Robert D. Otto

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 all ≤ 0.005), whereas sedentary herds were more differentiated (FST = 0.018–0.048). Geographical distances among sedentary herds limited gene flow. Historical estimates of gene flow were higher from migratory herds into sedentary herds (4Nm all > 9) 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.

VARIATION IN CALF BODY MASS IN MIGRATORY CARIBOU : THE ROLE OF HABITAT, CLIMATE, AND MOVEMENTS

Abstract Individual differences in body mass exert a major influence on several life-history traits of mammals. We investigated the factors influencing variation in body mass of calves of migratory caribou (Rangifer tarandus) at birth (June, 19 years of data) and in autumn (October, 15 years of data) in the Rivière-aux-Feuilles (Feuilles, 1991–2003) herd and the Rivière-George (George, 1978–2003) herd in Québec and Labrador, Canada. Mass at birth (hereafter, birth mass) did not differ between herds, possibly because part of their winter ranges overlapped. However, Feuilles calves were smaller in autumn than George calves, possibly reflecting differences in summer ranges. The birth mass of calves also varied with year, likely as the outcome of both intrinsic and extrinsic factors. Birth and autumn body mass were influenced positively by habitat quality in June, estimated by the normalized difference vegetation index. The North Atlantic Oscillation of the previous winter was positively correlated with autumn mass of the George calves. Previous winter snowfall was negatively related to the mass of George calves, and daily movement rates in summer were negatively correlated with the mass of calves of both herds in autumn. Birth mass was positively related with productivity in October in the George herd and also with productivity 3 and 4 years later, which corresponds to the beginning of reproduction of females. We suggest that a mechanism of delayed quality effect of the calves could have been involved in the decrease of fall productivity and population size of the George herd.

Body Size Variations in Caribou Ecotypes and Relationships With Demography

Abstract In many vertebrates size is one of the most influential and variable individual characteristics and a strong determinant of reproductive success. Body size is generally density dependent and decreases when intraspecific competition increases. Frequent and long-distance movements increase energy expenditures and, therefore, may also influence body size, particularly in highly mobile species. Caribou (Rangifer tarandus, also known as reindeer) exhibit tremendous variation in size and movements and thus represent an excellent candidate species to test the relationships between body size, population size, and movements. We analyzed body measurements of adult female caribou from 7 herds of the Québec-Labrador Peninsula, Canada, and we related their morphology to population size, movements, and annual ranges. The herds represented 3 ecotypes (migratory, montane, and sedentary). Ecotypes and herds differed in size (length), shape (roundness), and movements. The sedentary ecotype was larger and moved 4 to 7 times less than the migratory ecotype in the 1990s. At the start of a demographic growth period in the early 1960s, migratory caribou from the Rivière-George (hereafter George) herd had longer mandibles than caribou of the sedentary ecotype. Mandible length in the George herd declined in the 1980s after rapid population growth, while individuals performed extensive movements and the herds annual range increased. Migratory caribou then became shorter than sedentary caribou. After the George herd decline in the 1990s, mandible length increased again near levels of the 1980s. Caribou from the migratory Rivière-aux-Feuilles herd later showed a similar decline in mandible length during a period of population growth, associated with longer movements and increasing annual range. We hypothesize that the density-dependent effect observed on body size might have been exerted through summer habitat degradation and movement variations during herd growth. Our study has 2 important implications for caribou management: the distinctiveness of different populations and ecotypes, and the correlations between population trajectories and changes in body condition and habitat.

Body-condition dynamics in a northern ungulate gaining fat in winter

Individual condition generally depends on density and is partly determined by habitat quality and climate. We studied long-term trends in the condition and productivity of female caribou (Rangifer tarandus (L., 1758)) in two large migratory herds in the Quebec–Labrador peninsula (Canada), the George and the Feuilles herds. Females from the George herd were in better summer condition than those from the more abundant Feuilles herd in 2001–2002, while it was the opposite in 1988 when the Feuilles herd was less abundant than the George herd. Summer nutrition followed the same pattern between herds through time. Spring body condition of females in the George herd declined from 1976 to the mid-1980s during early population growth. Fall condition, however, did not change from 1983 to 2002 when caribou numbers first peaked and later declined. Pregnancy rates were inversely related to herd size in both herds. Vegetation quality (NDVI) in June was significantly related to body proteins in the fall. Albeit unusual ...

Winter Habitat Associations of a Low-Density Moose (Alces americanus) Population in Central Labrador

Abstract Alces americanus (Moose) are relatively new to Labrador, having only colonized the area in the late 1940s, and little is known about this population. We conducted large-scale aerial surveys for Moose in a 122,000-km2 area during winter 2000 and in a 29,900-km2 area in winter 2001. Moose densities were low in each area (1.6–3.0 Moose per 100 km2). Bull:cow ratios were nearly even and calf:cow ratios were relatively high, indicative of a population exposed to little hunting or predation pressure. Twinning rates were low, suggesting low range productivity. Moose used riparian areas and hardwood stands in higher proportion than their availability in winter (P < 0.05). Open habitats (conifer-lichen woodlands, bogs and fens, burned forest, and barren areas) were used in lower proportion than their availability. These data may provide the basis for developing habitat suitability maps for Moose in late winter across central Labrador.

Winter range drift in the George River Caribou Herd: a response to summer forage limitation?

Space use by the George River caribou herd (GRCH) changes in correspondence with migration patterns. The traditional range of this herd encompasses an area of approximately 900 000 km2. Range use is seasonal and includes travel to traditional calving grounds. Winter range use however, is more variable. The GRCH has grown rapidly from 5000 animals in 1954 to approximately 775 000 in 1993. Beginning in the mid 1980s, the calving and summer range habitats of the GRCH have deteriorated, resulting in a decline in physical condition and subsequent poor calf survival and low pregnancy rates. We assessed the importance of the winter range as a food source compensating for poor summer range quality through an evaluation of winter range drift and use intensity. We hypothesized that if winter ranges provide a compensatory source of forage, then George River caribou should avoid sites heavily used during the previous winter at a population level. Winter ranges for the GRCH were calculated using 4300 caribou locations obtained 1986-2000. We found that in spite of a doubling in net range area, the size of annual winter ranges did not increase, indicating the occurrence of range drift. Further, George River caribou exhibited avoidance of wintering areas at several spatial scales. Avoidance occurred across a use threshold, where the degree of use (or density) during the previous winter determined the level of avoidance during the subsequent winter. As the spatial scale decreased, the degree of avoidance increased. Caribou significantly avoided areas used the previous winter at spatial scales below and including 245 km2 (corresponding to a 75% use distribution). Results suggest winter foraging allows caribou suspend the effects of density-dependent summer forage limitation on herd productivity. As such, analysis of GRCH population trends should be considered in light of both summer and winter range resources.

Evaluation of satellite collar sample size requirements for mitigation of low-level military jet disturbance of the George River caribou herd

Wildlife radio-telemetry and tracking projects often determine a priori required sample sizes by statistical means or default to the maximum number that can be maintained within a limited budget. After initiation of such projects, little attention is focussed on effective sample size requirements, resulting in lack of statistical power. The Department of National Defence operates a base in Labrador, Canada for low level jet fighter training activities, and maintain a sample of satellite collars on the George River caribou (Rangifer tarandus caribou) herd of the region for spatial avoidance mitiga¬tion purposes. We analysed existing location data, in conjunction with knowledge of life history, to develop estimates of satellite collar sample sizes required to ensure adequate mitigation of GRCH. We chose three levels of probability in each of six annual caribou seasons. Estimated number of collars required ranged from 15 to 52, 23 to 68, and 36 to 184 for 50%, 75%, and 90% probability levels, respectively, depending on season. Estimates can be used to make more informed decisions about mitigation of GRCH, and, generally, our approach provides a means to adaptively assess radio collar sam¬ple sizes for ongoing studies.

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.

George’s Island, Labrador - A high-density predator-free refuge for a woodland caribou subpopulation?

The movement patterns and demographic parameters were measured for caribou (Rangifer tarandus caribou) on George’s Island (Labrador, Canada) to determine if the population is separate from the Mealy Mountain Caribou Herd. Movements between George’s Island caribou and nearby Mealy Mountain caribou were examined through satellite telemetry (April 2005 to April 2006). Demographic information was collected through aerial classification surveys. The predator-free island is currently maintaining a density of 22.5-26.5 caribou/km2. Female survival appears high and the recruitment rate in late fall-early spring was 19.0-29.2% calves. Mainland caribou moved very little throughout the year, travelling no more than 53.7 km on average from their initial collaring locations. Also, satellite data indicated no mixing between animals on George’s Island and the mainland. The elevated caribou density and high proportion of calves suggest that George’s Island could at times be acting as a predator-free recruitment area and that George’s Island may be a subpopulation from which animals disperse to the mainland.