Francis J. Singer

Density dependence, compensation, and environmental effects on elk calf mortality in Yellowstone National Park

We studied survival of radiocollared elk (Cervus elaphus) calves in Yellowstone National Park from 1987 to 1990, and survival of calves computed from population estimates from 1968 to 1992. We hypothesized that summer and winter survival of elk calves and mass of neonates were inversely related to population size, measures of environmental severity, and timing of births. Herd-wide survival estimates based on winter counts, reported harvests, and herd classifications, suggested that winter survival of elk calves was related inversely to estimated size of the elk population during winter (P = 0.0002), but we found no correlation with an index of winter severity (P = 0.51). Summer survival of elk calves also was correlated inversely with the estimated size of the elk population the previous winter (P = 0.03). Summer survival of radiocollared calves averaged 0.65 (n = 127 marked calves) from 1987 to 1990, the losses mostly due to predation (22%). Winter survival of calves averaged 0.72 (n = 88 marked calves entered the winter period), with losses due mostly to malnutrition (58%). Summer survival of radiocollared calves was positively correlated with estimated birth weight (P = 0.001). Survival of radiocollared calves during winters 1987-90 was correlated positively with early calving and mildness of the winter (in contrast to herd-wide survival estimates), and was inversely correlated with estimated elk population size that winter (P = 0.006). Winter survival of radiocollared calves was lower during 1988-89 following the drought and large fires than the other 3 winters (P < 0.001). Predation on elk calves during summer doubled after the drought and fires of 1988 (13% calf losses to predation before the fires vs. 29% after the fires). Potential compensation existed between components of calf mortality: predators killed more light (P = 0.041) and more late born calves (P = 0.146); calves were born later and lighter (P = 0.048) following severe weather conditions; and heavier born calves survived at a higher rate (P = 0.006). Our results are consistent with the hypothesis that density-dependent mortality of calves during winter due to malnutrition, and summer mortality of calves due to predation were partially compensatory but severe environmental conditions produced largely additive components to both summer (increased predation) and winter (increased malnutrition) mortality.

Effects of wild pig rooting in a deciduous forest

Action des porcs sauvages sur la faune superficielle, les elements nutritifs, la biomasse de la litiere forestiere et du sol

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.

Responses of desert bighorn sheep to increased human recreation.

Human recreation has been implicated in the decline of several populations of desert bighorn sheep (Ovis is canadensis nelsoni). Managers are concerned about the impact of increased recreation on desert bighorn sheep in Canyonlands National Park (NP), Utah, USA, where visitation increased 325% from 1979 to 1994. We compared behavioral responses of sheep to recreational activity between a low visitor use area and a high visitor use area during 1993 and 1994 by observing behavioral responses, distances moved, and duration of responses to vehicles, mountain bikers, and humans on foot. Hikers caused the most severe responses in desert bighorn sheep (animals fled in 61% of encounters), followed by vehicles (17% fled) and mountain bikers (6% fled), apparently because hikers were more likely to be in unpredictable locations and often directly approached sheep. We observed considerable individual heterogeneity in responses of bighorn sheep to the greater human use: some animals lived close to the road corridor and were apparently habituated to the human activities, but other animals avoided the road corridor. In the high-use area, we observed 3 radiocollared sheep that lived closer to the road than expected and found evidence of fewer responses to vehicles by females in spring, less response time of all sheep to vehicles in spring, and fewer responses to mountain bikers compared to the low-use area. Overall, there was an avoidance of the road corridor by most other bighorn sheep in the high-use area where all animals, on average, were found 39% farther from roads (490 ± 19 m vs. 354 ± 36 m) than in the low-use area. This avoidance of the road corridor by some animals represented 15% less use of potential suitable habitat in the high-use area over the low-use area. Increased sensitivity to hikers in the high-use area was suggested by a greater responsiveness by males in autumn and greater distance fled by females in spring. Responses of bighorn sheep were greater when human activity approached at the same elevation, when sheep were moving or standing, when female interactions occurred in spring and summer and male interactions occurred in autumn, and when sheep were farther from escape terrain. We recommend managers confine hikers to designated trails during spring lambing and the autumn rut in desert bighorn sheep habitat.

Dynamics of interacting Elk populations within and adjacent to Rocky Mountain National Park

We studied population subdivision and density-dependent and independent factors influencing population processes between 1965 and 2001 for elk (Cervus elaphus) inhabiting Rocky Mountain National Park (park) and the adjacent Estes Valley (town), Colorado, USA. Elk numbers within the park were held relatively constant by management controls until 1967, after which time they were allowed to increase without human interference. Radiotelemetry of 73 elk indicated limited exchange between the subpopulations; combined with clear distinctions in population dynamics, this suggests that these subpopulations are relatively independent despite the absence of physical barriers between them. The elk subpopulation within the park initially increased at 6.5%/year between 1968 and 1970, then growth gradually slowed-exhibiting density-dependent reductions both in calf survival and recruitment with increasing population size-and approached an estimated carrying capacity of 1,069 ± 55 (?±SE). Since 1991, this subpopulation has remained within ±5% of this equilibrium. The adjacent Estes Valley subpopulation grew at an estimated maximum 5-year average rate of 11.0% from 1979 to 1983 and is still increasing at 5.2%/year (1991-2001 average). Estimated town population currently is about 70% of our projected carrying capacity of 2,869 ±415 elk based on projection of observed calf recruitment decline with increasing population. Both carrying-capacity estimates are consistent with independent estimates based on forage biomass and energy considerations. Adult cow survival rate was not found to differ between park and town, and we estimated a constant rate of 0.913 [95% CI = 0.911, 0.915]. Bull survival rates increased in the park from 0.52 to 0.79 between 1965 and 2001, but remained constant at 0.42 [0.35, 0.47] in the Estes Valley. Colder winter temperatures were correlated with reduced calf recruitment (calves:cow at age 0.5 yr) and with reduced calf survival (between age 0.5 and 1.5 yr) in town. Recruitment of town elk also increased with warmer summer temperatures and greater summer precipitation. No weather covariates were significantly correlated with calf recruitment or survival in the park. Declining calf recruitment has been nearly linear and similar in both the park and town. In the park, calf survival responded little to density when the population was well below carrying capacity, but responded at an increasing rate as the population neared carrying capacity. This pattern may explain why calf survival response to density has not yet been detected in town. We estimated current combined population size of 3,049 [2759, 3369] elk in 2001. Elk in the town sector currently outnumber elk in the adjacent national park by almost 2:1 and are projected to increase by 46% before being nutritionally limited, suggesting that human-elk conflicts likely will increase in the absence of active management intervention.

Behavior of Mountain Goats in Relation to U.S. Highway 2, Glacier National Park, Montana

A study was conducted in 1975 on mountain goats (Oreamnos americanus) crossing a highway to visit a natural mineral lick. Eighty-seven successful crossings of the highway involving 692 mountain goats were observed, and an estimated 812 crossings occurred during the 1975 lick season. Crossing success was associated (P

Influences of herbivory and water on willow in elk winter range

Elimination of large predators and reduced hunter harvest have led to concerns that an increasing ell (Cervus elaphus) population may be adversely affecting vegetation on the low-elevation elk winter range of Rock Mountain National Park, Colorado, USA. Beaver (Castor canadensis) and their impoundments also have declined dramatically (94%) in the same area over the past 50 years coincident with a 20% decline in willow (Salix spp.) cover. From 1994 to 1998, we studied vegetation production responses of willow communities to elk herbivory and water availability. We estimated willow production by measuring current annual growth of shrubs in 9.3-m 2 circu lar plots, and we measured herbaceous production by clipping vegetation within 0.25-m 2 circular plots. Elk her bivory suppressed willow heights, leader lengths, annual production, and herbaceous productivity of willow corn munities. Water impoundment had a positive effect on herbaceous plant production, but little effect on shrubs, possibly because water tables were naturally high on the study sites even without beaver dams. Nevertheless, the winter range environment previously included more riparian willow habitat because of more stream area (47-69%) due to larger beaver populations. Elk herbivory appears to be the dominant force determining vegetation pro ductivity in willow sites, but the effects may he exacerbated by lowered water tables. Fewer elk or protection from browsing, and water enhancement for <10 years along with management to encourage elk movement away from willow communities, could possibly work as strategies to reestablish sustainable willow communities.

Physiological Assessment of Winter Nutritional Deprivation in Elk of Yellowstone National Park

During 13 January-29 March 1988, we assessed the extent of nutritional deprivation in cow elk (Cervus elaphus nelsoni) groups on the lower, middle, and upper Northern Range and on the Madison-Firehole Range in Yellowstone National Park by 4 sequential collections and chemical analyses of urine excreted in snow (snow-urine). Associated changes in elk density and calf : cow ratios also were estimated during early and late winter or spring. Decreasing (P=0.0001) potassium : creatinine (K:C) ratios throughout winter and increased (P=0.0001) urea nitrogen : creatinine (U:C) on all 4 areas indicated progressive nutritional deprivation and increasing net catabolism of lean body tissue, respectively. These ratios also varied among the 4 sampling areas during early (P=0.0001) and late winter (P<0.005)

Physiological responses of Yellowstone bison to winter nutritional deprivation

Because nutrition is critically related to other aspects of bison (Bison bison) ecology, and the winter ranges inhabited by bison in Yellowstone National Park (YNP) are ecologically diverse, it was important to determine if nutritional deprivation differences occurred among winter ranges. We used chemistry profiles of urine suspended in snow to compare nutritional deprivation of bison from january to April 1988 on 4 sampling areas of 3 winter ranges in YNP. Declining (P < 0.001) trends of urinary potassium: creatinine ratios in bison on all 4 sampling areas indicated progressive nutritional deprivation through late March. Concurrent increases (P ≤ 0.001) in mean urea nitrogen: creatinine ratios from late February through late March in 3 of 4 areas suggested that increased net catabolism was occurring

Activity in a hunted and an unhunted herd of Dall sheep

Abstract In three sampling areas (two unhunted, one hunted), we compared activity patterns of Dall sheep rams (Ovis dalli) of different horn size during the rut. In all three areas rams with horns 3 4 - curled and smaller rams were less frequently seen in ewe bands as the rut progressed. In one area where we documented activity budgets, we observed equivalent budgets for ewes and rams with horns 1 4 - curled . Older rams spent less time foraging than young rams, except that rams with horns 5 8 – 7 8 curled spent less time foraging than full-curl rams late in the rut. Display frequencies of 7 8 - curl rams were similar to those of full-curl rams and considerably higher than those of smaller-horned rams early in the rut. These results suggest that the rut is more energy costly for old rams than for young rams, but that it may be most costly for the next-to-largest horn-curl class rams (e.g. 7 8 - curl rams in unhunted populations) rather than the dominant rams (full-curl and larger). Rams in the next-to-largest horn-curl class in the hunted herd, however, did not acquire the higher display rates (e.g., 3 4 - curl rams had equally low display rates in both the hunted and unhunted herds).