Evidence for large-scale effects of competition: niche displacement in Canada lynx and bobcat

Determining the patterns, causes and consequences of character displacement is central to our understanding of competition in ecological communities. However, the majority of competition research has occurred over small spatial extents or focused on fine-scale differences in morphology or behaviour. The effects of competition on broad-scale distribution and niche characteristics of species remain poorly understood but critically important. Using range-wide species distribution models, we evaluated whether Canada lynx (Lynx canadensis) or bobcat (Lynx rufus) were displaced in regions of sympatry. Consistent with our prediction, we found that lynx niches were less similar to those of bobcat in areas of sympatry versus allopatry, with a stronger reliance on snow cover driving lynx niche divergence in the sympatric zone. By contrast, bobcat increased niche breadth in zones of sympatry, and bobcat niches were equally similar to those of lynx in zones of sympatry and allopatry. These findings suggest that competitively disadvantaged species avoid competition at large scales by restricting their niche to highly suitable conditions, while superior competitors expand the diversity of environments used. Our results indicate that competition can manifest within climatic niche space across species’ ranges, highlighting the importance of biotic interactions occurring at large spatial scales on niche dynamics.     

Fine‐scale habitat selection by sympatric Canada lynx and bobcat

The Canada lynx (Lynx canadensis) and the bobcat (Lynx rufus) are closely related species with overlap at their range peripheries, but the factors that limit each species and the interactions between them are not well understood. Habitat selection is a hierarchical process, in which selection at higher orders (geographic range, home range) may constrain selection at lower orders (within the home range). Habitat selection at a very fine scale within the home range has been less studied for both lynx and bobcat compared to selection at broader spatiotemporal scales. To compare this fourth‐order habitat selection by the two species in an area of sympatry, we tracked lynx and bobcat during the winters of 2017 and 2018 on the north shore of Lake Huron, Ontario. We found that both lynx and bobcat selected shallower snow, higher snowshoe hare abundance, and higher amounts of coniferous forest at the fourth order. However, the two species were spatially segregated at the second order, and lynx were found in areas with deeper snow, more snowshoe hare, and more coniferous forest. Taken together, our findings demonstrate that the lynx and bobcat select different resources at the second order, assorting along an environmental gradient in the study area, and that competition is unlikely to be occurring between the two species at finer scales.     

Growth trades off with habitat specialization

How differences in niche breadth evolve and are maintained remains largely unknown. The 'jack of all trades is master of none' model of resource specialization has been widely considered, but, to our knowledge, never before supported empirically. It invokes performance trade-offs associated with specialization. Specialists should outperform generalists on a subset of resources, but be unable to maintain high performance over a broader range of resources. By contrast, generalists should perform less well, on average, using a greater diversity of resources. We report such trade-offs among four coral goby species in the wild. Habitat specialists grew faster than generalists in one of two habitats. Average growth rates of generalists were less than that of specialists, but more consistent between habitats. Performance trade-offs associated with resource specialization could influence the evolution and maintenance of narrow niche breadth.     

Biogeography of the Primates of Guyana: Effects of Habitat Use and Diet on Geographic Distribution

Species that exploit a wide range of resources or habitats (generalists) tend to be widely distributed, whereas species that exploit a narrow range of resources or habitats (specialists) often have a limited distribution. The distribution patterns are thought to result from specialists using relatively smaller habitats than those exploited by generalists. I used data from 1,725 km of primate surveys that I conducted in Guyana to test these hypotheses. Habitat breadth is the total number of different habitat types occupied by each species. I used the total number of different food categories exploited by each species to measure dietary breadth. Geographic range size is correlated with habitat breadth but not with dietary breadth or body size for the 8 primate species in Guyana. Habitat generalists—red howlers and wedge-capped capuchins—range into all habitats. Habitat specialists—spider monkeys, brown bearded sakis, and golden-handed tamarins—range only into large habitats. Habitat generalists tend to be dietary type specialists in Guyana. I suggest that only habitat generalists can subsist on the low-quality foods in small habitats in Guyana. Conversely, habitat specialists tend to be dietary type generalists in Guyana. They must feed on a variety of food types in large habitats. However, using the number of food categories exploited as a measure of dietary breadth may be only a weak aspect of multidimensional niche. Researchers testing biogeographic hypotheses associated with dietary breadth should consider including multivariate indicators of both the types of food categories eaten and the number of plant species exploited.     

Signatures of selection in mammalian clock genes with coding trinucleotide repeats: Implications for studying the genomics of high‐pace adaptation

Climate change is predicted to affect the reproductive ecology of wildlife; however, we have yet to understand if and how species can adapt to the rapid pace of change. Clock genes are functional genes likely critical for adaptation to shifting seasonal conditions through shifts in timing cues. Many of these genes contain coding trinucleotide repeats, which offer the potential for higher rates of change than single nucleotide polymorphisms (SNPs) at coding sites, and, thus, may translate to faster rates of adaptation in changing environments. We characterized repeats in 22 clock genes across all annotated mammal species and evaluated the potential for selection on repeat motifs in three clock genes (NR1D1, CLOCK, and PER1) in three congeneric species pairs with different latitudinal range limits: Canada lynx and bobcat (Lynx canadensis and L. rufus), northern and southern flying squirrels (Glaucomys sabrinus and G. volans), and white‐footed and deer mouse (Peromyscus leucopus and P. maniculatus). Signatures of positive selection were found in both the interspecific comparison of Canada lynx and bobcat, and intraspecific analyses in Canada lynx. Northern and southern flying squirrels showed differing frequencies at common CLOCK alleles and a signature of balancing selection. Regional excess homozygosity was found in the deer mouse at PER1 suggesting disruptive selection, and further analyses suggested balancing selection in the white‐footed mouse. These preliminary signatures of selection and the presence of trinucleotide repeats within many clock genes warrant further consideration of the importance of candidate gene motifs for adaptation to climate change.     

Microbial generalist or specialist: Intraspecific variation and dormancy potential matter

Microbial generalists and specialists coexist in the soil environment while having distinctive impacts on microbial community dynamics. In microbial ecology, the underlying mechanisms as to why a species is a generalist or a specialist remain ambiguous. Herein, we collected soils across a national scale and identified bacterial generalists and specialists according to niche breadth at the species level (OTU level), and the single-nucleotide differences in each species were measured to investigate intraspecific variation (at zero-radius OTU level). Compared with that of the specialists, the intraspecific variation of the generalists was much higher, which ensured their wider niche breadth and lower variability. The higher asynchrony and different niche preferences of conspecific individuals and the higher dormancy potential within the generalists further contributed to their stability in varying environments. Besides, generalists were less controlled by environmental filtering, which was indicated by the stronger signature of stochastic processes in their assembly, and had higher diversification and transition rates that allowed them to adapt to environmental changes to a greater extent than specialists. Overall, this study provides a new comprehensive understanding of the rules of assembly and the evolutionary roles of bacterial generalists and specialists. It also highlights the importance of intraspecific variation and the dormancy potential in the stability of species.     

Variable effects of wolves on niche breadth and density of intraguild competitors

1. The parallel niche release hypothesis (PNR) indicates that reduced competition with dominant competitors results in greater density and niche breadth of subordinate competitors and which may support an adaptive advantage.
2. We assessed support for the PNR by evaluating relationships between variation in niche breadth and intra‐ and interspecific density (an index of competition) of wolves (Canis lupus) coyotes (C. latrans), and bobcats (Lynx rufus).
3. We estimated population density (wolf track surveys, coyote howl surveys, and bobcat hair snare surveys) and variability in space use (50% core autocorrelated kernel density home range estimators), temporal activity (hourly and overnight speed), and dietary (isotopic δ¹³C and δ¹⁵N) niche breadth of each species across three areas of varying wolf density in the Upper Peninsula of Michigan, USA, 2010–2019.
4. Densities of wolves and coyotes were inversely related, and increased variability in space use, temporal activity, and dietary niche breadth of coyotes was associated with increased coyote density and decreased wolf density supporting the PNR. Variability in space use and temporal activity of wolves and dietary niche breadth of bobcats also increased with increased intraspecific density supporting the PNR.
5. Through demonstrating decreased competition between wolves and coyotes and increased coyote niche breadth and density, our study provides multidimensional support for the PNR. Knowledge of the relationship between niche breadth and population density can inform our understanding of the role of competition in shaping the realized niche of species.

     

Seasonal Resource Selection of Canada Lynx in Managed Forests of the Northern Rocky Mountains

ABSTRACT We investigated seasonal patterns in resource selection of Canada lynx (Lynx canadensis) in the northern Rockies (western MT, USA) from 1998 to 2002 based on backtracking in winter (577 km; 10 M, 7 F) and radiotelemetry (630 locations; 16 M, 11 F) in summer. During winter, lynx preferentially foraged in mature, multilayer forests with Engelmann spruce (Picea engelmannii) and subalpine fir (Abies lasiocarpa) in the overstory and midstory. Forests used during winter were composed of larger diameter trees with higher horizontal cover, more abundant snowshoe hares (Lepus americanus), and deeper snow compared to random availability; multilayer, spruce-fir forests provided high horizontal cover with tree branching that touched the snow surface. During winter, lynx killed prey at sites with higher horizontal cover than that along foraging paths. Lynx were insensitive to snow depth or penetrability in determining where they killed prey. During summer, lynx broadened their resource use to select younger forests with high horizontal cover, abundant total shrubs, abundant small-diameter trees, and dense saplings, especially spruce-fir saplings. Based on multivariate logistic-regression models, resource selection occurred primarily at a fine spatial scale as was consistent with a sight-hunting predator in dense forests. However, univariate comparisons of patch-level metrics indicated that lynx selected homogenous spruce-fir patches, and avoided recent clear-cuts or other open patches. Given that lynx in Montana exhibit seasonal differences in resource selection, we encourage managers to maintain habitat mosaics. Because winter habitat may be most limiting for lynx, these mosaics should include abundant multistory, mature spruce-fir forests with high horizontal cover that are spatially well-distributed.     

Habitat utilization by coral reef fish: implications for specialists vs. generalists in a changing environment

1. The impact of environmental disturbance and habitat loss on associated species is expected to be dependent on a species' level of specialization. We examined habitat use and specialization of coral reef fish from the diverse and ecologically important family Pomacentridae, and determined which species are susceptible to declines in coral cover due to disturbance induced by crown-of-thorns seastar (COTS, Acanthaster planci L.). 2. A high proportion of pomacentrid species live in association with live coral as adults (40%) or juveniles (53%). Adults of many species had strong affiliations with branching corals, while juveniles favoured plating growth forms, reflecting the sizes of refuge provided by coral types. 3. Juveniles of species that associated with coral had narrower niche breadths than adult conspecifics, due to associations with specific coral types. The especially high coral association and narrower niche breadth of juveniles suggest that the presence of live coral is crucial for many species during early life history, and that disturbance-induced coral loss may have serious flow-on effects on adult abundance. 4. Microhabitat availability was a poor predictor of fish species abundance. Significant correlations between coverage of coral types and abundance of five adults and two juvenile species were detected; however, these relationships explained <35% and <10% of the variation in abundance of adult and juvenile species, respectively. 5. Niche breadth explained 74% of the variation in species' mean response to coral decline and it is clear that disturbance has a greater impact on resource specialists, suggesting that increasing frequency and intensity of coral loss will cause reef fish communities to become dominated by habitat generalists at the expense of coral-dwelling specialists.     

Trophic Niche in a Raptor Species: The Relationship between Diet Diversity,Habitat Diversity and Territory Quality

Recent research reports that many populations of species showing a wide trophic niche (generalists) are made up of both generalist individuals and individuals with a narrow trophic niche (specialists), suggesting trophic specializations at an individual level. If true, foraging strategies should be associated with individual quality and fitness. Optimal foraging theory predicts that individuals will select the most favourable habitats for feeding. In addition, the “landscape heterogeneity hypothesis” predicts a higher number of species in more diverse landscapes. Thus, it can be predicted that individuals with a wider realized trophic niche should have foraging territories with greater habitat diversity, suggesting that foraging strategies, territory quality and habitat diversity are inter-correlated. This was tested for a population of common kestrels Falco tinnunculus. Diet diversity, territory occupancy (as a measure of territory quality) and habitat diversity of territories were measured over an 8-year period. Our results show that: 1) territory quality was quadratically correlated with habitat diversity, with the best territories being the least and most diverse; 2) diet diversity was not correlated with territory quality; and 3) diet diversity was negatively correlated with landscape heterogeneity. Our study suggests that niche generalist foraging strategies are based on an active search for different prey species within or between habitats rather than on the selection of territories with high habitat diversity.     

Linking habitat specialization with species' traits in European birds

Ecological specialization provides information about adaptations of species to their environment. However, identification of traits representing the relevant dimensions of ecological space remains challenging. Here we endeavoured to explain how complex habitat specializations relate to various ecological traits of European birds. We employed phylogenetic generalized least squares and information theoretic approach statistically controlling for differences in geographic range size among species. Habitat specialists had narrower diet niche, wider climatic niche, higher wing length/tail length ratio and migrated on shorter distances than habitat generalists. Our results support an expected positive link between habitat and diet niche breadth estimates, however a negative relationship between habitat and climate niche breadths is surprising. It implies that habitat specialists occur mostly in spatially restricted environments with high climatic variability such as mountain areas. This, however, complicates our understanding of predicted impacts of climatic changes on avian geographical distributions. Our results further corroborate that habitat specialization reflects occupation of morphological space, when specialists depend more on manoeuvrability of the flight and are thus more closely associated to open habitats than habitat generalists. Finally, our results indicate that long distance movements might hamper narrow habitat preferences. In conclusion, we have shown that species’ distributions across habitats are informative about their positions along other axes of ecological space and can explain states of particular functional traits, however, our results also reveal that the links between different niche estimates cannot be always straightforwardly predicted.     

Demographic differences in diet breadth of Canada lynx during a fluctuation in prey availability

Population dynamics of specialist carnivores are closely linked to prey availability, but the extent of variability in diet breadth of individual carnivores relative to natural variability in the abundance of their primary prey is not well understood. Canada lynx (Lynx canadensis) specialize on snowshoe hares (Lepus americanus) and exhibit cyclic fluctuations in abundance that lag 1–2 years behind those of snowshoe hares. Declining hare densities spur demographic changes in lynx, but it is unclear whether a corresponding increase in diet breadth occurs: (1) broadly across a lynx population; (2) only among individuals who are able to effectively switch to alternative prey; or (3) only among individuals who cannot capture sufficient primary prey. We measured stable isotope ratios of lynx muscle tissue spanning a cyclic increase and decline in hare density (1998–2001) in Fort Providence, NT, Canada. We found that lynx cohorts responded differently to hare population change, with yearling animals having broader diets at low hare densities, while adults and dependent juveniles maintained a constant diet through the initial decline in hare density. This result was consistent irrespective of lynx sex and indicates that yearling lynx likely are forced to adopt a broader diet when primary prey densities decline. Our results imply that select cohorts of specialist carnivores can exhibit high dietary plasticity in response to changes in primary prey abundance, prompting the need to determine whether increased diet breadth in young lynx is a successful strategy for surviving through periods of snowshoe hare scarcity. In this way, cohort‐specific niche expansion could strongly affect the dynamics of organisms exhibiting population cycles.     

Competition and coexistence in sympatric bobcats and pumas

Space use and diets of sympatric bobcats Lynx rufus and pumas Puma concolor were compared using sign surveys and scat analysis during 1997–2002 in south-eastern Arizona, USA. Bobcats appeared to use grassland, scrub, riparian and woodland habitats equally, but pumas had higher activity in riparian and woodland habitats. There was little evidence that bobcats avoided pumas in space use. Bobcats ate primarily rodents (33% of items in scats), lagomorphs (32%) and ungulates (16%), whereas pumas ate primarily ungulates (69%) and carnivores (21%). Pumas had a narrower dietary niche breadth than bobcats, and puma diet overlapped bobcat diet by 56%, suggesting that pumas may be more vulnerable to changes in prey density than bobcats. Pumas also killed and consumed bobcats, indicating that interference competition may be manifesting through intraguild predation.     

Rarity in Australian desert lizards

Most species of Australian desert lizards are uncommon. Possible causes of rarity are examined, including body size as measured by snout–vent length (SVL), fecundity, number of sites occupied, habitat niche breadth, microhabitat niche breadth, dietary niche breadth, and average total niche overlap with other species. Rare species tend to be larger with lower fecundities than abundant species and they occur at fewer sites. Many, but not all, uncommon species are specialists, either in habitat, microhabitat, or diet. The niche breadth hypothesis, which states that abundant species should be generalists whereas specialized species should be rare, is tested, but rejected as a general explanation for rarity. Some uncommon species exhibit high overlap with other species suggesting that they may experience diffuse competition. However, no single cause of rarity can be identified, but rather each species has its own idiosyncratic reasons for being uncommon. Multivariate analyses show distinct ecological differences between abundant and uncommon species.     

The generalism–specialism debate: the role of generalists in the life and death of species

Specialisms on resources and for niches, leading to specialization, have been construed to be tantamount to speciation and vice versa, while the occurrence of true generalism in nature has also been questioned. We argue that generalism in resource use, biotope occupancy, and niche breadth not only exists, but also forms a crucial part in the evolution of specialists, representing a vital force in speciation and a more effective insurance against extinction. We model the part played by generalism and specialism in speciation and illustrate how a balance may be maintained between the number of specialists and generalists within taxa. The balance occurs as an ongoing cycle arising from turnover in the production of specialists and generalists, speciation, and species extinction. The nature of the balance depends on the type of resources exploited, biotopes, and niche space occupied. These vary between different regions and create taxonomic biases towards generalists or specialists. We envisage that the process may be sympatric/parapatric, although it is more likely initiated by allopatry driven by abiotic forces. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 725–737.     

Metabolic Flexibility as a Major Predictor of Spatial Distribution in Microbial Communities

A better understand the ecology of microbes and their role in the global ecosystem could be achieved if traditional ecological theories can be applied to microbes. In ecology organisms are defined as specialists or generalists according to the breadth of their niche. Spatial distribution is often used as a proxy measure of niche breadth; generalists have broad niches and a wide spatial distribution and specialists a narrow niche and spatial distribution. Previous studies suggest that microbial distribution patterns are contrary to this idea; a microbial generalist genus (Desulfobulbus) has a limited spatial distribution while a specialist genus (Methanosaeta) has a cosmopolitan distribution. Therefore, we hypothesise that this counter-intuitive distribution within generalist and specialist microbial genera is a common microbial characteristic. Using molecular fingerprinting the distribution of four microbial genera, two generalists, Desulfobulbus and the methanogenic archaea Methanosarcina, and two specialists, Methanosaeta and the sulfate-reducing bacteria Desulfobacter were analysed in sediment samples from along a UK estuary. Detected genotypes of both generalist genera showed a distinct spatial distribution, significantly correlated with geographic distance between sites. Genotypes of both specialist genera showed no significant differential spatial distribution. These data support the hypothesis that the spatial distribution of specialist and generalist microbes does not match that seen with specialist and generalist large organisms. It may be that generalist microbes, while having a wider potential niche, are constrained, possibly by intrageneric competition, to exploit only a small part of that potential niche while specialists, with far fewer constraints to their niche, are more capable of filling their potential niche more effectively, perhaps by avoiding intrageneric competition. We suggest that these counter-intuitive distribution patterns may be a common feature of microbes in general and represent a distinct microbial principle in ecology, which is a real challenge if we are to develop a truly inclusive ecology.     

Host specialist clownfishes are environmental niche generalists

Why generalist and specialist species coexist in nature is a question that has interested evolutionary biologists for a long time. While the coexistence of specialists and generalists exploiting resources on a single ecological dimension has been theoretically and empirically explored, biological systems with multiple resource dimensions (e.g. trophic, ecological) are less well understood. Yet, such systems may provide an alternative to the classical theory of stable evolutionary coexistence of generalist and specialist species on a single resource dimension. We explore such systems and the potential trade-offs between different resource dimensions in clownfishes. All species of this iconic clade are obligate mutualists with sea anemones yet show interspecific variation in anemone host specificity. Moreover, clownfishes developed variable environmental specialization across their distribution. In this study, we test for the existence of a relationship between host-specificity (number of anemones associated with a clownfish species) and environmental-specificity (expressed as the size of the ecological niche breadth across climatic gradients). We find a negative correlation between host range and environmental specificities in temperature, salinity and pH, probably indicating a trade-off between both types of specialization forcing species to specialize only in a single direction. Trade-offs in a multi-dimensional resource space could be a novel way of explaining the coexistence of generalist and specialists.     

Improved prediction of Canada lynx distribution through regional model transferability and data efficiency

The application of species distribution models (SDMs) to areas outside of where a model was created allows informed decisions across large spatial scales, yet transferability remains a challenge in ecological modeling. We examined how regional variation in animal‐environment relationships influenced model transferability for Canada lynx (Lynx canadensis), with an additional conservation aim of modeling lynx habitat across the northwestern United States. Simultaneously, we explored the effect of sample size from GPS data on SDM model performance and transferability. We used data from three geographically distinct Canada lynx populations in Washington (n = 17 individuals), Montana (n = 66), and Wyoming (n = 10) from 1996 to 2015. We assessed regional variation in lynx‐environment relationships between these three populations using principal components analysis (PCA). We used ensemble modeling to develop SDMs for each population and all populations combined and assessed model prediction and transferability for each model scenario using withheld data and an extensive independent dataset (n = 650). Finally, we examined GPS data efficiency by testing models created with sample sizes of 5%–100% of the original datasets. PCA results indicated some differences in environmental characteristics between populations; models created from individual populations showed differential transferability based on the populations'' similarity in PCA space. Despite population differences, a single model created from all populations performed as well, or better, than each individual population. Model performance was mostly insensitive to GPS sample size, with a plateau in predictive ability reached at ~30% of the total GPS dataset when initial sample size was large. Based on these results, we generated well‐validated spatial predictions of Canada lynx distribution across a large portion of the species'' southern range, with precipitation and temperature the primary environmental predictors in the model. We also demonstrated substantial redundancy in our large GPS dataset, with predictive performance insensitive to sample sizes above 30% of the original.     

Small cats in big trouble? Diet,activity, and habitat use of jungle cats and leopard cats in threatened dry deciduous forests,Cambodia

Dry deciduous dipterocarp forests (DDF) cover about 15%–20% of Southeast Asia and are the most threatened forest type in the region. The jungle cat (Felis chaus) is a DDF specialist that occurs only in small isolated populations in Southeast Asia. Despite being one of the rarest felids in the region, almost nothing is known about its ecology. We investigated the ecology of jungle cats and their resource partitioning with the more common leopard cats (Prionailurus bengalensis) in a DDF‐dominated landscape in Srepok Wildlife Sanctuary, Cambodia. We used camera‐trap data collected from 2009 to 2019 and DNA‐confirmed scats to determine the temporal, dietary and spatial overlap between jungle cats and leopard cats. The diet of jungle cats was relatively diverse and consisted of murids (56% biomass consumed), sciurids (15%), hares (Lepus peguensis; 12%), birds (8%), and reptiles (8%), whereas leopard cats had a narrower niche breadth and a diet dominated by smaller prey, primarily murids (73%). Nonetheless, dietary overlap was high because both felid species consumed predominantly small rodents. Both species were primarily nocturnal and had high temporal overlap. Two‐species occupancy modelling suggested jungle cats were restricted to DDF and had low occupancy, whereas leopard cats had higher occupancy and were habitat generalists. Our study confirmed that jungle cats are DDF specialists that likely persist in low numbers due to the harsh conditions of the dry season in this habitat, including annual fires and substantial decreases in small vertebrate prey. The lower occupancy and more diverse diet of jungle cats, together with the broader habitat use of leopard cats, likely facilitated the coexistence of these species. The low occupancy of jungle cats in DDF suggests that protection of large areas of DDF will be required for the long‐term conservation of this rare felid in Southeast Asia.     

When species’ ranges meet: assessing differences in habitat selection between sympatric large carnivores

Differentiation in habitat selection among sympatric species may depend on niche partitioning, species interactions, selection mechanisms and scales considered. In a mountainous area in Sweden, we explored hierarchical habitat selection in Global Positioning System-collared individuals of two sympatric large carnivore species; an obligate predator, the Eurasian lynx (Lynx lynx), and a generalist predator and scavenger, the wolverine (Gulo gulo). Although the species’ fundamental niches differ widely, their ranges overlap in this area where they share a prey base and main cause of mortality. Both lynx and wolverines selected for steep and rugged terrain in mountainous birch forest and in heaths independent of scale and available habitats. However, the selection of lynx for their preferred habitats was stronger when they were forming home ranges and they selected the same habitats within their home ranges independent of home range composition. Wolverines displayed a greater variability when selecting home ranges and habitat selection also varied with home range composition. Both species selected for habitats that promote survival through limited encounters with humans, but which also are rich in prey, and selection for these habitats was accordingly stronger in winter when human activity was high and prey density was low. We suggest that the observed differences between the species result primarily from different foraging strategies, but may also depend on differences in ranging and resting behaviour, home range size, and relative density of each species. Our results support the prediction that sympatric carnivores with otherwise diverging niches can select for the same resources when sharing main sources of food and mortality.