Geology topics
Publications and source records attributed to Layne G. Adams.
The killing of a cow Muskox ( Ovibos moschatus ) by two Wolves ( Canis lupus ) in 5 minutes during summer on Ellesmere Island is described. After two of the four feedings observed, one Wolf cached a leg and regurgitated food as far as 2.3 km away and probably farther. The implications of this behavior for deriving food-consumption estimates are discussed.
We tested prototype GPS collars on 8 free-ranging wolves ( Canis lupus ) and 3 white-tailed deer ( Odocoileus virginianus ) for varying periods between February and August 1997. We programmed the 920-gm collars to make a location attempt 6-96 times per day. The collars were designed to be remotely released from the animal and the data were then downloaded to a desktop computer. The collars produced 47-1,549 locations each during 11-41 days; locations were successful in 26-95% of the attempts (x̄ = 70%). Eight collars released successfully. Three collar-release failures were caused by condensation. Two collars had GPS antennas that were improperly attached and did not collect data. Life was as long as, or longer than, expected in 4 collars, less than expected in 5 collars, and unknown in 2 collars. Limitations of this type of collar include brief life if programmed at short location-attempt intervals (≤1 hr) and possible drop-off failure. Nevertheless, the large volume of data we collected with no field telemetry effort demonstrates the potential for this type of GPS collar to answer questions about movements of medium-sized mammals.
Cardiac chamber enlargement and hypertrophy are normal physiologic responses to repetitive endurance exercise activity in human beings and domestic dogs. Whether similar changes occur in wild animals as a consequence of increased activity is unknown. We found that free-ranging gray wolves ( Canis lupus , n =11), the archetypical endurance athlete, have electrocardiographic evidence of cardiac chamber enlargement and hypertrophy relative to sedentary captive gray wolves ( n =20), as demonstrated by significant increases in QRS duration, QT interval, and QT interval corrected for heart rate, a tendency towards increased Q, R, and S wave voltages in all leads, and a significant decrease in heart rate. We conclude that exercise activity level and therefore lifestyle affects physiologic variables in wild animals. An immediate consequence of this finding is that physiologic measurements obtained from a captive wild-animal population with reduced exercise activity level may not accurately reflect the normal physiologic state for free-ranging members of the same species.
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.
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.
Wolf packs generally consist of a breeding pair and their maturing offspring that help provision and protect pack young. Because the reproductive tenure in wolves is often short, reproductively mature offspring might replace their parents, resulting in sibling or parent-offspring matings. To determine the extent of incestuous pairings, we measured relatedness based on variability in 20 microsatellite loci of mated pairs, parent-offspring pairs, and siblings in two populations of gray wolves. Our 16 sampled mated pairs had values of relatedness not overlapping those of known parent-offspring or sibling dyads, which is consistent with their being unrelated or distantly related. These results suggest that full siblings or a parent and its offspring rarely mate and that incest avoidance is an important constraint on gray wolf behavioral ecology.
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 < 0.001) neonatal survival than did other calves. Grizzly bears (Ursus arctos), wolves (Canis lupus), and unknown large predators (i.e., grizzly bears or wolves) accounted for 49, 29, and 16% of the neonatal deaths, respectively. The rate of bear-caused mortalities declined (P < 0.001) with calf age, and bears killed few calves >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.
Management of gray wolves ( Canis lupus ) and their prey in interior Alaska has been controversial for three decades (Harbo and Dean 1983). Recently, debate was rekindled with renewed interest in wolf control to bolster two populations of caribou ( Rangifer tarandus ). Our research in Denali National Park provides insights into the declines in caribou numbers over the last few years that are the basis of recent wolf control proposals. Our observations of fluctuating populations also illustrate the complexity of managing these predator-prey systems to meet a diverse array of public interests. Wolves and caribou are two components of the large mammal community of Denali National Park that also includes grizzly bears ( Ursus arctos ), moose ( Alces alces ), and Dall sheep ( Ovis dalli ). With the 1980 park expansion to more than 18,800 km2 (7,300 mi2) of central Alaska, this large mammal system became the only one of its kind that is virtually unaffected by human harvest. Therefore, Denali provides a unique opportunity to understand the natural interactions of these species and serves as a baseline for comparison with areas where hunting or other active wildlife management occurs. We have studied Denali's wolves and caribou since 1986 to determine their numbers and status and understand their natural interactions in this protected subarctic ecosystem. Our studies began near the end of more than a decade of mostly light winter snowfalls of around 100 cm (39 in)/yr. Since winter 1988-89, we have experienced five consecutive winters with above-average snowfalls, including two record-setting years. During winters 1990-91 and 1992-93, more than 390 cm (154 in) of snow fell, four times as much as in the early years of our study. This change in snowfall had profound effects on the wildlife in central Alaska. The population trends of Denali's caribou and wolves are strong evidence of the natural fluctuations to be expected in species inhabiting such dynamic and variable environments.
The patterns of selection by wolves ( Canis lupu s) preying on moose ( Alces alces ), caribou ( Rangifer tarandus ), and Dall sheep ( Ovis dalli) in Denali National Park and Preserve, Alaska were studied from 1986 through early 1992. Wolves and their prey are legally protected or relatively unharvested in most of the area, and wolf numbers doubled during the study. Based on remains of 294 moose, 225 caribou, and 63 sheep, wolves killed calves and old adults disproportionately, and individuals with low marrow fat, jaw necrosis, or arthritis. Seasonal trends in proportions of various species, ages, and sex of kills were found. During the winters following winters of deep snowfalls, wolves greatly increased the proportion of caribou cows and calves taken. We conclude that in a natural system, wolves can survive on vulnerable prey even during moderate weather, and when snowfall exceeds average, they can respond by switching to newly vulnerable prey and greatly increasing their numbers.
Observations of wolf pack dynamics over a six-year period in Denali National Park and Preserve, Alaska, found high rates of intraspecific strife, wolf pack dissolution and new pack formation, and the acceptance of new wolves into established packs. These observations corroborate genetic studies that found more genetic links between packs, and more genetic diversity within packs, than would be expected if most packs were composed of an unrelated breeding pair and their offspring. Longevity of packs, stability of pack territories, and the incidence of inbreeding all appear to be less than previously suggested, even in the absence of significant human disturbance. The formation of new packs by two or more local dispersers, the acceptance of unrelated wolves into existing packs, and the presence of multiple breeding females within packs would tend to blur genetic distinctions between the packs in a population.
We investigated patterns of winter wolf predation, including prey selection, prey switching, kill rates, carcass utilization, and consumption rates for four wolf packs during three different study periods (March 1989, March 1990, and November 1990) in Gates of the Arctic National Park and Preserve, Alaska. Wolves killed predominantly caribou (165 caribou, seven moose, and five Dall sheep) even when moose and sheep were more abundant. Prey selection varied between study periods. More moose were killed in march 1989, a particularly deep snow year, and more sheep were killed in November 1990 than during other periods. Overall kill rates ranged from 0-8 days/ungulate killed ( x̅ = 2.0, SD = 1.6) and did not vary between study periods. Pack size and species killed explained significant variation in the length of time intervals between kills. Although caribou density varied nearly 40-fold between pack territories, it had little influence on predation characteristics except at low densities, when kill rates may have declined. Caribou distribution had marked effects on wolf predation rate.
During 1987-1991, 29 to 45 radio-collared caribou cows were monitored daily during calving each year and their calves were radio-collared (n = 147 calves) to investigate calf production and survival. We determined characteristics of wolf predation on caribou calves and, utilizing information from a companion wolf study, evaluated the role of spacing by caribou cows in minimizing wolf predation on neonates (calves < 15 days old) during a period when wolf abundance doubled. On average, 49% of the neonates died, ranging from 30% in 1987 to 71% in 1991. Overall, wolves killed 22% of the neonates produced and were the most important mortality agent. Wolves preyed on calves primarily during six days following the peak of calving and usually killed calves five to 15 days old. The mortality rate for neonates was strongly inversely correlated with average birthweight. Neonatal losses to wolves were also correlated with birthweight but not spring wolf density or mean calving elevation. Caribou concentrated on a calving ground when spring snow conditions allowed and adjusted their distribution on the calving ground depending on snow conditions and wolf distribution and abundance. Even though the wolf population doubled, the exposure of caribou calves to wolf predation did not increase, when spacing by caribou at the wolf pack territory scale was accounted for.
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, 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 numerical response, implicates an anti-regulatory effect of wolf predation on barren-ground caribou dynamics. 4. There was little potential for regulation caused by the multiplicative effect of increasing functional and numerical responses because of presence of alternative prey. This resulted in high wolf:caribou ratios at low prey densities which precluded the effects of an increasing functional response. 5. Inversely density-dependent predation by other predators, such as bears, reduces the potential for predators to regulate caribou populations at low densities, and small reductions in predation by one predator may have disproportionately large effects on the total predation rate.