Divergent estimates of herd‐wide caribou calf survival: Ecological factors and methodological biases

Population monitoring is a critical part of effective wildlife management, but methods are prone to biases that can hinder our ability to accurately track changes in populations through time. Calf survival plays an important role in ungulate population dynamics and can be monitored using telemetry and herd composition surveys. These methods, however, are susceptible to unrepresentative sampling and violations of the assumption of equal detectability, respectively. Here, we capitalized on 55 herd‐wide estimates of woodland caribou (Rangifer tarandus caribou) calf survival in Newfoundland, Canada, using telemetry (n = 1,175 calves) and 249 herd‐wide estimates of calf:cow ratios (C:C) using herd composition surveys to investigate these potential biases. These data included 17 herd‐wide estimates replicated from both methods concurrently (n = 448 calves and n = 17 surveys) which we used to understand which processes and sampling biases contributed to disagreement between estimates of herd‐wide calf survival. We used Cox proportional hazards models to determine whether estimates of calf mortality risk were biased by the date a calf was collared. We also used linear mixed‐effects models to determine whether estimates of C:C ratios were biased by survey date and herd size. We found that calves collared later in the calving season had a higher mortality risk and that C:C tended to be higher for surveys conducted later in the autumn. When we used these relationships to modify estimates of herd‐wide calf survival derived from telemetry and herd composition surveys concurrently, we found that formerly disparate estimates of woodland caribou calf survival now overlapped (within a 95% confidence interval) in a majority of cases. Our case study highlights the potential of under‐appreciated biases to impact our understanding of population dynamics and suggests ways that managers can limit the influence of these biases in the two widely applied methods for estimating herd‐wide survival.     

Extensive Predator Space Use Can Limit the Efficacy of a Control Program

ABSTRACT Reduced to small isolated groups by anthropogenic habitat losses or habitat modifications, populations of many endangered species are sensitive to additive sources of mortality, such as predation. Predator control is often one of the first measures considered when predators threaten survival of a population. Unfortunately, predator ecology is often overlooked because relevant data are difficult to obtain. For example, the endangered Gaspésie caribou (Rangifer tarandus caribou) has benefited from 2 periods of predator control that targeted black bears (Ursus americanus) and coyotes (Canis latrans) in an attempt to reduce predation on caribou calves. Despite a high trapping effort, the number of predators removed has remained stable over time. To assess impact of predator movements on efficacy of a control program, we studied space use of 24 black bears and 16 coyotes over 3 years in and around the Gaspésie Conservation Park, Quebec, Canada, using Global Positioning System radiocollars. Annual home ranges of black bears averaged 260 km² and 10 individuals frequented area used by caribou. Annual home ranges of resident coyotes averaged 121 km², whereas dispersing coyotes covered >2,600 km². Coyotes were generally located at lower altitudes than caribou. However, because coyotes undertook long-distance excursions, they overlapped areas used by caribou. Simulations based on observed patterns showed that 314 bears and 102 coyotes potentially shared part of their home range with areas used by female caribou during the calving period. Despite low densities of both predator species, extensive movement and use of nonexclusive territories seem to allow predators to rapidly occupy removal areas, demonstrating the need for recurrent predator removals. Our results underscore the necessity of considering complementary and alternative solutions to predator control to assure long-term protection of endangered species.     

Calving rate,calf survival rate,and habitat selection of forest-dwelling caribou in a highly managed landscape

Logging negatively affects the threatened forest-dwelling caribou (Rangifer tarandus caribou) through its positive effects on large predator populations. As recruitment is a key component of caribou population growth rate, we assessed calving rates of females and calf survival rates during the most critical period for calf survival, the calving period. We also identified causes of calf mortality and investigated the influence of predation risk, food availability, and human disturbance on habitat selection of females during the calving period at both the home-range and forest stand scales. We hypothesized that caribou should display habitat selection patterns to reduce predation risk at both scales. Using telemetry, we followed 22 females and their calves from 2004 to 2007 in a highly managed study area in Québec, Canada. Most females (78.5 ± 0.05 [SE]) gave birth each year, but only 46.3 ± 8.0% of the calves survived during the first 50 days following birth, and 57.3 ± 14.9% of them died from black bear (Ursus americanus) predation. At the home-range scale, caribou selected calving areas located at upper slope positions and avoided high road density areas. Surprisingly, they also selected the forested habitat type having the lowest lateral cover (mixed and deciduous stands) while avoiding the highest cover (regenerating conifer stands). At the forest stand scale, caribou selected areas located at relatively high elevations and with a lower basal area of black spruce trees. The selection of upper slope positions likely favored spatial segregation between calving females and wolves (Canis lupus) but not black bear. Our results suggest that calving females used areas from which they could visually detect approaching predators. While wolf avoidance appeared to be effective in a highly managed landscape, caribou did not appear to have adjusted their predator avoidance strategy to the recent increase in black bear abundance, who have benefited from increased food abundance. This situation requires focused attention from wildlife managers as logging activities are progressing towards the north within the core of forest-dwelling caribou range. © 2011 The Wildlife Society.     

Woodland caribou calf mortality in Newfoundland: insights into the role of climate,predation and population density over three decades of study

The rates and causes of juvenile mortality are central features of the dynamics and conservation of large mammals, like woodland caribou (Rangifer tarandus caribou (Gmelin, 1788)), but intrinsic and extrinsic factors may be modified by variations in animal abundance. We tested the influences of population size, climate, calf weight and sex on survival to 6 months of age of 1241 radio-collared caribou calves over three decades, spanning periods of population growth (1979–1997) and decline (2003–2012) in Newfoundland, Canada. Daily survival rates were higher and rose more quickly with calf age during the population growth period compared to the decline. Population size (negatively) and calf weight (positively) affected survival during the decline but neither had a detectable influence during the growth phase. Sex, climate and plant productivity (the latter two derived from the North Atlantic Oscillation and Normalized Difference Vegetation Index, respectively) exerted minimal influence during either phase. Predation was the dominant source of mortality. The mean percentage of calves killed by predators was 30 % higher during the decline compared to the growth phase. Black bears (Ursus americanus) and lynx (Lynx canadensis) were the major predators during the population increase but this changed during the decrease to black bears and coyotes (Canis latrans). Our findings are consistent with the hypothesis that Newfoundland caribou experienced phase-dependent survival mediated proximally by predation and competition for food.     

Fire,grazing history,lichen abundance,and winter distribution of caribou in Alaska's taiga

In the early 1990s the Nelchina Caribou (Rangifer tarandus) Herd (NCH) began a dramatic shift to its current winter range, migrating at least an additional 100 km beyond its historic range. We evaluated the impacts of fire and grazing history on lichen abundance and subsequent use and distribution by the NCH. Historic (prior to 1990) and current (2002) winter ranges of the NCH had similar vascular vegetation, lichen cover (P = 0.491), and fire histories (P = 0.535), but the former range had significantly less forage lichen biomass as a result of grazing by caribou. Biomass of forage lichens was twice as great overall (P = 0.031) and 4 times greater in caribou selected sites on the current range than in the historic range, greatly increasing availability to caribou. Caribou on the current range selected for stands with >20% lichen cover (P < 0.001), greater than 1,250 kg/ha (P < 0.001) forage lichen biomass and stands older than 80 yr postfire (P < 0.001). After fires, forage lichen cover and biomass seldom recovered sufficiently to attract caribou grazing until after ≥60 yr, and, as a group, primary forage lichen species did not reach maximum abundance until 180 yr postfire. Recovery following overgrazing can occur much more quickly because lichen cover, albeit mostly fragments, and organic substrates remain present. Our results provide benchmarks for wildlife managers assessing condition of caribou winter range and predicting effects of fires on lichen abundance and caribou distribution. Of our measurements of cover and biomass by species, densities and heights of trees, elevation, slope and aspect, only percentage cover by Cladonia amaurocraea, Cladina rangiferina, Flavocetraria cuculata, and lowbush cranberry (Vaccinium vitis-idaea) were necessary for predicting caribou use of winter range. © 2011 The Wildlife Society     

Predator densities and white-tailed deer fawn survival

Predation is the dominant source of mortality for white-tailed deer (Odocoileus virginianus) <6 months old throughout North America. Yet, few white-tailed deer fawn survival studies have occurred in areas with 4 predator species or have considered concurrent densities of deer and predator species. We monitored survival and cause-specific mortality from birth to 6 months for 100 neonatal fawns during 2013–2015 in the Upper Peninsula of Michigan, USA, while simultaneously estimating population densities of deer, American black bear (Ursus americanus), coyote (Canis latrans), bobcat (Lynx rufus), and gray wolf (Canis lupus). We estimated fawn predation risk in response to sex, birth mass, and date of birth. Six-month fawn survival pooled among years was 36%, and fawn mortality risk was not related to birth mass, date of birth, or sex. Estimated mean annual deer and predator densities were 334 fawns/100 km², 25.9 black bear/100 km², 23.8 coyotes/100 km², 3.8 bobcat/100 km², and 2.8 wolves/100 km². Despite lower estimated per-individual kill rates, coyotes and black bears were the leading sources of fawn mortality because they had greater densities relative to bobcats and wolves. Our results indicate that the presence of more predator species in a system is not entirely additive in its effect on fawn survival. © The Wildlife Society, 2019     

No long-term effect of black bear removal on elk calf recruitment in the southern Appalachians

In 2001 and 2002, 52 elk (Cervus canadensis; 21 males, 31 females), originally obtained from Elk Island National Park, Alberta, Canada, were transported and released into Cataloochee Valley in the northeastern portion of Great Smoky Mountains National Park (GRSM, Park), North Carolina, USA. The annual population growth rate (λ) was negative (0.996, 95% CI = 0.945–1.047) and predation by black bears (Ursus americanus) on elk calves was identified as an important determinant of population growth. From 2006 to 2008, 49 bears from the primary elk calving area (i.e., Cataloochee Valley) were trapped and translocated about 70 km to the southwestern portion of the Park just prior to elk calving. Per capita recruitment (i.e., the number of calves produced per adult female that survive to 1 year of age) increased from 0.306 prior to bear translocation (2001–2005) to 0.544 during years when bears were translocated (2006–2008) and λ increased to 1.118 (95% CI = 1.096–1.140). Our objective was to determine whether per capita calf recruitment rates after bear removal (2009–2019) at Cataloochee were similar to the higher rates estimated during bear removal (i.e., long-term response) or if they returned to rates before bear removal (i.e., short-term response), and how those rates compared with recruitment from portions of our study area where bears were not relocated. We documented 419 potential elk calving events and monitored 129 yearling and adult elk from 2001 to 2019. Known-fate models based on radio-telemetry and observational data supported calf recruitment returning to pre-2006 levels at Cataloochee (short-term response); recruitment of Cataloochee elk before and after bear relocation was lower (0.184) than during bear relocation (0.492). Recruitment rates of elk outside the removal area during the bear relocation period (0.478) were similar to before and after rates (0.420). In the Cataloochee Valley, cause-specific annual calf mortality rates due to predation by bears were 0.319 before, 0.120 during, and 0.306 after bear relocation. In contrast, the cause-specific annual mortality rate of calves in areas where bears were not relocated was 0.033 after the bear relocation period, with no bear predation on calves before or during bear relocation. The mean annual population growth rate for all monitored elk was 1.062 (95% CI = 0.979–1.140) after bear relocation based on the recruitment and survival data. Even though the effects of bear removal were temporary, the relocations were effective in achieving a short-term increase in elk recruitment, which was important for the reintroduction program given that the elk population was small and vulnerable to extirpation.     

Integrative use of spatial, genetic, and demographic analyses for investigating genetic connectivity between migratory, montane, and sedentary caribou herds

Genetic differentiation is generally assumed to be low in highly mobile species, but this simplistic view may obscure the complex conditions and mechanisms allowing genetic exchanges between specific populations. Here, we combined data from satellite-tracked migratory caribou (Rangifer tarandus), microsatellite markers, and demographic simulations to investigate gene flow mechanisms between seven caribou herds of eastern Canada. Our study included one montane, two migratory, and four sedentary herds. Satellite-tracking data indicated possibilities of high gene flow between migratory herds: overlap of their rutting ranges averaged 10% across years and 9.4% of females switched calving sites at least once in their lifetime. Some migratory individuals moved into the range of the sedentary herds, suggesting possibilities of gene flow between these herds. Genetic differentiation between herds was weak but significant (FST=0.015): migratory and montane herds were not significantly distinct (FST all9) than vice-versa (4Nm all<5), which suggests migratory herds had a demographic impact on sedentary herds. Demographic simulations showed that an effective immigration rate of 0.0005 was sufficient to obtain the empirical FST of 0.015, while a null immigration rate increased the simulated FST to >0.6. In conclusion, the weak genetic differentiation between herds cannot be obtained without some genetic exchanges among herds, as demonstrated by genetic and spatial data.     

Effects of black bear relocation on elk calf recruitment at Great Smoky Mountains National Park

Previous research from 2001 to 2006 on an experimentally released elk (Cervus elaphus) population at Great Smoky Mountains National Park (GSMNP or Park) indicated that calf recruitment (i.e., calves reaching 1 yr of age per adult female elk) was low (0.306, total SE = 0.090) resulting in low or negative population growth (λ = 0.996, 95% CI = 0.945–1.047). Black bear (Ursus americanus) predation was the primary calf mortality factor. From 2006 to 2008, we trapped and relocated 49 bears (30 of which were radiocollared) from the primary calving areas in the Park and radiomonitored 67 (28 M:39 F) adult elk and 42 calves to compare vital rates and population growth with the earlier study. A model with annual calf recruitment rate correlating with the number of bears relocated each year was supported (ΔAIC_c = 0.000; β = 0.070, 95% CI = 0.028–0.112) and a model with annual calf recruitment differing from before to during bear relocation revealed an increase to 0.544 (total SE = 0.098; β = −1.092, 95% CI = −1.180 to −0.375). Using vital rates and estimates of process standard errors observed during our study, 25-yr simulations maintained a mean positive growth rate in 100% of the stochastic trials with λ averaging 1.118 (95% CI = 1.096–1.140), an increase compared with rates before bear relocation. A life table response experiment revealed that increases in population growth were mostly (67.1%) due to changes in calf recruitment. We speculate that behavioral adaptation of the elk since release also contributed to the observed increases in recruitment and population growth. Our results suggest that managers interested in elk reintroduction within bear range should consider bear relocation as a temporary means of increasing calf recruitment. © 2011 The Wildlife Society.     

Longevity and reuse of black bear dens in managed forests of coastal British Columbia

We evaluated longevity and reuse of denning structures by American black bears (Ursus americanus) in coastal temperate rainforests of British Columbia from 1992 to 2010 to assess potential impacts of forest management on these critical habitat features over time. We identified 67 dens during a 4-yr intensive radio-telemetry study (1992–1995): 40 dens of 21 radio-collared black bears and 27 dens found incidentally. Dens occurred in or beneath large diameter trees or wooden structures derived from trees (i.e., logs, root boles, and stumps; = 143 cm diameter, SD = 49 cm). Longevity of dens varied by type, tree species that formed the den, and forest management that occurred at or near the den. Twenty-four of 28 dens of radio-collared bears that were monitored were still usable in 2010, whereas only 5 of 14 dens found incidentally were still usable in 2010. We assessed reuse of bear dens 3 times following initial identification: during the radio-telemetry study, again in 2000, and finally in 2010. Radio-collared bears reused dens from previous years on 7 of 25 potential occasions during the course of the radio-telemetry study. Upon assessment in 2000 and 2010, 17 of 24 (71%) available dens first used by radio-collared bears were reused at least once between 1993 and 2010. The high rate of reuse may indicate low availability of den structures in our study area. Because black bears in coastal British Columbia only used trees or structures derived from trees for winter dens and forest harvesting reduces the supply of these necessary structures, conservation and recruitment of suitable den trees is necessary if maintaining black bear populations is a management goal in these areas. © 2011 The Wildlife Society.     

Inferring parturition and neonate survival from movement patterns of female ungulates: a case study using woodland caribou

Analyses of animal movement data have primarily focused on understanding patterns of space use and the behavioural processes driving them. Here, we analyzed animal movement data to infer components of individual fitness, specifically parturition and neonate survival. We predicted that parturition and neonate loss events could be identified by sudden and marked changes in female movement patterns. Using GPS radio‐telemetry data from female woodland caribou (Rangifer tarandus caribou), we developed and tested two novel movement‐based methods for inferring parturition and neonate survival. The first method estimated movement thresholds indicative of parturition and neonate loss from population‐level data then applied these thresholds in a moving‐window analysis on individual time‐series data. The second method used an individual‐based approach that discriminated among three a priori models representing the movement patterns of non‐parturient females, females with surviving offspring, and females losing offspring. The models assumed that step lengths (the distance between successive GPS locations) were exponentially distributed and that abrupt changes in the scale parameter of the exponential distribution were indicative of parturition and offspring loss. Both methods predicted parturition with near certainty (>97% accuracy) and produced appropriate predictions of parturition dates. Prediction of neonate survival was affected by data quality for both methods; however, when using high quality data (i.e., with few missing GPS locations), the individual‐based method performed better, predicting neonate survival status with an accuracy rate of 87%. Understanding ungulate population dynamics often requires estimates of parturition and neonate survival rates. With GPS radio‐collars increasingly being used in research and management of ungulates, our movement‐based methods represent a viable approach for estimating rates of both parameters.     

Impacts of spruce budworm defoliation on the habitat of woodland caribou,moose, and their main predators

Forest logging has contributed to the decline of several woodland caribou populations by causing the fragmentation of mature coniferous stands. Such habitat alterations could be worsened by spruce budworm (SBW) outbreaks. Using 6201 vegetation plots from provincial inventories conducted after the last SBW outbreak (1968–1992) in boreal forests of Québec (Canada), we investigated the influence of SBW‐caused tree defoliation and mortality on understory vegetation layers relevant to woodland caribou and its main predators. We found a positive association between severe outbreaks and the cover of most groups of understory plant species, especially in stands that were dominated by balsam fir before the outbreak, where a high canopy openness particularly benefited relatively fast‐growing deciduous plants. Such increases in early successional vegetation could provide high‐quality forage for moose, which is likely to promote higher wolf densities and increase predation pressure on caribou. SBW outbreaks may thus negatively affect woodland caribou by increasing predation risk, the main factor limiting caribou populations in managed forests. For the near future, we recommend updating the criteria used to define critical caribou habitat to consider the potential impacts of spruce budworm defoliation.     

Variation in life history and demography of the American black bear

Variation in life history and demography across a species' range informs researchers about regional adaptations and affects whether managers can borrow information from other populations in decision-making. The American black bear (Ursus americanus) is a long-lived game species whose continued persistence depends on management of harvest and removal of habituated bears that come into conflict with humans. Understanding the demography of black bears guides efforts at management and conservation, yet detailed knowledge of many populations is typically lacking. I performed a hierarchical Bayesian meta-analysis of black bear demographic studies across the geographic range of the species to explore how vital rates vary across the range, what information they give us about population growth, and whether managers can justify borrowing information from other studies to inform management decisions. Cub, yearling, and adult survival and fecundity varied between eastern and western North America, whereas subadult survival did not show geographic structuring. Adult survival and fecundity appeared to trade off, with higher survival in the western portions of bears' range and higher fecundity in the east. Although adult survival had the highest elasticity, differences in reproduction drove differences in population growth rate. Mean population growth rate was higher in the east (0.99; 95% credible interval [CrI]: 0.96, 1.03) than the west (0.97; 95%CrI: 0.93, 1.01). Despite declining trends in the west, 34% of the distribution of population growth rate was >1, compared to 55% in the east. Further work needs to be done to address the cause of the apparent trade-off between adult survival and fecundity and explore how the estimated growth rates are likely to affect population status of black bears. Because population growth rates are close to 1 and small deviations could impact whether a population is considered increasing or decreasing, managers need to employ caution in borrowing vital rates from other populations. © 2011 The Wildlife Society.