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The cost of reproduction: differential resource specialization in female and male California sea otters

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 (δ 13 C) and nitrogen (δ 15 N) 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 isotopic 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.

California

Timescales alter the inferred strength and temporal consistency of intraspecific diet specialization

Many populations consist of individuals that differ substantially in their diets. Quantification of the magnitude and temporal consistency of such intraspecific diet variation is needed to understand its importance, but the extent to which different approaches for doing so reflect instantaneous vs. time-aggregated measures of individual diets may bias inferences. We used direct observations of sea otter individuals ( Enhydra lutris nereis ) to assess how: (1) the timescale of sampling, (2) under-sampling, and (3) the incidence- vs. frequency-based consideration of prey species affect the inferred strength and consistency of intraspecific diet variation. Analyses of feeding observations aggregated over hourly to annual intervals revealed a substantial bias associated with time aggregation that decreases the inferred magnitude of specialization and increases the inferred consistency of individuals’ diets. Time aggregation also made estimates of specialization more sensitive to the consideration of prey frequency, which decreased estimates relative to the use of prey incidence; time aggregation did not affect the extent to which under-sampling contributed to its overestimation. Our analyses demonstrate the importance of studying intraspecific diet variation with an explicit consideration of time and thereby suggest guidelines for future empirical efforts. Failure to consider time will likely produce inconsistent predictions regarding the effects of intraspecific variation on predator–prey interactions.

Oecologia

The interaction of intraspecific competition and habitat on individual diet specialization: a near range-wide examination of sea otters

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 population- and 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 isotopic 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 context-dependent 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.

Oecologia

Climatic controls of aboveground net primary production in semi-arid grasslands along a latitudinal gradient portend low sensitivity to warming

Although climate models forecast warmer temperatures with a high degree of certainty, precipitation is the primary driver of aboveground net primary production (ANPP) in most grasslands. Conversely, variations in temperature seldom are related to patterns of ANPP. Thus forecasting responses to warming is a challenge, and raises the question: how sensitive will grassland ANPP be to warming? We evaluated climate and multi-year ANPP data (67 years) from eight western US grasslands arrayed along mean annual temperature (MAT; ~7-14 °C) and mean annual precipitation (MAP; ~250-500 mm) gradients. Weused regression and analysis of covariance to assess relationships between ANPP and temperature, as well as precipitation (annual and growing season) to evaluate temperature sensitivity of ANPP. We also related ANPP to the standardized precipitation evaporation index (SPEI), which combines precipitation and evapotranspiration to better represent moisture available for plant growth. Regression models indicated that variation in growing season temperature was negatively related to total and graminoid ANPP, but precipitation was a stronger predictor than temperature. Growing season temperature was also a significant parameter in more complex models, but again precipitation was consistently a stronger predictor of ANPP. Surprisingly, neither annual nor growing season SPEI were as strongly related to ANPP as precipitation. We conclude that forecasted warming likely will affect ANPP in these grasslands, but that predicting temperature effects from natural climatic gradients is difficult. This is because, unlike precipitation, warming effects can be positive or negative and moderated by shifts in the C3/C4 ratios of plant communities.

Oecologia

C 3 and C 4 plant responses to increased temperatures and altered monsoonal precipitation in a cool desert on the Colorado Plateau, USA

Dryland ecosystems represent >40 % of the terrestrial landscape and support over two billion people; consequently, it is vital to understand how drylands will respond to climatic change. However, while arid and semiarid ecosystems commonly experience extremely hot and dry conditions, our understanding of how further temperature increases or altered precipitation will affect dryland plant communities remains poor. To address this question, we assessed plant physiology and growth at a long-term (7-year) climate experiment on the Colorado Plateau , USA, where the community is a mix of shallow-rooted C 3 and C 4 grasses and deep-rooted C 4 shrubs. The experiment maintained elevated-temperature treatments (+2 or +4 °C) in combination with altered summer monsoonal precipitation (+small frequent precipitation events or +large infrequent events). Increased temperature negatively affected photosynthesis and growth of the C 3 and C 4 grasses, but effects varied in their timing: +4 °C treatments negatively affected the C 3 grass early in the growing season of both years, while the negative effects of temperature on the C 4 grass were seen in the +2 and +4 °C treatments, but only during the late growing season of the drier year. Increased summer precipitation did not affect photosynthesis or biomass for any species, either in the year the precipitation was applied or the following year. Although previous research suggests dryland plants, and C 4 grasses in particular, may respond positively to elevated temperature, our findings from a cool desert show marked declines in C 3 and C 4 photosynthesis and growth, with temperature effects dependent on the degree of warming and growing-season precipitation.

Utah

Individual specialization in the foraging habits of female bottlenose dolphins living in a trophically diverse and habitat rich estuary

We examine individual specialization in foraging habits (foraging habitat and trophic level) of female bottlenose dolphins ( Tursiops truncatus ) resident in Sarasota Bay, Florida, USA, by analyzing time series of stable isotope (δ 15 N and δ 13 C) values in sequential growth layer groups within teeth. The isotope data provide a chronology of foraging habits over the lifetime of the individual and allowed us to show that female bottlenose dolphins exhibit a high degree of individual specialization in both foraging habitat and trophic level. The foraging habits used by adult females are similar to those they used as calves and may be passed down from mother to calf through social learning. We also characterized the foraging habits and home range of each individual by constructing standard ellipses from isotope values and dolphin sightings data (latitude and longitude), respectively. These data show that Sarasota Bay bottlenose dolphins forage within a subset of the habitats in which they are observed. Moreover, females with similar observational standard ellipses often possessed different foraging specializations. Female bottlenose dolphins may demonstrate individual specialization in foraging habits because it reduces some of the cost of living in groups, such as competition for prey.

Florida

Linking degradation status with ecosystem vulnerability to environmental change

Environmental change can cause regime shifts in ecosystems, potentially threatening ecosystem services. It is unclear if the degradation status of ecosystems correlates with their vulnerability to environmental change, and thus the risk of future regime shifts. We assessed resilience in acidified (degraded) and circumneutral (undegraded) lakes with long-term data (1988–2012), using time series modeling. We identified temporal frequencies in invertebrate assemblages, which identifies groups of species whose population dynamics vary at particular temporal scales. We also assessed species with stochastic dynamics, those whose population dynamics vary irregularly and unpredictably over time. We determined the distribution of functional feeding groups of invertebrates within and across the temporal scales identified, and in those species with stochastic dynamics, and assessed attributes hypothesized to contribute to resilience. Three patterns of temporal dynamics, consistent across study lakes, were identified in the invertebrates. The first pattern was one of monotonic change associated with changing abiotic lake conditions. The second and third patterns appeared unrelated to the environmental changes we monitored. Acidified and the circumneutral lakes shared similar levels and patterns of functional richness, evenness, diversity, and redundancy for species within and across the observed temporal scales and for stochastic species groups. These similar resilience characteristics suggest that both lake types did not differ in vulnerability to the environmental changes observed here. Although both lake types appeared equally vulnerable in this study, our approach demonstrates how assessing systemic vulnerability by quantifying ecological resilience can help address uncertainty in predicting ecosystem responses to environmental change across ecosystems.

Oecologia

Detecting mismatches of bird migration stopover and tree phenology in response to changing climate

Migratory birds exploit seasonal variation in resources across latitudes, timing migration to coincide with the phenology of food at stopover sites. Differential responses to climate in phenology across trophic levels can result in phenological mismatch; however, detecting mismatch is sensitive to methodology. We examined patterns of migrant abundance and tree flowering, phenological mismatch, and the influence of climate during spring migration from 2009 to 2011 across five habitat types of the Madrean Sky Islands in southeastern Arizona, USA. We used two metrics to assess phenological mismatch: synchrony and overlap. We also examined whether phenological overlap declined with increasing difference in mean event date of phenophases. Migrant abundance and tree flowering generally increased with minimum spring temperature but depended on annual climate by habitat interactions. Migrant abundance was lowest and flowering was highest under cold, snowy conditions in high elevation montane conifer habitat while bird abundance was greatest and flowering was lowest in low elevation riparian habitat under the driest conditions. Phenological synchrony and overlap were unique and complementary metrics and should both be used when assessing mismatch. Overlap declined due to asynchronous phenologies but also due to reduced migrant abundance or flowering when synchrony was actually maintained. Overlap declined with increasing difference in event date and this trend was strongest in riparian areas. Montane habitat specialists may be at greatest risk of mismatch while riparian habitat could provide refugia during dry years for phenotypically plastic species. Interannual climate patterns that we observed match climate change projections for the arid southwest, altering stopover habitat condition.

Arizona

Effects of climate and plant phenology on recruitment of moose at the southern extent of their range

Climate plays a fundamental role in limiting the range of a species, is a key factor in the dynamics of large herbivores, and is thought to be involved in declines of moose populations in recent decades. We examined effects of climate and growing-season phenology on recruitment (8–9 months old) of young Shiras moose ( Alces alces shirasi ) over three decades, from 18 herds, across a large geographic area encompassing much of the southern extent of their range. Recruitment declined in 8 of 18 herds during 1980–2009, whereas others did not exhibit a temporal trend (none showed a positive trend). During those three decades, seasonal temperatures increased, spring–summer precipitation decreased, and spring occurred earlier, became shorter in duration, and green-up occurred faster. Recruitment was influenced negatively by warm temperatures during the year before young were born, but only for herds with declining recruitment. Dry spring–summers of the previous year and rapid rates of spring green-up in the year of birth had similar negative influences across declining and stable herds. Those patterns indicate both direct (year t ) and delayed (year t− 1 ) effects of weather and plant phenology on recruitment of young, which we hypothesize was mediated through effects on maternal nutritional condition. Suppressed nutrition could have been induced by (1) increased thermoregulatory costs associated with warming temperatures and (2) shortened duration of availability of high-quality forage in spring. Progressive reductions in net energetic gain for species that are sensitive to climate may continue to hamper individual fitness and population dynamics.

Oecologia

Variable nutrient stoichiometry (carbon:nitrogen:phosphorus) across trophic levels determines community and ecosystem properties in an oligotrophic mangrove system

Our study investigated the carbon:nitrogen:phosphorus (C:N:P) stoichiometry of mangrove island of the Mesoamerican Barrier Reef (Twin Cays, Belize). The C:N:P of abiotic and biotic components of this oligotrophic ecosystem was measured and served to build networks of nutrient flows for three distinct mangrove forest zones (tall seaward fringing forest, inland dwarf forests and a transitional zone). Between forest zones, the stoichiometry of primary producers, heterotrophs and abiotic components did not change significantly, but there was a significant difference in C:N:P, and C, N, and P biomass, between the functional groups mangrove trees, other primary producers, heterotrophs, and abiotic components. C:N:P decreased with increasing trophic level. Nutrient recycling in the food webs was highest for P, and high transfer efficiencies between trophic levels of P and N also indicated an overall shortage of these nutrients when compared to C. Heterotrophs were sometimes, but not always, limited by the same nutrient as the primary producers. Mangrove trees and the primary tree consumers were P limited, whereas the invertebrates consuming leaf litter and detritus were N limited. Most compartments were limited by P or N (not by C), and the relative depletion rate of food sources was fastest for P. P transfers thus constituted a bottleneck of nutrient transfer on Twin Cays. This is the first comprehensive ecosystem study of nutrient transfers in a mangrove ecosystem, illustrating some mechanisms (e.g. recycling rates, transfer efficiencies) which oligotrophic systems use in order to build up biomass and food webs spanning various trophic levels.

Twin Cays

Pulse-drought atop press-drought: unexpected plant responses and implications for dryland ecosystems

In drylands, climate change is predicted to cause chronic reductions in water availability (press-droughts) through reduced precipitation and increased temperatures as well as increase the frequency and intensity of short-term extreme droughts (pulse-droughts). These changes in precipitation patterns may have profound ecosystem effects, depending on the sensitivities of the dominant plant functional types (PFTs). Here we present the responses of four Colorado Plateau PFTs to an experimentally imposed, 4-year, press-drought during which a natural pulse-drought occurred. Our objectives were to (1) identify the drought sensitivities of the PFTs, (2) assess the additive effects of the press- and pulse-drought, and (3) examine the interactive effects of soils and drought. Our results revealed that the C 3 grasses were the most sensitive PFT to drought, the C 3 shrubs were the most resistant, and the C 4 grasses and shrubs had intermediate drought sensitivities. Although we expected the C 3 grasses would have the greatest response to drought, the higher resistance of C 3 shrubs relative to the C 4 shrubs was contrary to our predictions based on the higher water use efficiency of C 4 photosynthesis. Also, the additive effects of press- and pulse-droughts caused high morality in C 3 grasses, which has large ecological and economic ramifications for this region. Furthermore, despite predictions based on the inverse texture hypothesis, we observed no interactive effects of soils with the drought treatment on cover or mortality. These results suggest that plant responses to droughts in drylands may differ from expectations and have large ecological effects if press- and pulse-droughts push species beyond physiological and mortality thresholds.

Oecologia

Trophic dynamics of shrinking Subarctic lakes: naturally eutrophic waters impart resilience to rising nutrient and major ion concentrations

Shrinking lakes were recently observed for several Arctic and Subarctic regions due to increased evaporation and permafrost degradation. Along with lake drawdown, these processes often boost aquatic chemical concentrations, potentially impacting trophic dynamics. In particular, elevated chemical levels may impact primary productivity, which may in turn influence populations of primary and secondary consumers. We examined trophic dynamics of 18 shrinking lakes of the Yukon Flats, Alaska, that had experienced pronounced increases in nutrient (>200 % total nitrogen, >100 % total phosphorus) and ion concentrations (>100 % for four major ions combined) from 1985-1989 to 2010-2012, versus 37 stable lakes with relatively little chemical change over the same period. We found that phytoplankton stocks, as indexed by chlorophyll concentrations, remained unchanged in both shrinking and stable lakes from the 1980s to 2010s. Moving up the trophic ladder, we found significant changes in invertebrate abundance across decades, including decreased abundance of five of six groups examined. However, these decadal losses in invertebrate abundance were not limited to shrinking lakes, occurring in lakes with stable surface areas as well. At the top of the food web, we observed that probabilities of lake occupancy for ten waterbird species, including adults and chicks, remained unchanged from the period 1985-1989 to 2010-2012. Overall, our study lakes displayed a high degree of resilience to multi-trophic cascades caused by rising chemical concentrations. This resilience was likely due to their naturally high fertility, such that further nutrient inputs had little impact on waters already near peak production.

Alaska

Annual grass invasion in sagebrush-steppe: The relative importance of climate, soil properties and biotic interactions

The invasion by winter-annual grasses (AGs) such as Bromus tectorum into sagebrush steppe throughout the western USA is a classic example of a biological invasion with multiple, interacting climate, soil and biotic factors driving the invasion, although few studies have examined all components together. Across a 6000-km 2 area of the northern Great Basin, we conducted a field assessment of 100 climate, soil, and biotic (functional group abundances, diversity) factors at each of 90 sites that spanned an invasion gradient ranging from 0 to 100 % AG cover. We first determined which biotic and abiotic factors had the strongest correlative relationships with AGs and each resident functional group. We then used regression and structural equation modeling to explore how multiple ecological factors interact to influence AG abundance. Among biotic interactions, we observed negative relationships between AGs and biodiversity, perennial grass cover, resident species richness, biological soil crust cover and shrub density, whereas perennial and annual forb cover, tree cover and soil microbial biomass had no direct linkage to AG. Among abiotic factors, AG cover was strongly related to climate (increasing cover with increasing temperature and aridity), but had weak relationships with soil factors. Our structural equation model showed negative effects of perennial grasses and biodiversity on AG cover while integrating the negative effects of warmer climate and positive influence of belowground processes on resident functional groups. Our findings illustrate the relative importance of biotic interactions and climate on invasive abundance, while soil properties appear to have stronger relationships with resident biota than with invasives.

Oregon

Detecting grizzly bear use of ungulate carcasses using global positioning system telemetry and activity data

Global positioning system (GPS) wildlife collars have revolutionized wildlife research. Studies of predation by free-ranging carnivores have particularly benefited from the application of location clustering algorithms to determine when and where predation events occur. These studies have changed our understanding of large carnivore behavior, but the gains have concentrated on obligate carnivores. Facultative carnivores, such as grizzly/brown bears ( Ursus arctos ), exhibit a variety of behaviors that can lead to the formation of GPS clusters. We combined clustering techniques with field site investigations of grizzly bear GPS locations ( n = 732 site investigations; 2004–2011) to produce 174 GPS clusters where documented behavior was partitioned into five classes (large-biomass carcass, small-biomass carcass, old carcass, non-carcass activity, and resting). We used multinomial logistic regression to predict the probability of clusters belonging to each class. Two cross-validation methods—leaving out individual clusters, or leaving out individual bears—showed that correct prediction of bear visitation to large-biomass carcasses was 78–88%, whereas the false-positive rate was 18–24%. As a case study, we applied our predictive model to a GPS data set of 266 bear-years in the Greater Yellowstone Ecosystem (2002–2011) and examined trends in carcass visitation during fall hyperphagia (September–October). We identified 1997 spatial GPS clusters, of which 347 were predicted to be large-biomass carcasses. We used the clustered data to develop a carcass visitation index, which varied annually, but more than doubled during the study period. Our study demonstrates the effectiveness and utility of identifying GPS clusters associated with carcass visitation by a facultative carnivore.

Greater Yellowstone Ecosystem

Life history plasticity does not confer resilience to environmental change in the mole salamander (Ambystoma talpoideum)

Plasticity in life history strategies can be advantageous for species that occupy spatially or temporally variable environments. We examined how phenotypic plasticity influences responses of the mole salamander, Ambystoma talpoideum, to disturbance events at the St. Marks National Wildlife Refuge (SMNWR), FL, USA from 2009 to 2014. We observed periods of extensive drought early in the study, in contrast to high rainfall and expansive flooding events in later years. Flooding facilitated colonization of predatory fishes to isolated wetlands across the refuge. We employed multistate occupancy models to determine how this natural experiment influenced the occurrence of aquatic larvae and paedomorphic adults and what implications this may have for the population. We found that, in terms of occurrence, responses to environmental variation differed between larvae and paedomorphs, but plasticity (i.e. the ability to metamorphose rather than remain in aquatic environment) was not sufficient to buffer populations from declining as a result of environmental perturbations. Drought and fish presence negatively influenced occurrence dynamics of larval and paedomorphic mole salamanders and, consequently, contributed to observed short-term declines of this species. Overall occurrence of larval salamanders decreased from 0.611 in 2009 to 0.075 in 2014 and paedomorph occurrence decreased from 0.311 in 2009 to 0.121 in 2014. Although variation in selection pressures has likely maintained this polyphenism previously, our results suggest that continued changes in environmental variability and the persistence of fish in isolated wetlands could lead to a loss of paedomorphosis in the SMNWR population and, ultimately, impact regional persistence in the future.

Oecologia

Response of aboveground carbon balance to long-term, experimental enhancements in precipitation seasonality is contingent on plant community type in cold-desert rangelands

Semi-arid rangelands are important carbon (C) pools at global scales. However, the degree of net C storage or release in water-limited systems is a function of precipitation amount and timing, as well as plant community composition. In northern latitudes of western North America, C storage in cold-desert ecosystems could increase with boosts in wintertime precipitation, in which climate models predict, due to increases in wintertime soil water storage that enhance summertime productivity. However, there are few long-term, manipulative field-based studies investigating how rangelands will respond to altered precipitation amount or timing. We measured aboveground C pools and fluxes at leaf, soil, and ecosystem scales over a single growing season in plots that had 200 mm of supplemental precipitation added in either winter or summer for the past 21 years, in shrub- and exotic-bunchgrass-dominated garden plots. At our cold-desert site (298 mm precipitation during the study year), we hypothesized that increased winter precipitation would stimulate the aboveground C uptake and storage relative to ambient conditions, especially in plots containing shrubs. Our hypotheses were generally supported: ecosystem C uptake and long-term biomass accumulation were greater in winter- and summer-irrigated plots compared to control plots in both vegetation communities. However, substantial increases in the aboveground biomass occurred only in winter-irrigated plots that contained shrubs. Our findings suggest that increases in winter precipitation will enhance C storage of this widespread ecosystem, and moreso in shrub- compared to grass-dominated communities.

Idaho

Habitat degradation affects the summer activity of polar bears

Understanding behavioral responses of species to environmental change is critical to forecasting population-level effects. Although climate change is significantly impacting species’ distributions, few studies have examined associated changes in behavior. Polar bear ( Ursus maritimus ) subpopulations have varied in their near-term responses to sea ice decline. We examined behavioral responses of two adjacent subpopulations to changes in habitat availability during the annual sea ice minimum using activity data. Location and activity sensor data collected from 1989 to 2014 for 202 adult female polar bears in the Southern Beaufort Sea (SB) and Chukchi Sea (CS) subpopulations were used to compare activity in three habitat types varying in prey availability: (1) land; (2) ice over shallow, biologically productive waters; and (3) ice over deeper, less productive waters. Bears varied activity across and within habitats with the highest activity at 50–75% sea ice concentration over shallow waters. On land, SB bears exhibited variable but relatively high activity associated with the use of subsistence-harvested bowhead whale carcasses, whereas CS bears exhibited low activity consistent with minimal feeding. Both subpopulations had fewer observations in their preferred shallow-water sea ice habitats in recent years, corresponding with declines in availability of this substrate. The substantially higher use of marginal habitats by SB bears is an additional mechanism potentially explaining why this subpopulation has experienced negative effects of sea ice loss compared to the still-productive CS subpopulation. Variability in activity among, and within, habitats suggests that bears alter their behavior in response to habitat conditions, presumably in an attempt to balance prey availability with energy costs.

Oecologia

Examining spatial patterns of selection and use for an altered predator guild

Anthropogenic disturbances have altered species’ distributions potentially impacting interspecific interactions. Interference competition is when one species denies a competing species access to a resource. One mechanism of interference competition is aggression, which can result in altered space-use of a subordinate species due to the threat of harm, otherwise known as a ‘landscape of fear’. Alternatively, subordinates might outcompete dominant species in resource-poor environments via a superior ability to extract resources. Our goal was to evaluate spatial predictions of the ‘landscape of fear’ hypothesis for a carnivore guild in Newfoundland, Canada, where coyotes recently immigrated. Native Newfoundland carnivores include red foxes, Canada lynx, and black bears. We predicted foxes and lynx would avoid coyotes because of their larger size and similar dietary niches. We used scat-detecting dogs and genetic techniques to locate and identify predator scats. We then built resource selection functions and tested for avoidance by incorporating predicted values of selection for the alternative species into the best supported models of each species. We found multiple negative relationships, but notably did not find avoidance by foxes of areas selected by coyotes. While we did find that lynx avoided coyotes, we also found a reciprocal relationship. The observed patterns suggest spatial partitioning and not coyote avoidance, although avoidance could still be occurring at different spatial or temporal scales. Furthermore, Newfoundland’s harsh climate and poor soils may swing the pendulum of interspecific interactions from interference competition to exploitative competition, where subordinates outcompete dominant competitors through a superior ability to extract resources.

Newfoundland