Impacts of climate change on the seasonal distribution of migratory caribou

Arctic ecosystems are especially vulnerable to global climate change as temperature and precipitation regimes are altered. An ecologically and socially highly important northern terrestrial species that may be impacted by climate change is the caribou, Rangifer tarandus . We predicted the current and potential future occurrence of two migratory herds of caribou [Rivière George herd (RG) and Rivière-aux-Feuilles (RAF) herd] under a Canadian General Circulation Model climate change scenario, across all seasons in the Québec–Labrador peninsula, using climatic and habitat predictor variables. Argos satellite-tracking collars have been deployed on 213 caribou between 1988 and 2003 with locations recorded every 4–5 days. In addition, we assembled a database of climate (temperature, precipitation, snowfall, timing and length of growing season) and habitat data obtained from the SPOT VEGETATION satellite sensor. Logistic regression models indicated that both climatic and physical habitat variables were significant predictors of current migratory caribou occurrence. Migratory caribou appeared to prefer regions with higher snowfall and lichen availability in the fall and winter. In the summer, caribou preferred cooler areas likely corresponding to a lower prevalence of insects, and they avoided disturbed and recently burnt areas. Climate change projections using climate data predicted an increased range for the RAF herd and decreased range for the RG herd during 2040–2069, limiting the herds to northeastern regions of the Québec–Labrador peninsula. Direct and indirect consequences of climate change on these migratory caribou herds possibly include alteration in habitat use, migration patterns, foraging behaviour, and demography, in addition to social and economic stress to arctic and subarctic native human populations.     

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.     

Spatial synchrony of recruitment in mountain-dwelling woodland caribou

Spatial synchrony in population dynamics is a ubiquitous feature across a range of taxa. Understanding factors influencing this synchrony may shed light on important drivers of population dynamics. Three mechanisms influence the degree of spatial synchrony between populations: dispersal, shared predators, and spatial environmental covariance (the Moran effect). We assessed demographic spatial synchrony in recruitment (calf:cow ratio) of 10 northern mountain caribou herds in the Yukon Territory, Canada (1982–2008). Shared predators and dispersal were ruled out as causal mechanisms of spatial recruitment synchrony in these herds and therefore any spatial synchrony should be due to the Moran effect. We also assessed the degree of spatial synchrony in April snow depth to represent environmental variability. The regional average spatial synchrony in detrended residuals of April snow depth was 0.46 (95% CI 0.37 to 0.55). Spatial synchrony in caribou recruitment was weak at 0.13 (95% CI −0.06 to 0.32). The spatial scale of synchrony in April snow depth and caribou recruitment was 330.2 km (95% CI 236.3 to 370.0 km) and 170.0 km (95% CI 69.5 to 282.8 km), respectively. We also investigated how the similarity in terrain features between herds influenced the degree of spatial synchrony using exponential decay models. Only the difference in elevation variability between herds during calving was supported by the data. Herds with more similar elevation variability may track snowmelt ablation patterns in a more similar fashion, which would subsequently result in more synchronized predation rates on calves and/or nutritional effects impacting juvenile survival. Interspecific interactions with predators and alternate prey may also influence spatial synchrony of recruitment in these herds.     

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.     

A data-driven demographic model to explore the decline of the Bathurst caribou herd

The Bathurst herd of barren-ground caribou (Rangifer tarandus groenlandicus) in the Canadian central arctic declined from an estimated 203,800 to 16,400 breeding females from 1986 to 2009, with the most rapid decline from 2006 to 2009. A key research and management question was whether the decline was mainly due to decreases in productivity alone or also due to reduced adult female survival. Investigating causes of the decline was hampered by a lack of direct estimates of caribou demographic parameters. We developed a demographic model that could be objectively fitted to field data to explore the mechanisms for the Bathurst decline, with a focus on the recent accelerated decline from 2006 to 2009. Our modeling indicated that the decline was driven by increasing negative trends in adult female and calf survival rates and possibly reduced fecundity The effect of a constant hunter harvest on the declining herd was one potential cause for the recent accelerated decline in adult survival. The demographic model detected negative trends in adult female survival that were not detected using standalone analyses of collar-based survival data. The model allowed rigorous interpretation of trends in productivity by controlling for the simultaneous influence of trends in adult, calf, and yearling survival and adult fecundity on field-based calf–cow ratios. Stochastic simulations suggested that large increases in adult survival and productivity would be needed for the herd to recover. Our methods enable objective modeling of caribou demography that can assist in caribou management based upon all sources of available data. © 2011 The Wildlife Society.     

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     

Prioritization of landscape connectivity for the conservation of Peary caribou

Adequate connectivity between discontinuous habitat patches is crucial for the persistence of metapopulations across space and time. Loss of landscape connectivity is often a direct result of fragmentation caused by human activities but also can be caused indirectly through anthropogenic climate change. Peary caribou (Rangifer tarandus pearyi) are widely dispersed across the islands of the Canadian Arctic Archipelago and rely on sea ice to move seasonally between island habitats throughout their range. Seasonal connectivity provided by sea ice is necessary to maintain genetic diversity and to facilitate dispersal and recolonization of areas from which caribou have been extirpated. We used least‐cost path analysis and circuit theory to model connectivity across Peary caribou range, and future climate projections to investigate how this connectivity might be affected by a warming climate. Further, we used measures of current flow centrality to estimate the role of High Arctic islands in maintaining connectivity between Peary caribou populations and to identify and prioritize those islands and linkages most important for conservation. Our results suggest that the Bathurst Island complex plays a critical role in facilitating connectivity between Peary caribou populations. Large islands, including Banks, Victoria, and Ellesmere have limited roles in connecting Peary caribou. Without rigorous greenhouse gas emission reductions our projections indicate that by 2100 all connectivity between the more southern Peary caribou populations will be lost for important spring and early‐winter movement periods. Continued connectivity across the Canadian Arctic Archipelago, and possibly Peary caribou persistence, ultimately hinges on global commitments to limit climate change. Our research highlights priority areas where, in addition to emission reductions, conservation efforts to maintain connectivity would be most effective.     

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.     

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.     

No evidence of inbreeding depression in fast declining herds of migratory caribou

Identifying inbreeding depression early in small and declining populations is essential for management and conservation decisions. Correlations between heterozygosity and fitness (HFCs) provide a way to identify inbreeding depression without prior knowledge of kinship among individuals. In Northern Quebec and Labrador, the size of two herds of migratory caribou (Rivière‐George, RG and Rivière‐aux‐Feuilles, RAF) has declined by one to two orders of magnitude in the last three decades. This raises the question of a possible increase in inbreeding depression originating from, and possibly contributing to, the demographic decline in those populations. Here, we tested for the association of genomic inbreeding indices (estimated with 22,073 SNPs) with body mass and survival in 400 caribou sampled in RG and RAF herds between 1996 and 2016. We found no association of individual heterozygosity or inbreeding coefficient with body mass or annual survival. Furthermore, those genomic inbreeding indices remained stable over the period monitored. These results suggest that the rapid and intense demographic decline of the herds did not cause inbreeding depression in those populations. Although we found no evidence for HFCs, if demographic decline continues, it is possible that such inbreeding depression would be triggered.     

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.     

Effects of weather and hunting on wild reindeer population dynamics in Hardangervidda National Park

Wild reindeer have a range that extends across the circumpolar region. In the last few decades, however, populations of wild reindeer have been on the decline. The reasons for these declines are poorly understood, but are suggested to be linked to both local and global climatic factors, disease, and human interference. Hardangervidda plateau in Norway is home to the largest wild reindeer population in Europe, and is at the southern end of its European range. This population is therefore of particular importance, particularly in the light of climate change. We investigated how weather and hunting have affected the wild reindeer population in Hardangervidda over the last two decades. Our findings suggest that the wild reindeer population in Hardangervidda is most affected by winter temperature and hunting, where colder temperatures and lower harvest rates typically result in higher growth rates. We did not find significant evidence for linear density dependence. Our results show trends across Hardangervidda, and give an indication of how region-wide weather and hunting pressure can affect the wild reindeer population. As new data emerge, future investigations should look into the existence and nature of density dependence and the influence of other weather and human disturbance related factors.     

The role of predation in the decline and extirpation of woodland caribou

To select appropriate recovery strategies for endangered populations, we must understand the dynamics of small populations and distinguish between the possible causes that drive such populations to low numbers. It has been suggested that the pattern of population decline may be inversely density-dependent with population growth rates decreasing as populations become very small; however, empirical evidence of such accelerated declines at low densities is rare. Here we analyzed the pattern of decline of a threatened population of woodland caribou (Rangifer tarandus caribou) in British Columbia, Canada. Using information on the instantaneous rate of increase relative to caribou density in suitable winter foraging habitat, as well as on pregnancy rates and on causes and temporal distribution of mortalities from a sample of 349 radiocollared animals from 15 subpopulations, we tested 3 hypothesized causes of decline: (a) food regulation caused by loss of suitable winter foraging habitat, (b) predation-sensitive foraging caused by loss of suitable winter foraging habitat and (c) predation with caribou being secondary prey. Population sizes of caribou subpopulations ranged from <5 to >500 individuals. Our results showed that the rates of increase of these subpopulations varied from −0.1871 to 0.0496 with smaller subpopulations declining faster than larger subpopulations. Rates of increase were positively related to the density of caribou in suitable winter foraging habitat. Pregnancy rates averaged 92.4% ±2.24 and did not differ among subpopulations. In addition, we found predation to be the primary cause of mortality in 11 of 13 subpopulations with known causes of mortality and predation predominantly occurred during summer. These results are consistent with predictions that caribou subpopulations are declining as a consequence of increased predation. Recovery of these woodland caribou will thus require a multispecies perspective and an appreciation for the influence of inverse density dependence on population trajectories.