Bruce W. Dale

Functional response of wolves preying on barren-ground caribou in a multiple-prey ecosystem

1. We investigated the functional response of wolves (Canis lupus) to varying abundance of ungulate prey to test the hypothesis that switching from alternate prey to preferred prey results in regulation of a caribou (Rangifer tarandus) population at low densities. 2. We determined prey selection, kill rates, and prey abundance for four wolf packs during three 30-day periods in March 1989, March 1990 and November 1990, and created a simple discrete model to evaluate the potential for the expected numerical and observed functional responses of wolves to regulate caribou populations. 3. We observed a quickly decelerating type II functional response that, in the absence of a numerical response, implicates an anti-regulatory effect of wolf predation on barren-ground caribou dynamics

Caribou calf mortality in Denali National Park, Alaska

Calf mortality is a major component of caribou (Rangifer tarandus) population dynamics, but little is known about the timing or causes of calf losses, or of characteristics that predispose calves to mortality. During 1984-87, we radiocollared 226 calves (≤3 days old) in the Denali Caribou Herd (DCH), an unhunted population utilized by a natural complement of predators, to determine the extent, timing, and causes of calf mortality and to evaluate influences of year, sex, birthdate, and birth mass on those losses. Overall, 39% of radio-collared calves died as neonates (≤ 15 days old), and 98% of those deaths were attributed to predation. Most neonatal deaths (85%) occurred within 8 days of birth. Few deaths occurred after the neonatal period (5, 10, and 0% of calves instrumented died during 16-30, 31-150, and >150 days of age, respectively). Survival of neonates was lower (P = 0.038) in 1985, following a severe winter, than during the other 3 years. In years other than 1985, calves born during the peak of calving (approx 50% of the total, born 5-8 days after calving onset) experienced higher (P 10 days old. Wolf predation was not related (P > 0.05) to calf age and peaked 10 days after onset of calving. Grizzly bear and wolf predation on neonates during the calving season was a limiting factor for the Denali Caribou Herd.

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 witl 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.

Population Dynamics and Harvest Characteristics of Wolves in the Central Brooks Range, Alaska

Abstract Our understanding of wolf (Canis lupus) population dynamics in North America comes largely from studies of protected areas, at-risk populations, and wolf control programs, although most North American wolves experience moderate levels of regulated harvest. During 1986–1992, we investigated the population dynamics and harvests of wolves in the newly created Gates of the Arctic National Park and Preserve in northern Alaska, USA, where wolves were harvested by local residents. Our objectives were to determine wolf abundance, estimate important vital rates (i.e., productivity, survival, emigration), and characterize wolf harvests. We monitored 50 radiocollared wolves in 25 packs over 4 years (Apr 1987–Apr 1991) to assess patterns of dispersal, emigration, survival and mortality causes in the wolf population. We determined pack sizes, home ranges, and pups per pack in autumn (1 Oct) for instrumented wolf packs, and calculated wolf densities in autumn and spring (15 Apr) based on the number of wolves in instrumented packs and the aggregate area those packs inhabited. We also gathered information from local hunters and trappers on the timing, location, methods, and sex–age composition of wolf harvests during 6 winter harvest seasons (Aug 1987–Apr 1992). Wolf densities averaged 6.6 wolves per 1,000 km2 and 4.5 wolves per 1,000 km2 in autumn and spring, respectively, and spring densities increased by 5% per year during our study. On average, pups constituted 50% of the resident wolf population each autumn. An estimated 12% of the population was harvested annually. Natural mortality, primarily intraspecific strife, equaled 11% per year. Young wolves emigrated from the study area at high annual rates (47% and 27% for yearlings and 2-yr-olds, respectively), and we estimated the emigration rate for the population at ≥19% annually. Yearlings and 2-year-olds were lost from the population at rates of 60% per year and 45% per year, respectively, primarily as a result of emigration; mortality was the principal cause of the 26% annual loss of wolves ≥3 years old. On average, 47 wolves were harvested each winter from our study population, or twice the harvest we estimated from survival analyses of radiocollared wolves (23 wolves/yr). We suggest that the additional harvested wolves were transients, including local dispersers and migrants from outside the study area. Trapping harvest was well-distributed throughout the trapping season (Nov–Apr), whereas shooting harvest occurred mainly in February and March. Of 35 individuals who harvested wolves in the area, 6 accounted for 66% of the harvest. We analyzed information from North American wolf populations and determined that annual rates of increase have an inverse, curvilinear relationship with human-caused mortality (r2 = 0.68, P < 0.001) such that population trends were not correlated with annual human take ≤29% (P = 0.614). We provide evidence that wolf populations compensate for human exploitation ≤29% primarily via adjustments in dispersal components (i.e., local dispersal, emigration, and immigration), whereas responses in productivity or natural mortality have little or no role in offsetting harvests. Given the limited effects of moderate levels of human take on wolf population trends and biases in assessing wolf populations and harvests resulting from the existence of transient wolves, the risks of reducing wolf populations inadvertently through regulated harvest are quite low.

Simulating the influences of various fire regimes on caribou winter habitat

Caribou are an integral component of high-latitude ecosystems and represent a major subsistence food source for many northern people. The availability and quality of winter habitat is critical to sustain these caribou populations. Caribou commonly use older spruce woodlands with adequate terrestrial lichen, a preferred winter forage, in the understory. Changes in climate and fire regime pose a significant threat to the long-term sustainability of this important winter habitat. Computer simulations performed with a spatially explicit vegetation succession model (ALFRESCO) indicate that changes in the frequency and extent of fire in interior Alaska may substantially impact the abundance and quality of winter habitat for caribou. We modeled four different fire scenarios and tracked the frequency, extent, and spatial distribution of the simulated fires and associated changes to vegetation composition and distribution. Our results suggest that shorter fire frequencies (i.e., less time between recurring fires) on the winter range of the Nelchina caribou herd in eastern interior Alaska will result in large decreases of available winter habitat, relative to that currently available, in both the short and long term. A 30% shortening of the fire frequency resulted in a 3.5-fold increase in the area burned annually and an associated 41% decrease in the amount of spruce-lichen forest found on the landscape. More importantly, simulations with more frequent fires produced a relatively immature forest age structure, compared to that which currently exists, with few stands older than 100 years. This age structure is at the lower limits of stand age classes preferred by caribou from the Nelchina herd. Projected changes in fire regime due to climate warming and/or additional prescribed burning could substantially alter the winter habitat of caribou in interior Alaska and lead to changes in winter range use and/or population dynamics.

Ranking Alaska Moose Nutrition: Signals to Begin Liberal Antlerless Harvests

Abstract We focused on describing low nutritional status in an increasing moose (Alces alces gigas) population with reduced predation in Game Management Unit (GMU) 20A near Fairbanks, Alaska, USA. A skeptical public disallowed liberal antlerless harvests of this moose population until we provided convincing data on low nutritional status. We ranked nutritional status in 15 Alaska moose populations (in boreal forests and coastal tundra) based on multiyear twinning rates. Data on age-of-first-reproduction and parturition rates provided a ranking consistent with twinning rates in the 6 areas where comparative data were available. Also, short-yearling mass provided a ranking consistent with twinning rates in 5 of the 6 areas where data were available. Data from 5 areas implied an inverse relationship between twinning rate and browse removal rate. Only in GMU 20A did nutritional indices reach low levels where justification for halting population growth was apparent, which supports prior findings that nutrition is a minor factor limiting most Alaska moose populations compared to predation. With predator reductions, the GMU 20A moose population increased from 1976 until liberal antlerless harvests in 2004. During 1997–2005, GMU 20A moose exhibited the lowest nutritional status reported to date for wild, noninsular, North American populations, including 1) delayed reproduction until moose reached 36 months of age and the lowest parturition rate among 36-month-old moose (29%, n = 147); 2) the lowest average multiyear twinning rates from late-May aerial surveys (x̄ = 7%, SE = 0.9%, n = 9 yr, range = 3–10%) and delayed twinning until moose reached 60 months of age; 3) the lowest average mass of female short-yearlings in Alaska (x̄ = 155 ± 1.6 [SE] kg in the Tanana Flats subpopulation, up to 58 kg below average masses found elsewhere); and 4) high removal (42%) of current annual browse biomass compared to 9–26% elsewhere in boreal forests. When average multiyear twinning rates in GMU 20A (sampled during 1960–2005) declined to <10% in the mid- to late 1990s, we began encouraging liberal antlerless harvests, but only conservative annual harvests of 61–76 antlerless moose were achieved during 1996–2001. Using data in the context of our broader ranking system, we convinced skeptical citizen advisory committees to allow liberal antlerless harvests of 600–690 moose in 2004 and 2005, with the objective of halting population growth of the 16,000–17,000 moose; total harvests were 7–8% of total prehunt numbers. The resulting liberal antlerless harvests served to protect the moose populations health and habitat and to fulfill a mandate for elevated yield. Liberal antlerless harvests appear justified to halt population growth when multiyear twinning rates average ≤10% and ≥1 of the following signals substantiate low nutritional status: <50% of 36-month-old moose are parturient, average multiyear short-yearling mass is <175 kg, or >35% of annual browse biomass is removed by moose.

Calf mortality and population growth in the Delta caribou herd after wolf control

Abstract A program to control wolves (Canis lupus) in interior Alaska in 1993 and 1994 did not result in expected increases in calf survival in the Delta caribou (Rangifer tarandus) herd (DCH). Therefore, the Alaska Department of Fish and Game conducted a study to determine causes of calf mortality during 1995–1997 and monitored recruitment, mortality, and population size annually in the DCH for 6 years after wolf control ended. Despite removal of 60–62% of the autumn 1993 wolf population, wolves still killed 25% of 166 radiocollared calves between birth in mid- to late May and 30 September during 1995–1997. Although autumn calf:cow ratios in the DCH increased after wolf control, similar increases in calf:cow ratios occurred in the adjacent Denali Herd, where wolves were not controlled. Calf:cow ratios following wolf control in 1993 and 1994 were lower than ratios obtained in the same area after wolf control from 1976–1982. We identified 4 factors that contributed to continued low calf:cow ratios in the DCH following the 1993–1994 wolf control program: 1) other predators in combination (i.e., golden eagles [Aquila chrysaetos] and grizzly bears [Ursus arctos]) were the most significant mortality source for caribou calves, 2) the temporal and spatial extent for wolf removal was inadequate to effectively reduce wolf predation, 3) in 1987 the DCH shifted its main calving area, a move that may have increased predation by golden eagles and grizzly bears, and 4) natality rates and nutritional condition of caribou declined during the 5 years before wolf control coincident with a density-dependent population decline. We conclude that wolf control within the range of the DCH failed because the wolf trapping program did not remove enough wolves and was not conducted long enough to substantially reduce predation by wolves on caribou calves. In addition, wolves that lived outside the control area were responsible for about 40% of the wolf-caused mortality to collared caribou calves, and significant numbers of calves died from unknown, neonatal causes.

Stochastic and Compensatory Effects Limit Persistence of Variation in Body Mass of Young Caribou

Abstract 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.

Serologic survey for canine coronavirus in wolves from Alaska

Wolves (Canis lupus) were captured in three areas of Interior Alaska (USA). Four hundred twenty-five sera were tested for evidence of exposure to canine coronavirus by means of an indirect fluorescent antibody procedure. Serum antibody prevalence averaged 70% (167/240) during the spring collection period and 25% (46/185) during the autumn collection period. Prevalence was 0% (0/42) in the autumn pup cohort (age 4–5 mo), and 60% (58/97) in the spring pup cohort (age 9–10 mo). Prevalence was lowest in the Eastern Interior study area. A statistical model indicates that prevalence increased slightly each year in all three study areas. These results indicate that transmission occurs primarily during the winter months, antibody decay is quite rapid, and reexposure during the summer is rare.

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

Abstract 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 (FST = 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.