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Caribou distribution during the post-calving period in relation to infrastructure in the Prudhoe Bay oil field, Alaska

There is concern that caribou ( Rangifer tarandus ) may avoid roads and facilities (i.e., infrastructure) in the Prudhoe Bay oil field (PBOF) in northern Alaska, and that this avoidance can have negative effects on the animals. We quantified the relationship between caribou distribution and PBOF infrastructure during the post-calving period (mid-June to mid-August) with aerial surveys from 1990 to 1995. We conducted four to eight surveys per year with complete coverage of the PBOF. We identified active oil field infrastructure and used a geographic information system (GIS) to construct ten 1 km wide concentric intervals surrounding the infrastructure. We tested whether caribou distribution is related to distance from infrastructure with a chi-squared habitat utilization-availability analysis and log-linear regression. We considered bulls, calves, and total caribou of all sex/age classes separately. The habitat utilization-availability analysis indicated there was no consistent trend of attraction to or avoidance of infrastructure. Caribou frequently were more abundant than expected in the intervals close to infrastructure, and this trend was more pronounced for bulls and for total caribou of all sex/age classes than for calves. Log-linear regression (with Poisson error structure) of numbers of caribou and distance from infrastructure were also done, with and without combining data into the 1 km distance intervals. The analysis without intervals revealed no relationship between caribou distribution and distance from oil field infrastructure, or between caribou distribution and Julian date, year, or distance from the Beaufort Sea coast. The log-linear regression with caribou combined into distance intervals showed the density of bulls and total caribou of all sex/age classes declined with distance from infrastructure. Our results indicate that during the post-calving period: 1) caribou distribution is largely unrelated to distance from infrastructure; 2) caribou regularly use habitats in the PBOF; 3) caribou often occur close to infrastructure; and 4) caribou do not appear to avoid oil field infrastructure.

Alaska

Collection and analysis of traditional ecological knowledge about a population of arctic tundra caribou

Aboriginal peoples want their ecological knowledge used in the management of wildlife populations. To accomplish this, management agencies will need regional summaries of aboriginal knowledge about long-term changes in the distribution and abundance of wildlife populations and ecological factors that influence those changes. Between 1983 and 1994, we developed a method for collecting Inuit knowledge about historical changes in a caribou ( Rangifer tarandus ) population on southern Baffin Island from c. 1900 to 1994. Advice from Inuit allowed us to collect and interpret their oral knowledge in culturally appropriate ways. Local Hunters and Trappers Associations (HTAs) and other Inuit identified potential informants to maximize the spatial and temporal scope of the study. In the final interview protocol, each informant (i) established his biographical map and time line, (ii) described changes in caribou distribution and density during his life, and (iii) discussed ecological factors that may have caused changes in caribou populations. Personal and parental observations of caribou distribution and abundance were reliable and precise. Inuit who had hunted caribou during periods of scarcity provided more extensive information than those hunters who had hunted mainly ringed seals ( Phoca hispida ); nevertheless, seal hunters provided information about coastal areas where caribou densities were insufficient for the needs of caribou hunters. The wording of our questions influenced the reliability of informants' answers; leading questions were especially problematic. We used only information that we considered reliable after analyzing the wording of both questions and answers from translated transcripts. This analysis may have excluded some reliable information because informants tended to understate certainty in their recollections. We tried to retain the accuracy and precision inherent in Inuit oral traditions; comparisons of information from several informants and comparisons with published and archival historical reports indicate that we retained these qualities of Inuit knowledge.

Nunavut

Central Arctic caribou and petroleum development: Distributional, nutritional, and reproductive implications

We synthesize findings from cooperative research on effects of petroleum development on caribou ( Rangifer tarandus granti ) of the Central Arctic Herd (CAH). The CAH increased from about 6000 animals in 1978 to 23000 in 1992, declined to 18 000 by 1995, and again increased to 27 000 by 2000. Net calf production was consistent with changes in herd size. In the Kuparuk Development Area (KDA), west of Prudhoe Bay, abundance of calving caribou was less than expected within 4 km of roads and declined exponentially with road density. With increasing infrastructure, high-density calving shifted from the KDA to inland areas with lower forage biomass. During July and early August, caribou were relatively unsuccessful in crossing road/pipeline corridors in the KDA, particularly when in large, insect-harassed aggregations; and both abundance and movements of females were lower in the oil field complex at Prudhoe Bay than in other areas along the Arctic coast. Female caribou exposed to petroleum development west of the Sagavanirktok River may have consumed less forage during the calving period and experienced lower energy balance during the midsummer insect season than those under disturbance-free conditions east of the river. The probable consequences were poorer body condition at breeding and lower parturition rates for western females than for eastern females (e.g., 1988–94: 64% vs. 83% parturient, respectively; p = 0.003), which depressed the productivity of the herd. Assessments of cumulative effects of petroleum development on caribou must incorporate the complex interactions with a variable natural environment.

Arctic

Aggression and coexistence in female caribou

Female caribou ( Rangifer tarandus ) are highly gregarious, yet there has been little study of the behavioral mechanisms that foster coexistence. Quantifying patterns of aggression between male and female, particularly in the only cervid taxa where both sexes grow antlers, should provide insight into these mechanisms. We asked if patterns of aggression by male and female caribou followed the pattern typically noted in other polygynous cervids, in which males display higher frequencies and intensity of aggression. From June to August in 2011 and 2012, we measured the frequency and intensity of aggression across a range of group sizes through focal animal sampling of 170 caribou (64 males and 106 females) on Adak Island in the Aleutian Archipelago, Alaska. Males in same-sex and mixed-sex groups and females in mixed-sex groups had higher frequencies of aggression than females in same-sex groups. Group size did not influence frequency of aggression. Males displayed more intense aggression than females. Frequent aggression in mixed-sex groups probably reflects lower tolerance of males for animals in close proximity. Female caribou were less aggressive and more gregarious than males, as in other polygynous cervid species.

Alaska

Evaluating potential effects of an industrial road on winter habitat of caribou in North-Central Alaska

Worldwide, some caribou (Rangifer tarandus) populations are experiencing declines due partially to the expansion of industrial development. Caribou can exhibit behavioral avoidance of development, leading to indirect habitat loss, even if the actual footprint is small. Thus, it is important to understand before construction begins how much habitat might be affected by proposed development. In northern Alaska, an industrial road that has been proposed to facilitate mining transects a portion of the Western Arctic caribou herd's winter range. To understand how winter habitat use might be affected by the road, we estimated resource selection patterns during winter for caribou in a study area surrounding the proposed road. We assessed the reductions of habitat value associated with three proposed routes at three distance thresholds for disturbance. High-value winter habitat tended to occur in locally rugged areas that have not burned recently and have a high density of lichen and early dates of spring snowmelt. We found that 1.5% to 8.5% (146-848 km2) of existing high-value winter habitat in our study area might be reduced in quality. The three alternative routes were only marginally different. Our results suggest that the road would have minimal direct effects on high-value winter habitat; however, additional cumulative impacts to caribou (e.g., increased access by recreationists and hunters) should be considered before the full effects of the road can be estimated.

Alaska

Moose, caribou, and grizzly bear distribution in relation to road traffic in Denali National Park, Alaska

Park managers are concerned that moose ( Alces alces ), caribou ( Rangifer tarandus ), and grizzly bears ( Ursus arctos ) may be avoiding areas along the 130 km road through Denali National Park as a result of high traffic volume, thus decreasing opportunities for visitors to view wildlife. A wildlife monitoring system was developed in 1996 that used 19 landscape level view sheds, stratified into four sections based on decreasing traffic along the road corridor. Data were collected from 22 samplings of all viewsheds during May-August in 1996 and 1997. In 1997, nine backcountry viewsheds were established in three different areas to determine whether density estimates for each species in the backcountry were higher than those for the same animals in similar road-corridor areas. Densities higher than those in the road corridor were found in one backcountry area for moose and in two backcountry areas for grizzly bears. None of the backcountry areas showed a higher density of caribou. We tested hypotheses that moose, caribou, and grizzly bear distributions were unrelated to the road and traffic. Moose sightings were lower than expected within 300 m of the road. More caribou and grizzly bears than expected occurred between 601 and 900 m from the road, while more moose and fewer caribou than expected occurred between 900 and 1200 m from the road. Bull moose in stratum 1 were distributed farther from the road than bulls and cows in stratum 4; cows in stratum 1 and bul1s in stratum 2 were distributed farther from the road than cows in stratum 4. Grizzly bears in stratum 2 were distributed farther from the road than bears in stratum 3. The distribution of moose sightings suggests traffic avoidance, but the spatial pattern of preferred forage may have had more of an influence. Caribou and grizzly bear distributions indicated no pattern of traffic avoidance.

Alaska

Stochastic and compensatory effects limit persistence of variation in body mass of young caribou

Nutritional restriction during growth can have short- and long-term effects on fitness; however, animals inhabiting uncertain environments may exhibit adaptations to cope with variation in food availability. We examined changes in body mass in free-ranging female caribou ( Rangifer tarandus ) by measuring mass at birth and at 4, 11, and 16 months of age to evaluate the relative importance of seasonal nutrition to growth, the persistence of cohort-specific variation in body mass through time, and compensatory growth of individuals. Relative mean body mass of cohorts did not persist through time. Compensatory growth of smaller individuals was not observed in summer; however, small calves exhibited more positive change in body mass than did large calves. Compensation occurred during periods of nutritional restriction (winter) rather than during periods of rapid growth (summer) thus differing from the conventional view of compensatory growth.

Journal of Mammalogy

Habitat selection by a focal predator (Canis lupus) in a multiprey ecosystem of the northern Rockies

Large predators respond to land cover and physiography that maximize the likelihood of encountering prey. Using locations from global positioning system-collared wolves (Canis lupus), we examined whether land cover, vegetation productivity or change, or habitat-selection value for ungulate prey species themselves most influenced patterns of selection by wolves in a large, intact multiprey system of northern British Columbia. Selection models based on land cover, in combination with topographical features, consistently outperformed models based on indexes of vegetation quantity and quality (using normalized difference vegetation index) or on selection value to prey species (moose [Alces americanus], elk [Cervus elaphus], woodland caribou [Rangifer tarandus], and Stone's sheep [Ovis dalli stonei]). Wolves generally selected for shrub communities and high diversity of cover across seasons and avoided conifer stands and non-vegetated areas and west aspects year-round. Seasonal selection strategies were not always reflected in use patterns, which showed highest frequency of use in riparian, shrub, and conifer classes. Patterns of use and selection for individual wolf packs did not always conform to global models, and appeared related to the distribution of land cover and terrain within respective home ranges. Our findings corroborate the biological linkages between wolves and their habitat related to ease of movement and potential prey associations. ?? American 2011 Society of Mammalogists.

Journal of Mammalogy

Diversity of nitrogen isotopes and protein status in caribou: implications for monitoring northern ungulates

Nutritional condition is an important determinant of productivity and survival in caribou (Rangifer tarandus). We used samples of excreta (n = 1,150) to estimate diet composition from microhistology and 2 isotopic proxies of protein status for 2 ecotypes of caribou in 4 herds in late winter (2006–2008). Isotopes of nitrogen (δ 15 N in parts per thousand [‰]) from excreta samples (urea, diet, and body N) were used to estimate indexes of protein status: the proportion of urea N derived from body N (p-UN) and the difference between the δ15N of the body and urinary urea (Δbody-urea). We examined dietary and terrain characteristics, δ 15 N, p-UN, and Δ body-urea by ecotype, herd, year, and foraging site. Multiple regression and an information-theoretic approach were used to evaluate correlates of protein status at each foraging site. The dietary and terrain characteristics of foraging sites did not vary by ecotype or herd (P > 0.108); diets were dominated by lichens (68% ± 14.1% SD). The δ 15 N of urea, diet, body N, p-UN, and Δ body-urea varied among foraging sites within each herd (P < 0.001). Although highly variable, the δ 15 N of urinary urea was typically low (−4.68‰ ± 2.67‰ SD). Dietary N also had low δ 15 N (−4.18‰ ± 0.92‰ SD), whereas body N was generally heavier in 15 N (2.20‰ ± 1.56‰ SD) than urinary urea or the diet. Both measures of protein status were similarly diverse between ecotypes and among herds, which limited their applicability to monitor protein status at the population level. Although we observed limitations to interpreting estimates of p-UN from highly vagile ungulates, the Δbody-urea may prove to be a useful index of protein status at smaller spatial and temporal scales. Indeed, a portion of the observed variance (r 2 = 0.26) in Δbody-urea at each foraging site was explained by the proportion of shrubs in the winter diet. There remains potential in using δ 15 N in excreta as a noninvasive tool for evaluating protein status in northern ungulates; however, considerable analytical and sampling challenges remain for applying these isotopic approaches at large scales.

Alaska

Range overlap and individual movements during breeding season influence genetic relationships of caribou herds in south-central Alaska

North American caribou ( Rangifer tarandus ) herds commonly exhibit little nuclear genetic differentiation among adjacent herds, although available evidence supports strong demographic separation, even for herds with seasonal range overlap. During 1997–2003, we studied the Mentasta and Nelchina caribou herds in south-central Alaska using radiotelemetry to determine individual movements and range overlap during the breeding season, and nuclear and mitochondrial DNA (mtDNA) markers to assess levels of genetic differentiation. Although the herds were considered discrete because females calved in separate regions, individual movements and breeding-range overlap in some years provided opportunity for male-mediated gene flow, even without demographic interchange. Telemetry results revealed strong female philopatry, and little evidence of female emigration despite overlapping seasonal distributions. Analyses of 13 microsatellites indicated the Mentasta and Nelchina herds were not significantly differentiated using both traditional population-based analyses and individual-based Bayesian clustering analyses. However, we observed mtDNA differentiation between the 2 herds (F STM = 0.041, P < 0.001). Although the Mentasta and Nelchina herds exhibit distinct population dynamics and physical characteristics, they demonstrate evidence of gene flow and thus function as a genetic metapopulation.

Alaska

Spatial occupancy models for large data sets

Since its development, occupancy modeling has become a popular and useful tool for ecologists wishing to learn about the dynamics of species occurrence over time and space. Such models require presence–absence data to be collected at spatially indexed survey units. However, only recently have researchers recognized the need to correct for spatially induced overdisperison by explicitly accounting for spatial autocorrelation in occupancy probability. Previous efforts to incorporate such autocorrelation have largely focused on logit-normal formulations for occupancy, with spatial autocorrelation induced by a random effect within a hierarchical modeling framework. Although useful, computational time generally limits such an approach to relatively small data sets, and there are often problems with algorithm instability, yielding unsatisfactory results. Further, recent research has revealed a hidden form of multicollinearity in such applications, which may lead to parameter bias if not explicitly addressed. Combining several techniques, we present a unifying hierarchical spatial occupancy model specification that is particularly effective over large spatial extents. This approach employs a probit mixture framework for occupancy and can easily accommodate a reduced-dimensional spatial process to resolve issues with multicollinearity and spatial confounding while improving algorithm convergence. Using open-source software, we demonstrate this new model specification using a case study involving occupancy of caribou (Rangifer tarandus) over a set of 1080 survey units spanning a large contiguous region (108 000 km 2 ) in northern Ontario, Canada. Overall, the combination of a more efficient specification and open-source software allows for a facile and stable implementation of spatial occupancy models for large data sets.

Ecology

Deer exposed to exceptionally high concentrations of lead near the Continental Mine in Idaho, USA

Habitat surrounding the inactive Continental Mine in northern Idaho, USA, supports bear ( Ursus arctos, Ursus americanus ), moose ( Alces alces ), elk ( Cervus elaphus ), woodland caribou ( Rangifer tarandus caribou ), and abundant mule ( Odocoileus hemionus ) and white-tailed deer ( Odocoileus virginianus ). Tailings on the mining site were capped and remediated in 2003 to reduce environmental exposure of surrounding soil and sediments of Blue Joe Creek, downslope of the mine. Before capping, the mean Pb concentration in deer pellets collected on-site was 920 mg/kg of Pb (dry wt). This exposure, if chronic, would be comparable to an exposure that could be lethal to cattle or horses. Surprisingly, the mean pellet Pb concentration of 950 mg/kg in 2004 was as high as it was before remediation, and it was related to a high rate of soil ingestion. Mean soil content of the pellets collected from the capped site in 2004 was 22% dry weight, estimated from the acid-insoluble ash, a marker of soil ingestion. Clumps of sand and bits of rock were observed inside some of the pellets, and Pb concentrations in the pellets were correlated (p < 0.05) with soil content. Although terrestrial risk assessments generally estimate exposure from diets and from incidentally ingested soil, the deer at this site were directly ingesting contaminated soil or mining waste. The mean Pb concentration of this ingested soil was estimated as 6,700 mg/kg and the maximum as 25,000 mg/kg, well above the Pb concentrations measured in the remediated cap. The deer seemed to be ingesting soil or mining waste from one or more small but highly contaminated sources located beyond the remediated cap.

Idaho

A Bayesian random effects discrete-choice model for resource selection: Population-level selection inference

Modeling the probability of use of land units characterized by discrete and continuous measures, we present a Bayesian random-effects model to assess resource selection. This model provides simultaneous estimation of both individual- and population-level selection. Deviance information criterion (DIC), a Bayesian alternative to AIC that is sample-size specific, is used for model selection. Aerial radiolocation data from 76 adult female caribou (Rangifer tarandus) and calf pairs during 1 year on an Arctic coastal plain calving ground were used to illustrate models and assess population-level selection of landscape attributes, as well as individual heterogeneity of selection. Landscape attributes included elevation, NDVI (a measure of forage greenness), and land cover-type classification. Results from the first of a 2-stage model-selection procedure indicated that there is substantial heterogeneity among cow-calf pairs with respect to selection of the landscape attributes. In the second stage, selection of models with heterogeneity included indicated that at the population-level, NDVI and land cover class were significant attributes for selection of different landscapes by pairs on the calving ground. Population-level selection coefficients indicate that the pairs generally select landscapes with higher levels of NDVI, but the relationship is quadratic. The highest rate of selection occurs at values of NDVI less than the maximum observed. Results for land cover-class selections coefficients indicate that wet sedge, moist sedge, herbaceous tussock tundra, and shrub tussock tundra are selected at approximately the same rate, while alpine and sparsely vegetated landscapes are selected at a lower rate. Furthermore, the variability in selection by individual caribou for moist sedge and sparsely vegetated landscapes is large relative to the variability in selection of other land cover types. The example analysis illustrates that, while sometimes computationally intense, a Bayesian hierarchical discrete-choice model for resource selection can provide managers with 2 components of population-level inference: average population selection and variability of selection. Both components are necessary to make sound management decisions based on animal selection.

Journal of Wildlife Management

Intraspecific mitochondrial DNA variation in North American cervids

Intraspecific variation in mitochondrial DNA of North American cervids was assessed with restriction enzymes to determine relationships among populations and subspecies. No variation was detected in moose ( Alces alces ) and little in elk ( Cervus elaphus ). Caribou ( Rangifer tarandus ), white-tailed deer ( Odocoileus virginianus ), and mule deer ( Odocoileus hemionus ) possessed considerable variation. Characteristic genotypes exist in caribou and white-tailed deer from different geographic areas although subspecies are not discernable as distinct mtDNA assemblages. Except for O. hemionus , intraspecific mtDNA sequence divergences are small (<2%). Subspecies of mule deer have divergent mtDNA (7%) and are the only subspecies of cervids with distinct genotypes.

Journal of Mammalogy

Timing and synchrony of parturition in Alaska caribou

Timing of parturition of caribou ( Rangifer tarandus ) varies within populations, but the relative influences of nutritional condition of females during the autumn breeding season and during gestation on that variation is not known. We determined timing of parturition of caribou in Denali National Park, Alaska, during 1984–1995, which had wide variation in snowfall that influenced nutritional condition and productivity of females. The first young were observed each year between 4 and 15 May. Annual median dates of parturition for radiocollared females during 1987–1995 varied from 13 to 21 May. Synchrony of births (i.e., proportion of births during 7 days centered on the median date) did not vary significantly among years ( P > 0.05). Females ≤4 years old were more likely to give birth after the annual median date than older females ( P = 0.005). Dates of parturition were significantly correlated with masses of females in the previous autumn ( P = 0.034) but not in late winter ( P = 0.155). Births tended to be earlier for parous females that were not lactating during the previous breeding season compared to those that were lactating ( P = 0.030). Annual onsets of parturition varied significantly with snowfall in late winter prior to conception ( P = 0.012) but not with snowfall during gestation. Timing of parturition in our study appeared to be a function of physical condition of breeding females in autumn.

Journal of Mammalogy

Reoccurrence of caribou in Minnesota

At least two caribou ( Rangifer tarandus ) were observed in Cook Co. of northeastern Minnesota from October 1980 through April 1981. This is the first record of caribou in the area since 1924. The nearest records of this species were 224 km NE of the present sightings, and the closest known established population of caribou is 240 km away.

Minnesota

Reproductive performance of female Alaskan caribou

We examined the reproductive performance of female caribou ( Rangifer tarandus granti ) in relation to age, physical condition, and reproductive experience for 9 consecutive years (1987-95) at Denali National Park, Alaska, during a period of wide variation in winter snowfall. Caribou in Denali differed from other cervid populations where reproductive performance has been investigated, because they occur at low densities (≥0.3/km 2 ) and experience high losses of young to predation. Females first gave birth at 2-6 years old; 56% of these females were 3 years old. Average annual natality rates increased from 27% for 2-year-olds to 100% for 7-year-olds, remained high for 7-13-year-olds (98%), and then declined for females ≥14 years old. Females ≥2 years old that failed to reproduce were primarily sexually immature (76%). Reproductive pauses of sexually mature females occurred predominantly in young (3-6 yr old) and old (≥14 yr old) females. Natality increased with body mass for 10-month-old females weighed 6 months prior to the autumn breeding season (P = 0.007), and for females >1 year old and weighed during autumn (late Sep-early Nov; P = 0.003). Natality for 2-, 3-, 4-, and 6-year-olds declined with increasing late-winter snowfall (Feb-May; P ≤ 0.039) during the winter prior to breeding. In most years, a high percentage of sexually mature females reproduced, and lactation status at the time of breeding did not influence productivity the following year. However, following particularly high snowfall during February-September 1992, productivity was reduced in 1993 for cows successfully rearing calves to autumn the previous year. High losses of calves to predators in 1992 may have increased productivity in 1993. Losses of young-of-the-year to predation prior to the annual breeding season can be an important influence on subsequent productivity for ungulate populations where productivity varies with lactation status of females at the time of breeding.

Journal of Wildlife Management

Grizzly bears and calving caribou: What is the relation with river corridors?

Researchers have debated the effect of the Trans-Alaska Pipeline (TAP) and associated developments to caribou ( Rangifer tarandus ) of the central Arctic herd (CAH) since the 1970s. Several studies have demonstrated that cows and calves of the CAH avoided the TAP corridor because of disturbance associated with the pipeline, whereas others have indicated that female caribou of the CAH avoided riparian habitats closely associated with the pipeline. This avoidance was explained as a predator-avoidance strategy. We investigated the relation between female caribou and grizzly bear ( Ursus arctos ) use of river corridors on the yet undisturbed calving grounds of the Porcupine caribou herd (PCH) in northeastern Alaska. On the coastal plain, caribou were closer to river corridors than expected (P = 0.038), but bear use of river corridors did not differ from expected (P = 0.740). In the foothills, caribou use of river corridors did not differ from expected (P = 0.520), but bears were farther from rivers than expected (P = 0.001). Our results did not suggest an avoidance of river corridors by calving caribou or a propensity for bears to be associated with riparian habitats, presumably for stalking or ambush cover. We propose that PCH caribou reduce the risks of predation to neonates by migrating to a common calving grounds, where predator swamping is the operational antipredator strategy. Consequently, we hypothesize that nutritional demands, not predator avoidance strategies, ultimately regulate habitat use patterns (e.g., use of river corridors) of calving PCH caribou.

Alaska