Nathaniel D. Rayl

Phase‐dependent climate–predator interactions explain three decades of variation in neonatal caribou survival

Climate can have direct and indirect effects on population dynamics via changes in resource competition or predation risk, but this influence may be modulated by density- or phase-dependent processes. We hypothesized that for ungulates, climatic conditions close to parturition have a greater influence on the predation risk of neonates during population declines, when females are already under nutritional stress triggered by food limitation. We examined the presence of phase-dependent climate-predator (PDCP) interactions on neonatal ungulate survival by comparing spatial and temporal fluctuations in climatic conditions, cause-specific mortality and per capita resource limitation. We determined cause-specific fates of 1384 caribou (Rangifer tarandus) from 10 herds in Newfoundland, spanning more than 30 years during periods of numerical increase and decline, while exposed to predation from black bears (Ursus americanus) and coyotes (Canis latrans). We conducted Cox proportional hazards analysis for competing risks, fit as a function of weather metrics, to assess pre- and post-partum climatic influences on survival on herds in population increase and decline phases. We used cumulative incidence functions to compare temporal changes in risk from predators. Our results support our main hypothesis; when caribou populations increased, weather conditions preceding calving were the main determinants of cause-specific mortality, but when populations declined, weather conditions during calving also influenced predator-driven mortality. Cause-specific analysis showed that weather conditions can differentially affect predation risk between black bears and coyotes with specific variables increasing the risk from one species and decreasing the risk from the other. For caribou, nutritional stress appears to increase predation risk on neonates, an interaction which is exacerbated by susceptibility to climatic events. These findings support the PDCP interactions framework, where maternal body condition influences susceptibility to climate-related events and, subsequently, risk from predation.

Mapping the distribution of a prey resource: neonate caribou in Newfoundland

Abstract The recently diminished caribou (Rangifer tarandus) population in insular Newfoundland, Canada, has been severely limited by predation on newborn calves. These neonates are patchily distributed on the landscape; therefore, to adequately understand predator–prey interactions, the temporal and geographic extent of the distribution of caribou calves must be identified so that current areas with calves can be differentiated from areas without calves. We used telemetry locations of 309 caribou calves and 100 adult females from 4 herds, 2008–2010, to estimate the spatiotemporal distribution of caribou calves during the time when they were most vulnerable to predation, to evaluate the predictability of the calf resource among years, and to assess the degree of aggregation during calving. Patterns of calf distribution were predictable in time and space from year to year, with an average distributional overlap of 68% between years. The dispersion of female caribou during calving varied among herds from highly aggregated (8% and 20% of herd range) to more dispersed (50–70% of herd range). Postcalving (up to 9 weeks) distributions also varied among herds; the 2 more-dispersed herds remained sedentary, whereas both highly aggregated herds migrated away from their calving grounds at the end of June. The most-aggregated herd remained so as it migrated from its calving ground, whereas the less-aggregated herd spread out and moved in a variety of predictable directions. Dispersion and movement patterns varied with forest cover; herds with less forest cover in their range were more aggregated and migratory than herds with more forest cover.

Spatial and Temporal Habitat Use of an Asian Elephant in Sumatra.

Simple Summary A wild Sumatran elephant radio-monitored near a conservation center from August 2007–May 2008 used medium- and open-canopy land cover more than expected, but closed canopy forests were used more during the day than at night. When in closed canopy forests, elephants spent more time near the forest edge. Effective elephant conservation strategies in Sumatra need to focus on forest restoration of cleared areas and providing a forest matrix that includes various canopy types. Abstract Increasingly, habitat fragmentation caused by agricultural and human development has forced Sumatran elephants into relatively small areas, but there is little information on how elephants use these areas and thus, how habitats can be managed to sustain elephants in the future. Using a Global Positioning System (GPS) collar and a land cover map developed from TM imagery, we identified the habitats used by a wild adult female elephant (Elephas maximus sumatranus) in the Seblat Elephant Conservation Center, Bengkulu Province, Sumatra during 2007–2008. The marked elephant (and presumably her 40–60 herd mates) used a home range that contained more than expected medium canopy and open canopy land cover. Further, within the home range, closed canopy forests were used more during the day than at night. When elephants were in closed canopy forests they were most often near the forest edge vs. in the forest interior. Effective elephant conservation strategies in Sumatra need to focus on forest restoration of cleared areas and providing a forest matrix that includes various canopy types.

Climate change can alter predator–prey dynamics and population viability of prey

For many organisms, climate change can directly drive population declines, but it is less clear how such variation may influence populations indirectly through modified biotic interactions. For instance, how will climate change alter complex, multi-species relationships that are modulated by climatic variation and that underlie ecosystem-level processes? Caribou (Rangifer tarandus), a keystone species in Newfoundland, Canada, provides a useful model for unravelling potential and complex long-term implications of climate change on biotic interactions and population change. We measured cause-specific caribou calf predation (1990–2013) in Newfoundland relative to seasonal weather patterns. We show that black bear (Ursus americanus) predation is facilitated by time-lagged higher summer growing degree days, whereas coyote (Canis latrans) predation increases with current precipitation and winter temperature. Based on future climate forecasts for the region, we illustrate that, through time, coyote predation on caribou calves could become increasingly important, whereas the influence of black bear would remain unchanged. From these predictions, demographic projections for caribou suggest long-term population limitation specifically through indirect effects of climate change on calf predation rates by coyotes. While our work assumes limited impact of climate change on other processes, it illustrates the range of impact that climate change can have on predator–prey interactions. We conclude that future efforts to predict potential effects of climate change on populations and ecosystems should include assessment of both direct and indirect effects, including climate–predator interactions.

Den abandonment and transitional day bed use by black bears Ursus americanus in Newfoundland

The use of day beds for extended periods during the transition into and out of the physiological state of hibernation has been documented in many bear populations, but has never been quantifi ed. Additionally, den abandonment by black bears Ursus americanus has rarely been observed at northern latitudes except after den visits by researchers. In three areas on the northern island of Newfoundland, where male and female black bears spent an average of 158 and 178 d denning, respectively, we identified den sites and extended-use day beds (occupied continuously for 6–26 d) remotely using GPS collars, and here provide the first systematic description of the use of these day beds by bears. We documented den abandonment in 6 (9%; 3 F, 3 M) of 67 bear-winters (6 [14%] of 44 radio-collared bears) and the use of extended-use transitional day beds in 16 (24%) of 67 bear-winters (15 [34%] of 44 radio-collared bears, 8 F, 7 M). In 5 of 10 instances bears left their fall day beds on days with > 15 mm of rain (mean = 28.2 mm, range = 15.6–63.6 mm), which was more than would be expected by chance (p < 0.01). We had more than one year of denning data for 17 bears, 6 (35%) of which reused den sites in diff erent years. Further, we observed some bears using day bed and den sites interchangeably. Though we hypothesized that environmental (flooding) or anthropogenic disturbance (researcher-, forestry-related, or recreational) may have played a role in den abandonment, we found no such relationships, nor was there a difference in the rate of abandonment or day bed use between male and female bears. We could not assess the eff ects of microhabitat attributes, condition, or reproductive status, but acknowledge that these factors may have played a role in den changes.

Spatiotemporal heterogeneity in prey abundance and vulnerability shapes the foraging tactics of an omnivore

Prey abundance and prey vulnerability vary across space and time, but we know little about how they mediate predator-prey interactions and predator foraging tactics. To evaluate the interplay between prey abundance, prey vulnerability and predator space use, we examined patterns of black bear (Ursus americanus) predation of caribou (Rangifer tarandus) neonates in Newfoundland, Canada using data from 317 collared individuals (9 bears, 34 adult female caribou, 274 caribou calves). During the caribou calving season, we predicted that landscape features would influence calf vulnerability to bear predation, and that bears would actively hunt calves by selecting areas associated with increased calf vulnerability. Further, we hypothesized that bears would dynamically adjust their foraging tactics in response to spatiotemporal changes in calf abundance and vulnerability (collectively, calf availability). Accordingly, we expected bears to actively hunt calves when they were most abundant and vulnerable, but switch to foraging on other resources as calf availability declined. As predicted, landscape heterogeneity influenced risk of mortality, and bears displayed the strongest selection for areas where they were most likely to kill calves, which suggested they were actively hunting caribou. Initially, the per-capita rate at which bears killed calves followed a type-I functional response, but as the calving season progressed and calf vulnerability declined, kill rates dissociated from calf abundance. In support of our hypothesis, bears adjusted their foraging tactics when they were less efficient at catching calves, highlighting the influence that predation phenology may have on predator space use. Contrary to our expectations, however, bears appeared to continue to hunt caribou as calf availability declined, but switched from a tactic of selecting areas of increased calf vulnerability to a tactic that maximized encounter rates with calves. Our results reveal that generalist predators can dynamically adjust their foraging tactics over short time-scales in response to changing prey abundance and vulnerability. Further, they demonstrate the utility of integrating temporal dynamics of prey availability into investigations of predator-prey interactions, and move towards a mechanistic understanding of the dynamic foraging tactics of a large omnivore.

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.