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

The quantification of individuality is a common research theme in the fields of population, community, and evolutionary ecology. The potential for individuality to arise is likely context-dependent, and the influence of habitat characteristics on its prevalence has received less attention than intraspecific competition. We examined individual diet specialization in 16 sea otter (Enhydra lutris) populations from southern California to the Aleutian Islands in Alaska. Because population histories, relative densities, and habitat characteristics vary widely among sites, we could examine the effects of intraspecific competition and habitat on the prevalence of individual diet specialization. Using observed diet data, we classified half of our sites as rocky substrate habitats and the other half containing a mixture of rocky and unconsolidated (soft) sediment substrates. We used stable isotope data to quantify populationand individual-level diet variation. Among rocky substrate sites, the slope [±standard error (SE)] of the positive significant relationship between the within-individual component (WIC) and total isotopie niche width (TINW) was shallow (0.23 ± 0.07) and negatively correlated with sea otter density. In contrast, the slope of the positive WIC/TINW relationship for populations inhabiting mixed substrate habitats was much higher (0.53 ± 0.14), suggesting a low degree of individuality, irrespective of intraspecific competition. Our results show that the potential for individuality to occur as a result of increasing intraspecific competition is contextdependent and that habitat characteristics, which ultimately influence prey diversity, relative abundance, and the range of skillsets required for efficient prey procurement, are important in determining when and where individual diet specialization occurs in nature.

Among-individual diet variation is common in natural populations and may occur at any trophic level within a food web. Yet, little is known about its variation among trophic levels and how such variation could affect phenotypic divergence within populations. In this study we investigate the relationships between trophic position (the population's range and average) and among-individual diet variation. We test for diet variation among individuals and across size classes of Eurasian perch (Perca fluviatilis), a widespread predatory freshwater fish that undergoes ontogenetic niche shifts. Second, we investigate among-individual diet variation within fish and invertebrate populations in two different lake communities using stable isotopes. Third, we test potential evolutionary implications of population trophic position by assessing the relationship between the proportion of piscivorous perch (populations of higher trophic position) and the degree of phenotypic divergence between littoral and pelagic perch sub-populations. We show that amongindividual diet variation is highest at intermediate trophic positions, and that this high degree of among-individual variation likely causes an increase in the range of trophic positions among individuals. We also found that phenotypic divergence was negatively related to trophic position in a population. This study thus shows that trophic position is related to and may be important for among-individual diet variation as well as to phenotypic divergence within populations.

Issue Title: Special Feature on Individual-level niche specialization

Many ecologically generalized populations are composed of relatively specialized individuals that selectively consume a subset of their population's diet, a phenomenon known as 'individual specialization'. The Niche Variation Hypothesis posits that this individual specialization can arise during ecological release if niche expansion occurs mainly through diet divergence among individuals, leading to greater morphological variation. Most tests of this hypothesis have searched for correlations between niche width and morphological variance, but this approach rests on the untested assumption that within-population morphological diversity is highly correlated with ecological diversity. Here, we test whether intrapopulation diet variation is correlated with intrapopulation morphological variation, across 12 lacustrine populations of three-spine stickleback. First, we use behavioral observations, isotopes, and gut contents to show that, within populations, individuals differ in microhabitat use and diet. Second, we show that some populations exhibit more diet variation than others, as evidenced by differences in both isotopie and gut content variation among individuals. Finally, we confirm that populations with greater dietary variation are more morphologically variable. However, this relationship is only significant when total morphological variance is examined, not for individual morphological traits. This discordance may reflect among-population differences in the relationship between individual morphology and diet. Because morphology-diet relationships can differ among populations, morphological variance may be a poor predictor of actual diet variation when diverse populations are being compared.

Many mobile marine species are presumed to utilize a broad spectrum of habitats, but this seemingly generalist life history may arise from conspecifics specializing on distinct habitat alternatives to exploit foraging, resting/refuge, or reproductive opportunities. We acoustically tagged 34 red drum, and mapped sand, seagrass, marsh, or oyster (across discrete landscape contexts) use by each uniquely coded individual. Using 144,000 acoustic detections, we recorded differences in habitat use among red drum: proportional use of seagrass habitat ranged from 0 to 100 %, and use of oyster-bottom types also varied among fish. WIC/TNW and IS metrics (previously applied vis-à-vis diet specialization) consistently indicated that a typical red drum overlapped >70 % with population-level niche exploitation. Monte Carlo permutations showed these values were lower than expected had fish drawn from a common habitat-use distribution, but longitudinal comparisons did not provide evidence of temporally consistent individuality, suggesting that differences among individuals were plastic and not reflective of true specialization. Given the range of acoustic detections we captured (from tens to 1,000s per individual), which are substantially larger sample sizes than in many diet studies, we extended our findings by serially reducing or expanding our data in simulations to evaluate samplesize effects. We found that the results of null hypothesis testing for specialization were highly dependent on sample size, with thresholds in the relationship between sample size and associated P-values. These results highlight opportunities and potential caveats in exploring individuality in habitat use. More broadly, exploring individual specialization in fine-scale habitat use suggests that, for mobile marine species, movement behaviors over shorter (≤weeks), but not longer (≥months), timescales may serve as an underlying mechanism for other forms of resource specialization.

Individual niche specialization (INS) is increasingly recognized as an important component of ecological and evolutionary dynamics. However, most studies that have investigated INS have focused on the effects of niche width and inter- and intraspecific competition on INS in small-bodied species for short time periods, with less attention paid to INS in large-bodied reptilian predators and the effects of available prey types on INS. We investigated the prevalence, causes, and consequences of INS in foraging behaviors across different populations of American alligators (Alligator mississippiensis), the dominant aquatic apex predator across the southeast US, using stomach contents and stable isotopes. Gut contents revealed that, over the short term, although alligator populations occupied wide ranges of the INS spectrum, general patterns were apparent. Alligator populations inhabiting lakes exhibited lower INS than coastal populations, likely driven by variation in habitat type and available prey types. Stable isotopes revealed that over longer time spans alligators exhibited remarkably consistent use of variable mixtures of carbon pools (e.g., marine and freshwater food webs). We conclude that INS in large-bodied reptilian predator populations is likely affected by variation in available prey types and habitat heterogeneity, and that INS should be incorporated into management strategies to efficiently meet intended goals. Also, ecological models, which typically do not consider behavioral variability, should include INS to increase model realism and applicability.

Although intra-population variation in niches is a widespread phenomenon with important implications for ecology, evolution and management of a range of animal species, the causes and consequences of this variation remain poorly understood. We used stable isotope analysis to characterise foraging niches and to investigate the causes and consequences of individual niche variation in the European badger, a mustelid mammal that lives in territorial social groups, but forages alone. We found that the degree of individual niche variation within social groups was negatively related to the availability of farmland habitats, which represent an important foraging habitat for badgers; and was positively related to territory size, supporting the idea that resource limitation and ecological opportunity lead to increased individual specialisation. We also found that the degree of individual specialisation related to an individual's body condition and that this effect varied with ecological context; such that specialisation had a stronger positive relationship with body condition in social groups with reduced availability of key farmland habitats. Body condition was also related to the utilisation of specific resources (woodland invertebrates), but again this relationship varied with the availability of farmland foraging habitats. This study supports the idea that resource availability plays an important role in determining patterns of individual niche variation, and identifies the potential adaptive consequences of specialised foraging strategies.

The focus of evolutionary behavioural ecologists has recently turned towards understanding the causes and consequences of behavioural consistency, manifesting either as animal personality (consistency in a single behaviour) or behavioural syndrome (consistency across more behaviours). Behavioural type (mean individual behaviour) has been linked to life-history strategies, leading to the emergence of the integrated pace-of-life syndrome (POLS) theory. Using Rana dalmatina tadpoles as models, we tested if behavioural consistency and POLS could be detected during the early ontogenesis of this amphibian. We targeted two ontogenetic stages and measured activity, exploration and risk-taking in a common garden experiment, assessing both individual behavioural type and intraindividual behavioural variation. We observed that activity was consistent in all tadpoles, exploration only became consistent with advancing age and risk-taking only became consistent in tadpoles that had been tested, and thus disturbed, earlier. Only previously tested tadpoles showed trends indicative of behavioural syndromes. We found an activity—age at metamorphosis POLS in the previously untested tadpoles irrespective of age. Relative growth rate correlated positively with the intra-individual variation of activity of the previously untested older tadpoles. In previously tested older tadpoles, intra-individual variation of exploration correlated negatively and intra-individual variation of risk-taking correlated positively with relative growth rate. We provide evidence for behavioural consistency and POLS in predator- and conspecific-naive tadpoles. Intraindividual behavioural variation was also correlated to life history, suggesting its relevance for the POLS theory. The strong effect of moderate disturbance related to standard behavioural testing on later behaviour draws attention to the pitfalls embedded in repeated testing.

Intraspecific variation in behavior and diet can have important consequences for population and ecosystem dynamics. Here, we examine how differences in reproductive investment and spatial ecology influence individual diet specialization in male and female southern sea otters (Enhydra lutris nereis). We hypothesize that greater reproductive constraints and smaller home ranges of females lead to more pronounced intraspecific competition and increased specialization. We integrate stable carbon (δ¹³C) and nitrogen (δ¹³N) isotope analysis of sea otter vibrissae with long-term observational studies of five subpopulations in California. We define individual diet specialization as low ratios of within-individual variation (WIC) to total population niche width (TNW). We compare isotopie and observational based metrics of WIC/TNW for males and females to data on population densities, and movement patterns using both general linear and linear mixed-effects models. Consistent with our hypothesis, increasing population density is associated with increased individual diet specialization by females but not by males. Additionally, we find the amount of coastline in a sea otter's home range positively related with individual dietary variability, with increased range span resulting in weaker specialization for both males and females. We attribute our results to sex-based differences in movement, with females needing to specialize in their small ranges to maximize energy gain, and posit that the paradigm of individual prey specialization in sea otters with increased intraspecific competition may be a pattern driven largely by females. Our work highlights a potentially broader role of sex in the mechanistic pressures promoting and maintaining diet specialization.