Shannon M. Barber-Meyer

Elk Calf Survival and Mortality Following Wolf Restoration to Yellowstone National Park

Abstract We conducted a 3-year study (May 2003–Apr 2006) of mortality of northern Yellowstone elk (Cervus elaphus) calves to determine the cause for the recruitment decline (i.e., 33 calves to 13 calves/100 adult F) following the restoration of wolves (Canis lupus). We captured, fit with radiotransmitters, and evaluated blood characteristics and disease antibody seroprevalence in 151 calves ≤6 days old (68M:83F). Concentrations (𝑥̄, SE) of potential condition indicators were as follows: thyroxine (T4; 13.8 μg/dL, 0.43), serum urea nitrogen (SUN; 17.4 mg/dL, 0.57), γ-glutamyltransferase (GGT; 66.4 IU/L, 4.36), gamma globulins (GG; 1.5 g/dL, 0.07), and insulin-like growth factor-1 (IGF-1; 253.6 ng/mL, 9.59). Seroprevalences were as follows: brucellosis (Brucella abortus; 3%), bovine-respiratory syncytial virus (3%), bovine-viral-diarrhea virus type 1 (25%), infectious-bovine rhinotracheitis (58%), and bovine parainfluenza-3 (32%). Serum urea nitrogen, GGT, GG, and IGF-1 varied with year; T4, SUN, and GG varied with age (P ≤ 0.01); and SUN varied by capture area (P = 0.02). Annual survival was 0.22 (SE = 0.035, n = 149) and varied by calving area but not year. Neonates captured in the Stephens Creek/Mammoth area of Yellowstone National Park, USA, had annual survival rates >3× higher (0.54) than those captured in the Lamar Valley area (0.17), likely due to the higher predator density in Lamar Valley. Summer survival (20 weeks after radiotagging) was 0.29 (SE = 0.05, n = 116), and calving area, absolute deviation from median birth date, and GG were important predictors of summer survival. Survival during winter (Nov–Apr) was 0.90 (SE = 0.05, n = 42), and it did not vary by calving area or year. Sixty-nine percent (n = 104) of calves died within the first year of life, 24% (n = 36) survived their first year, and 7% (n = 11) had unknown fates. Grizzly bears (Ursus arctos) and black bears (Ursus americanus) accounted for 58–60% (n = 60–62) of deaths, and wolves accounted for 14–17% (n = 15–18). Summer predation (95% of summer deaths) increased, and winter malnutrition (0% of winter deaths) decreased, compared with a similar study during 1987–1990 (72% and 58%, respectively). Physiological factors (e.g., low levels of GG) may predispose calves to predation. Also, the increase in bear numbers since wolf restoration and spatial components finer than the northern range should be considered when trying to determine the causes of the northern Yellowstone elk decline. This is the first study to document the predation impacts from reintroduced wolves on elk calf mortality in an ecosystem already containing established populations of 4 other major predators (i.e., grizzly and black bears, cougars [Puma concolor], and coyotes [Canis latrans]). The results are relevant to resource managers of the Yellowstone ecosystem in understanding the dynamics of the elk population, in providing harvest quota recommendations for local elk hunts to the Montana Department of Fish, Wildlife and Parks, the United States Fish and Wildlife Service regarding wolf and grizzly bear recovery, and to all areas worldwide where predators are increasing, by providing managers with information about potential carnivore impacts on elk populations.

Neonatal mortality of elk driven by climate, predator phenology and predator community composition

1. Understanding the interaction among predators and between predation and climate is critical to understanding the mechanisms for compensatory mortality. We used data from 1999 radio-marked neonatal elk (Cervus elaphus) calves from 12 populations in the north-western United States to test for effects of predation on neonatal survival, and whether predation interacted with climate to render mortality compensatory. 2. Weibull survival models with a random effect for each population were fit as a function of the number of predator species in a community (3-5), seven indices of climatic variability, sex, birth date, birth weight, and all interactions between climate and predators. Cumulative incidence functions (CIF) were used to test whether the effects of individual species of predators were additive or compensatory. 3. Neonatal elk survival to 3 months declined following hotter previous summers and increased with higher May precipitation, especially in areas with wolves and/or grizzly bears. Mortality hazards were significantly lower in systems with only coyotes (Canis latrans), cougars (Puma concolor) and black bears (Ursus americanus) compared to higher mortality hazards experienced with gray wolves (Canis lupus) and grizzly bears (Ursus horribilis). 4. In systems with wolves and grizzly bears, mortality by cougars decreased, and predation by bears was the dominant cause of neonatal mortality. Only bear predation appeared additive and occurred earlier than other predators, which may render later mortality by other predators compensatory as calves age. Wolf predation was low and most likely a compensatory source of mortality for neonatal elk calves. 5. Functional redundancy and interspecific competition among predators may combine with the effects of climate on vulnerability to predation to drive compensatory mortality of neonatal elk calves. The exception was the evidence for additive bear predation. These results suggest that effects of predation by recovering wolves on neonatal elk survival, a contentious issue for management of elk populations, may be less important than the composition of the predator community. Future studies would benefit by synthesizing overwinter calf and adult-survival data sets, ideally from experimental studies, to test the roles of predation in annual compensatory and additive mortality of elk.

Testing global positioning system telemetry to study wolf predation on deer fawns

Abstract We conducted a pilot study to test the usefulness of Global Positioning System (GPS) collars for investigating wolf (Canis lupus) predation on white-tailed deer (Odocoileus virginianus) fawns. Using GPS collars with short location-attempt intervals on 5 wolves and 5 deer during summers 2002–2004 in northeastern Minnesota, USA, demonstrated how this approach could provide new insights into wolf hunting behavior of fawns. For example, a wolf traveled ≥1.5–3.0 km and spent 20–22 hours in the immediate vicinity of known fawn kill sites and ≥0.7 km and 8.3 hours at scavenging sites. Wolf travel paths indicated that wolves intentionally traveled into deer summer ranges, traveled ≥0.7–4.2 km in such ranges, and spent <1–22 hours per visit. Each pair of 3 GPS-collared wolf pack members were located together for ≤6% of potential locations. From GPS collar data, we estimated that each deer summer range in a pack territory containing 5 wolves ≥1 year old and hunting individually would be visited by a wolf on average every 3–5 days. This approach holds great potential for investigating summer hunting behavior of wolves in areas where direct observation is impractical or impossible.

Survey of Selected Pathogens and Blood Parameters of Northern Yellowstone Elk: Wolf Sanitation Effect Implications

ABSTRACT The restoration or conservation of predators could reduce seroprevalences of certain diseases in prey if predation selectively removes animals exhibiting clinical signs. We assessed disease seroprevalences and blood parameters of 115 adult female elk (Cervus elaphus) wintering on the northern range of Yellowstone National Park [YNP] during 2000–2005 and compared them to data collected prior to wolf (Canis lupus) restoration (WR) in 1995 and to two other herds in Montana to assess this prediction. Blood parameters were generally within two standard deviations of the means observed in other Montana herds (Gravelly-Snowcrest [GS] and Garnet Mountain [GM]), but Yellowstone elk had higher seroprevalences of parainfluenza-3 virus (95% CI YNP  =  61.1–78.6, GS  =  30.3–46.5) and bovine-virus-diarrhea virus type 1 (95% CI YNP  =  15.9–31.9, GM  =  0). In comparisons between pre-wolf restoration [pre-WR] (i.e., prior to 1995) seroprevalences with those post-wolf restoration [post-WR] in Yellowstone, we found lower seroprevalences for some disease-causing agents post-wolf restoration (e.g., bovine-virus-diarrhea virus type-1 [95% CI pre-WR  =  73.1–86.3, post-WR  =  15.9–31.9] and bovine-respiratory syncytial virus [95% CI pre-WR  =  70.0–83.8, post-WR  =  0]), but similar (e.g., Brucella abortus [95% CI pre-WR  =  0–4.45, post-WR  =  0–4.74] and epizootic hemorrhagic disease virus [95% CI pre-WR  =  0, post-WR  =  0]) or higher for others (e.g., Anaplasma marginale [95% CI pre-WR  =  0, post-WR  =  18.5–38.7] and Leptospira spp. [95% CI pre-WR  =  0.5–6.5, post-WR  =  9.5–23.5]). Though we did not detect an overall strong predation effect through reduced disease seroprevalence using retrospective comparisons with sparse data, our reference values will facilitate future assessments of this issue.

Wolf (Canis lupus) Generation Time and Proportion of Current Breeding Females by Age

Information is sparse about aspects of female wolf (Canis lupus) breeding in the wild, including age of first reproduction, mean age of primiparity, generation time, and proportion of each age that breeds in any given year. We studied these subjects in 86 wolves (113 captures) in the Superior National Forest (SNF), Minnesota (MN), during 1972–2013 where wolves were legally protected for most of the period, and in 159 harvested wolves from throughout MN wolf range during 2012–2014. Breeding status of SNF wolves were assessed via nipple measurements, and wolves from throughout MN wolf range, by placental scars. In the SNF, proportions of currently breeding females (those breeding in the year sampled) ranged from 19% at age 2 to 80% at age 5, and from throughout wolf range, from 33% at age 2 to 100% at age 7. Excluding pups and yearlings, only 33% to 36% of SNF females and 58% of females from throughout MN wolf range bred in any given year. Generation time for SNF wolves was 4.3 years and for MN wolf range, 4.7 years. These findings will be useful in modeling wolf population dynamics and in wolf genetic and dog-domestication studies.

Weekly Summer Diet of Gray Wolves (Canis lupus) in Northeastern Minnesota

Abstract Wolves (Canis lupus) are opportunistic predators and will capitalize on available abundant food sources. However, wolf diet has primarily been examined at monthly, seasonal, or annual scales, which can obscure short-term responses to available food. We examined weekly wolf diet from late June to early October by collecting scats from a single wolf pack in northeastern Minnesota. During our 15 wk study, nonungulate food types constituted 58% of diet biomass. Deer (Odocoileus virginianus) fawns were a major food item until mid-July after which berries (primarily Vaccinium and Rubus spp.) composed 56–83% of weekly diet biomass until mid-August. After mid-August, snowshoe hares (Lepus americanus) and adult deer were the primary prey. Weekly diet diversity approximately doubled from June to October as wolves began using several food types in similar proportions as the summer transitioned into fall. Recreational hunting of black bears (Ursus americanus) contributed to weekly wolf diet in the fall as wolves consumed foods from bear bait piles and from gut piles/carcasses of successfully harvested or fatally wounded bears. To our knowledge, we are the first to examine wolf diet via scat analysis at weekly intervals, which enabled us to provide a detailed description of diet plasticity of this wolf pack, as well as the rapidity with which wolves can respond to new available food sources.

Differential wolf-pack-size persistence and the role of risk when hunting dangerous prey

Risk to predators hunting dangerous prey is an emerging area of research and could account for possible persistent differences in gray wolf (Canis lupus) pack sizes. We documented significant differences in long-term wolf-pack-size averages and variation in the Superior National Forest (SNF), Denali National Park and Preserve, Yellowstone National Park, and Yukon, Canada (p<0.01). The SNF differences could be related to the wolves’ risk when hunting primary prey, for those packs (N=3) hunting moose (Alces americanus) were significantly larger than those (N=10) hunting white-tailed deer (Odocoileus virginianus) (F1,8=16.50, p=0.004). Our data support the hypothesis that differential pack-size persistence may be perpetuated by differences in primary prey riskiness to wolves, and we highlight two important extensions of this idea: (1) the potential for wolves to provision and defend injured packmates from other wolves and (2) the importance of less-risky, buffer prey to pack-size persistence and year-to-year variation.

Trophic cascades from wolves to grizzly bears or changing abundance of bears and alternate foods

This is a Forum article commenting on: Ripple, W. J., Beschta, R. L., Fortin, J. K., & Robbins, C. T. (2014) Trophic cascades from wolves to grizzly bears in Yellowstone. Journal of Animal Ecology, 83, 223-233. Comparisons Ripple et al. (2014) used to demonstrate increased fruit availability and consumption by grizzly bears post-wolf reintroduction are flawed and tenuous at best. Importantly, a more parsimonious (than trophic cascades) hypothesis, not sufficiently considered by Ripple et al., exists and is better supported by available data I review.

INTERLEUKIN-6 AND TUMOR NECROSIS FACTOR-ALPHA VALUES IN ELK NEONATES

Abstract Serological indicators of general condition would be helpful for monitoring or assessing ungulate wildlife. Toward that end, we report the 1st reference values for 2 cytokines, interleukin-6 (IL-6) and tumor necrosis factor-alpha (TNF-α), in neonatal elk (Cervus elaphus). We obtained blood samples from 140 calves ≤ 6 days old in Yellowstone National Park during summer 2003–2005. IL-6 values ranged from 0 to 1.21 pg/ml with a median of 0.03 pg/ml. TNF-α values ranged from 0 to 225.43 pg/ml with a median of 1.85 pg/ml. IL-6 and TNF-α concentrations were not significant predictors of elk calf survival through 21 days. Development of ungulate-based IL-6 and TNF-α assays that provide greater sensitivity than cross-reacting human-based assays could be helpful in monitoring ungulate condition and health status comparisons among herds. Such information could provide indirect assessments of range quality or environmental influences among herds.

Evaluation of a Formula that Categorizes Female Gray Wolf Breeding Status by Nipple Size

Abstract The proportion by age class of wild Canis lupus (Gray Wolf) females that reproduce in any given year remains unclear; thus, we evaluated the applicability to our long-term (1972–2013) data set of the Mech et al. (1993) formula that categorizes female Gray Wolf breeding status by nipple size and time of year. We used the formula to classify Gray Wolves from 68 capture events into 4 categories (yearling, adult non-breeder, former breeder, current breeder). To address issues with small sample size and variance, we created an ambiguity index to allow some Gray Wolves to be classed into 2 categories. We classified 20 nipple measurements ambiguously: 16 current or former breeder, 3 former or adult non-breeder, and 1 yearling or adult non-breeder. The formula unambiguously classified 48 (71%) of the nipple measurements; based on supplemental field evidence, at least 5 (10%) of these were incorrect. When used in conjunction with an ambiguity index we developed and with corrections made for classifications involving very large nipples, and supplemented with available field evidence, the Mech et al. (1993) formula provided reasonably reliable classification of breeding status in wild female Gray Wolves.