Mungos mungo), using stable isotope analysis of banded mongoose whiskers to quantify individual and group foraging niche. Individual foraging niche size declined with increasing group size, despite all groups having a similar overall niche size. Our findings support the prediction that competition promotes niche partitioning within social groups and suggest that individual foraging specialisation may play an important role in the formation of stable social groupings.     

Testing the niche variation hypothesis with a measure of body condition

Individual variation and fitness are cornerstones of evolution by natural selection. The niche variation hypothesis (NVH) posits that when interspecific competition is relaxed, intraspecific competition should drive niche expansion by selection favoring use of novel resources and that among‐individual variation should confer a selective advantage. Population‐level niche expansion could be achieved by all individuals using all available resources, or by each individual using a unique combination of resources, thereby increasing among‐individual dietary niche variation. Although individual variation can lead to species‐level evolutionary and ecological change, observed variation does not ensure a beneficial outcome. We used carbon and nitrogen stable isotope analysis of claw keratin and a Bayesian stable isotope mixing model to estimate the summer (July–September) assimilated diet of individual female black Ursus americanus and brown U. arctos bears. We quantified variation in dietary niche in both populations, and assessed diet relative to percentage body fat. We hypothesized that if the NVH held, percentage body fat would be similar for individuals of the same species across much of the dietary range of observed proportional salmon contributions to individual bear diets. Although we found greater differences in dietary niches between than within species, we observed greater among‐individual dietary variation in the brown bear population. Moreover, we found that within each species individual female bears achieved similar ranges of percentage body fat at various levels of salmon in the diet. Our results provide support for the NVH. Linking individual dietary niches to measures of physiological condition related to fitness can offer new insights into eco‐evolutionary processes related to food resource use.     

Between‐individual variation drives the seasonal dynamics in the trophic niche of a Neotropical marsupial

The dynamics of population niches result from the variation in resource use within individuals and also from the variation between individuals. The prevalence of one mechanism or the other leads to competing hypotheses about the major mechanisms underlying the empirical observations of the contraction/expansion dynamics of the trophic niche in natural populations. In this study, we investigated how within‐ and between‐individual variation in resource use shapes the food niche dynamics of the woolly mouse opossum, Marmosa paraguayana (Didelphimorphia: Didelphidae), in a remnant of the highly seasonal Cerrado in south‐eastern Brazil. To do so, we analysed the faecal samples of live‐trapped individuals to determine their diets within the wet and dry seasons. In addition to a seasonal shift in the composition of the diet, the population trophic niche was significantly wider during the dry season than the wet season. This expansion resulted from larger between‐individual variation in the dry season that was not related to sex preferences, whereas the individual niche widths did not significantly increase from the wet to the dry seasons. Our findings add to the growing list of animal populations that show individual‐level variation in resource use. Furthermore, these results represent a pattern of individual‐level response to seasonal changes that is different from patterns reported for other organisms. We suggest that a pathway to build more realistic foraging models and produce more accurate predictions on population and community dynamics is to consider between‐individual variation and short‐term niche dynamics.     

Overlap and partitioning of the ecological and isotopic niches

Recently, it was proposed that stable isotope patterns can be used to quantify the width of the ecological niche of animals. However, the potential effects of habitat use on isotopic patterns of consumers have not been fully explored and consequently isotopic patterns may yield deceptive estimates of niche width. Here, we simulated four different scenarios of a consumer foraging across an isotopically heterogeneous landscape to test the combined effects of habitat and diet selection on the widths of the isotopic niche. We then modeled the actions of a naïve researcher who randomly sampled consumers from the simulated populations, and used these results to assess the overlap and partitioning of the isotopic and the ecological niches when habitat‐derived differences among isotope signatures are not considered. Our results suggest that populations of dietary specialists exhibited broader isotopic niches than populations composed of dietary generalists, and habitat generalists exhibited narrower isotopic niche widths compared with populations of individuals that foraged in specific habitats. The conversion of isotopic niches to ecological niches without knowledge of foraging behavior and habitat‐derived isotopic differences transformed an informative δ‐space into ‘a blurry p‐space’. Therefore, knowledge of habitat‐derived differences in stable isotope values and understanding of habitat use and individual foraging behavior are critical for the correct quantification of the ecological niche.     

Foraging and farming as niche construction: stable and unstable adaptations

All forager (or hunter-gatherer) societies construct niches, many of them actively by the concentration of wild plants into useful stands, small-scale cultivation, burning of natural vegetation to encourage useful species, and various forms of hunting, collectively termed 'low-level food production'. Many such niches are stable and can continue indefinitely, because forager populations are usually stable. Some are unstable, but these usually transform into other foraging niches, not geographically expansive farming niches. The Epipalaeolithic (final hunter-gatherer) niche in the Near East was complex but stable, with a relatively high population density, until destabilized by an abrupt climatic change. The niche was unintentionally transformed into an agricultural one, due to chance genetic and behavioural attributes of some wild plant and animal species. The agricultural niche could be exported with modifications over much of the Old World. This was driven by massive population increase and had huge impacts on local people, animals and plants wherever the farming niche was carried. Farming niches in some areas may temporarily come close to stability, but the history of the last 11,000 years does not suggest that agriculture is an effective strategy for achieving demographic and political stability in the world's farming populations.     

Beyond ecological opportunity: Prey diversity rather than abundance shapes predator niche variation

1. Ecological opportunity (i.e. the diversity of available resources) has a pivotal role in shaping niche variation and trophic specialisation of animals. However, ecological opportunity can be described with regard to both diversity and abundance of resources. The degree to which these two components contribute to niche variation remains unexplored.
2. To address this, we used an extensive dataset on fish diet and benthic invertebrate diversity and density from 73 sampling events in three Norwegian rivers in order to explore realised trophic niches and the response of dietary niche variation along gradients of resource diversity (potential trophic niches), resource density (as a proxy of resource abundance) and fish density (as a proxy of inter‐ and intra‐specific competition) in a freshwater top predator (the brown trout, Salmo trutta L.).
3. Linear models indicated that individual and population niche variation increased with increasing ecological opportunity in terms of prey diversity. However, no simple cause‐and‐effect associations between niche indices and prey abundance were found. Our multiple regression analyses indicated that the abundance of certain resources (e.g. Chironomidae) can interact with prey diversity to determine individual and population realised trophic niches. Niche variation (within‐individual component and inter‐individual diet variation) decreased with increasing inter‐ and intra‐specific competition.
4. This study extends prevailing trophic ecology theory by identifying diversity, rather than density, of available prey resources as a primary driver of niche variation in fish of temperate riverine systems with no extensive resource limitation. The study also shows that ecological opportunity may mask the direction of the effect (compression or expansion) of competition on niche variation when food resources are diverse.
5. Our study provides novel empirical insight to the driving forces behind niche variation and reveals that diversity, rather than density, of available prey resources may be a primary driver of niche variation in freshwater fish. Our study supports the view that a broader potential trophic niche promotes broader realised trophic niche variation by individuals, which leads to individual niche diversification by opening access to alternatives resources, resulting in a concomitant rise in the realised trophic niche width of the population.

     

Trophic niche width,offspring condition and immunity in a raptor species

Strategies developed by organisms to maximize foraging efficiency have a strong influence on fitness. The way in which the range of food resources is exploited has served to classify species, populations and individuals from more specialist (narrow trophic niche) to more generalist (broad trophic niche). Recent studies have provided evidence that many of the considered generalist species/populations are actually composed of different specialist individuals (individual specialization). Even the existence of generalism as an adaptive strategy has been questioned. In this study, we investigated the relationship between trophic niche width, individual quality and offspring viability in a population of common kestrel Falco tinnunculus during 4 years. We showed that the diet of kestrels varied significantly among years and that individuals of better quality fed their offspring with a higher diversity of prey species and a higher amount of food. Moreover, body condition and immune response of nestlings were positively correlated with diversity of prey delivered by parents. Our study suggests that generalism has the potential to increase fitness and that broadening the trophic niche may be an adaptive strategy in unpredictable environments.     

Temporal stability of niche use exposes sympatric Arctic charr to alternative selection pressures

There is now strong evidence that foraging niche specialisation plays a critical role in the very early stages of resource driven speciation. Here we test critical elements of models defining this process using a known polymorphic population of Arctic charr from subarctic Norway. We test the long-term stability of niche specialisation amongst foraging predators and discuss the possibility that contrasting foraging specialists are exposed to differing selection regimes. Inter-individual foraging niche stability was measured by combining two time-integrated ecological tracers of the foraging niche (each individual’s δ¹³C and δ¹⁵N stable isotope (SI) signatures and their food borne parasite fauna) with a short-term measure of foraging niche use (stomach contents composition). Three dietary subgroups of predators were identified, including zooplankton, gammarid and benthivore specialists foragers. Zooplanktivorous specialists had muscle low in δ ¹³C, a high abundance of parasites transmitted from pelagic copepods, a smaller head, longer snout and a more slender body-form than gammaridivorous specialist individuals which had muscle more enriched in δ ¹³C and high abundance of parasites transmitted from benthic Gammarus. Benthivorous individuals were intermediate between the other two foraging groups according to muscle SI-signals (δ¹³C) and loadings of parasites transmitted from both copepods and Gammarus. The close relationship between subgroups identified by stomach contents, time-integrated tracers of niche use (SI and parasites) and functional trophic morphology (niche adaptations) demonstrate a long-term temporally stable niche use of each individual predator. Differential habitat use and contrasting parasite communities and loadings, show differential exposure to different suites of selection pressures for different foraging specialists. Results also show that individual specialisation in trophic behaviour and thus exposure to different suites of selection pressures are stable over time, and thus provide a platform for disruptive selection to operate within this sympatric system.     

Stable isotope series from elephant ivory reveal lifetime histories of a true dietary generalist

Longitudinal studies have revealed how variation in resource use within consumer populations can impact their dynamics and functional significance in communities. Here, we investigate multi-decadal diet variations within individuals of a keystone megaherbivore species, the African elephant (Loxodonta africana), using serial stable isotope analysis of tusks from the Kruger National Park, South Africa. These records, representing the longest continuous diet histories documented for any extant species, reveal extensive seasonal and annual variations in isotopic--and hence dietary--niches of individuals, but little variation between them. Lack of niche distinction across individuals contrasts several recent studies, which found relatively high levels of individual niche specialization in various taxa. Our result is consistent with theory that individual mammal herbivores are nutritionally constrained to maintain broad diet niches. Individual diet specialization would also be a costly strategy for large-bodied taxa foraging over wide areas in spatio-temporally heterogeneous environments. High levels of within-individual diet variability occurred within and across seasons, and persisted despite an overall increase in inferred C(4) grass consumption through the twentieth century. We suggest that switching between C(3) browsing and C(4) grazing over extended time scales facilitates elephant survival through environmental change, and could even allow recovery of overused resources.     

Landscape heterogeneity drives intra‐population niche variation and reproduction in an arctic top predator

While intra‐population variability in resource use is ubiquitous, little is known of how this measure of niche diversity varies in space and its role in population dynamics. Here we examined how heterogeneous breeding environments can structure intra‐population niche variation in both resource use and reproductive output. We investigated intra‐population niche variation in the Arctic tundra ecosystem, studying peregrine falcon (Falco peregrinus tundrius, White) breeding within a terrestrial‐marine gradient near Rankin Inlet, Nunavut, Canada. Using stable isotope analysis, we found that intra‐population niches varied at the individual level; we examined within‐nest and among‐nest variation, though only the latter varied along the terrestrial‐marine gradient (i.e., increased among‐nest variability among birds nesting within the marine environment, indicating higher degree of specialization). Terrestrial prey species (small herbivores and insectivores) were consumed by virtually all falcons. Falcons nesting within the marine environment made use of marine prey (sea birds), but depended heavily on terrestrial prey (up to 90% of the diet). Using 28‐years of peregrine falcon nesting data, we found a positive relationship between the proportion of terrestrial habitat surrounding nest sites and annual nestling production, but no relationship with the likelihood of successfully rearing at least one nestling reaching 25 days old. Annually, successful inland breeders raised 0.47 more young on average compared to offshore breeders, which yields potential fitness consequences for this long‐living species. The analyses of niche and reproductive success suggest a potential breeding cost for accessing distant terrestrial prey, perhaps due to additional traveling costs, for those individuals with marine nest site locations. Our study indicates how landscape heterogeneity can generate proximate (niche variation) and ultimate (reproduction) consequences on a population of generalist predator. We also show that within‐individual and among‐individual variation are not mutually exclusive, but can simultaneously arise and structure intra‐population niche variation.     

Determining trophic niche width: an experimental test of the stable isotope approach

Determining the trophic niche width of an animal population and the relative degree to which a generalist population consists of dietary specialists are long‐standing problems of ecology. It has been proposed that the variance of stable isotope values in consumer tissues could be used to quantify trophic niche width of consumer populations. However, this promising idea has not yet been rigorously tested. By conducting controlled laboratory experiments using model consumer populations (Daphnia sp., Crustacea) with controlled diets, we investigated the effect of individual‐ and population‐level specialisation and generalism on consumer δ¹³C mean and variance values. While our experimental data follow general expectations, we extend current qualitative models to quantitative predictions of the dependence of isotopic variance on dietary correlation time, a measure for the typical time over which a consumer changes its diet. This quantitative approach allows us to pinpoint possible procedural pitfalls and critical sources of measurement uncertainty. Our results show that the stable isotope approach represents a powerful method for estimating trophic niche widths, especially when taking the quantitative concept of dietary correlation time into account.     

Natal habitat effects drive density—dependent scaling of dispersal decisions

The dispersal behavior of a species is critical for the stability and persistence of its populations across a landscape. How population density affects dispersal decisions is important for predicting these dynamics, as the form of density‐dependent dispersal influences the stability and persistence of populations. Natal habitat experience often has strong impacts on individual dispersal behavior as well, but its influence on density‐dependent dispersal behaviors remains unexplored. Here we address this conceptual gap in two experiments separately examining habitat selection and emigration from recently colonized patches for two species of flour beetle Tribolium sp. We found that interactions between the quality of habitat experienced during natal development and current habitat for dispersal capable adults can strongly affect the form of density dependence, including reversing the direction of nonlinearities (accelerating to decelerating), or even negating the influence of population density for individual dispersal decisions. Across heterogeneous landscapes, where individuals from different populations may experience different natal habitats, this altering of density‐dependent relationships is predicted by theory to fundamentally influence regional population dynamics. Our results indicate that species which occur across heterogeneous environments, such as during conservation reintroductions, or as invasive species spread, have much potential for natal experience to interact with density dependence and influence local and regional population dynamics.     

Synergistic selection between ecological niche and mate preference primes diversification

The ecological niche and mate preferences have independently been shown to be important for the process of speciation. Here, we articulate a novel mechanism by which ecological niche use and mate preference can be linked to promote speciation. The degree to which individual niches are narrow and clustered affects the strength of divergent natural selection and population splitting. Similarly, the degree to which individual mate preferences are narrow and clustered affects the strength of divergent sexual selection and assortative mating between diverging forms. This novel perspective is inspired by the literature on ecological niches; it also explores mate preferences and how they may contribute to speciation. Unlike much comparative work, we do not search for evolutionary patterns using proxies for adaptation and sexual selection, but rather we elucidate how ideas from niche theory relate to mate preference, and how this relationship can foster speciation. Recognizing that individual and population niches are conceptually and ecologically linked to individual and population mate preference functions will significantly increase our understanding of rapid evolutionary diversification in nature. It has potential to help solve the difficult challenge of testing the role of sexual selection in the speciation process. We also identify ecological factors that are likely to affect individual niche and individual mate preference in synergistic ways and as a consequence to promote speciation. The ecological niche an individual occupies can directly affect its mate preference. Clusters of individuals with narrow, differentiated niches are likely to have narrow, differentiated mate preference functions. Our approach integrates ecological and sexual selection research to further our understanding of diversification processes. Such integration may be necessary for progress because these processes seem inextricably linked in the natural world.     

Niches in the Anthropocene: passerine assemblages show niche expansion from natural to urban habitats

Human‐mediated habitat transformation is increasingly evident around the world. Yet, how this transformation influences species’ niche width and overlap remains unclear. On the one hand, human‐mediated habitat transformation promotes increased species similarity through trait‐based filtering, and an increased prevalence of generalist species with broad niches, resulting in functional homogenization. On the other hand, species that colonize transformed habitats could use empty niches, resulting in decreased species similarity and an expansion of assemblage‐level niche space. Here we explore these two alternatives in eight highly diverse passerine assembles in natural, rural and urban habitats in south and southwest China, a rapidly developing region of the world. Based on stable isotopes, we found that species’ niche width increased from natural to human‐made habitats, but there were no differences in niche overlap among habitats. Therefore, we found evidence for niche expansion, with generalists appearing to use empty niches created by human habitat modification, and with assemblages being comprised of complementary species. Further research is needed to determine whether increased between‐ or within‐individual niche variation is the main driver of niche expansion in transformed habitats.