Modeling Cumulative Effects of Climate and Development on Moose,Wolf, and Caribou Populations

Wildlife models focused solely on a single strong influence (e.g., habitat components, wildlife harvest) are limited in their ability to detect key mechanisms influencing population change. Instead, we propose integrated modeling in the context of cumulative effects assessment using multispecies population dynamics models linked to landscape-climate simulation at large spatial and temporal scales. We developed an integrated landscape and population simulation model using ALCES Online as the model-building platform, and the model accounted for key ecological components and relationships among moose (Alces alces), grey wolves (Canis lupus nubilus), and woodland caribou (Rangifer tarandus caribou) in northern Ontario, Canada. We simulated multiple scenarios over 5 decades (beginning 2020) to explore sensitivity to climate change and land use and assessed effects at multiple scales. The magnitude of effect and the relative importance of key factors (climate change, roads, and habitat) differed depending on the scale of assessment. Across the full extent of the study area (654,311km² [ecozonal scale]), the caribou population declined by 26% largely because of climate change and associated predator-prey response, which led to caribou range recession in the southern part of the study area. At the caribou range scale (108,378 km²), which focused on 2 herds in the northern part of the study area, climate change led to a 10% decline in the population and development led to an additional 7% decline. At the project scale (8,331 km²), which was focused more narrowly on the landscape surrounding 4 proposed mines, the caribou population declined by 29% largely in response to simulated development. Given that observed caribou population dynamics were sensitive to the cumulative effects of climate change, land use, interspecific interactions, and scale, insights from the analysis might not emerge under a less complex model. Our integrated modeling framework provides valuable support for broader regional assessments, including estimation of risk to caribou and Indigenous food security, and for developing and evaluating potential caribou recovery strategies. © 2021 The Authors. The Journal of Wildlife Management published by Wiley Periodicals LLC on behalf of The Wildlife Society.     

Adapt or die—Response of large herbivores to environmental changes in Europe during the Holocene

Climate warming and human landscape transformation during the Holocene resulted in environmental changes for wild animals. The last remnants of the European Pleistocene megafauna that survived into the Holocene were particularly vulnerable to changes in habitat. To track the response of habitat use and foraging of large herbivores to natural and anthropogenic changes in environmental conditions during the Holocene, we investigated carbon (δ¹³C) and nitrogen (δ¹⁵N) stable isotope composition in bone collagen of moose (Alces alces), European bison (Bison bonasus) and aurochs (Bos primigenius) in Central and Eastern Europe. We found strong variations in isotope compositions in the studied species throughout the Holocene and diverse responses to changing environmental conditions. All three species showed significant changes in their δ¹³C values reflecting a shift of foraging habitats from more open in the Early and pre‐Neolithic Holocene to more forest during the Neolithic and Late Holocene. This shift was strongest in European bison, suggesting higher plasticity, more limited in moose, and the least in aurochs. Significant increases of δ¹⁵N values in European bison and moose are evidence of a diet change towards more grazing, but may also reflect increased nitrogen in soils following deglaciation and global temperature increases. Among the factors explaining the observed isotope variations were time (age of samples), longitude and elevation in European bison, and time, longitude and forest cover in aurochs. None of the analysed factors explained isotope variations in moose. Our results demonstrate the strong influence of natural (forest expansion) and anthropogenic (deforestation and human pressure) changes on the foraging ecology of large herbivores, with forests playing a major role as a refugial habitat since the Neolithic, particularly for European bison and aurochs. We propose that high flexibility in foraging strategy was the key for survival of large herbivores in the changing environmental conditions of the Holocene.     

Parasitism of the deer ked,Lipoptena cervi,on the moose,Alces alces,in eastern Finland

The deer ked, Lipoptena cervi L. (Diptera: Hippoboscidae), is an ectoparasitic fly that spread to Finland in the early 1960s from the southeast across the Soviet border. It is currently a common parasite of the moose, Alces alces (Artiodactyla: Cervidae), in the southern part of the country and its area of distribution is gradually spreading to Finnish Lapland, where it will come into contact with another potential cervid host, the semi‐domesticated reindeer, Rangifer tarandus tarandus. The aim of this study was to determine the intensity of deer ked parasitism on the moose in eastern Finland. Whole skins of 23 moose were examined for the presence of deer keds, which were extracted and their total numbers estimated. The intensity of deer ked parasitism was correlated to the age, sex, skin area and anatomical region of the host. Bulls had the highest total number of keds (10616 ± 1375) and the highest deer ked density (35.7 ± 4.4 keds/dm² of skin). Cows had a higher total number of keds than calves (3549 ± 587 vs. 1730 ± 191), but ked densities on cows and calves were roughly equal (11.8 ± 1.7 vs. 9.4 ± 1.1 keds/dm² of skin). The density of keds was highest on the anterior back, followed by the posterior back, front limbs, abdomen, head and hind limbs. The sex ratio of deer keds was close to equal (male : female, 1.0 : 1.1). After they had consumed blood, male keds were heavier than females. As the total numbers and densities of deer keds were higher than reported previously on moose or for any other louse fly species, the effects of parasitism on the health of the host species should be determined.     

Living on the Edge: Viability of Moose in Northeastern Minnesota

ABSTRACT North temperate species on the southern edge of their distribution are especially at risk to climate-induced changes. One such species is the moose (Alces alces), whose continental United States distribution is restricted to northern states or northern portions of the Rocky Mountain cordillera. We used a series of matrix models to evaluate the demographic implications of estimated survival and reproduction schedules for a moose population in northeastern Minnesota, USA, between 2002 and 2008. We used data from a telemetry study to calculate adult survival rates and estimated calf survival and fertility of adult females by using results of helicopter surveys. Estimated age- and year-specific survival rates showed a sinusoidal temporal pattern during our study and were lower for younger and old-aged animals. Estimates of annual adult survival (when assumed to be constant for ages >1.7 yr old) ranged from 0.74 to 0.85. Annual calf survival averaged 0.40, and the annual ratio of calves born to radiocollared females averaged 0.78. Point estimates for the finite rate of increase (λ) from yearly matrices ranged from 0.67 to 0.98 during our 6-year study, indicative of a long-term declining population. Assuming each matrix to be equally likely to occur in the future, we estimated a long-term stochastic growth rate of 0.85. Even if heat stress is not responsible for current levels of survival, continuation of this growth rate will ultimately result in a northward shift of the southern edge of moose distribution. Population growth rate, and its uncertainty, was most sensitive to changes in estimated adult survival rates. The relative importance of adult survival to population viability has important implications for harvest of large herbivores and the collection of information on wildlife fertility.     

Science and Values Influencing Predator Control for Alaska Moose Management

ABSTRACT We encourage informed and transparent decision-making processes concerning the recently expanded programs in Alaska, USA, to reduce predation on moose (Alces alces). The decision whether to implement predator control ultimately concerns what society should value; therefore, policymakers, not objective biologists, play a leadership role. From a management and scientific standpoint, biological support for these predator-control programs requires convincing evidence that 1) predators kill substantial numbers of moose that would otherwise mostly live and be available for harvest, 2) low predation can facilitate reliably higher harvests of moose, 3) given less predation, habitats can sustain more moose and be protected from too many moose, and 4) sustainable populations of Alaska's brown bears (Ursus arctos), black bears (Ursus americanus), and wolves (Canis lupus) will exist in and out of control areas. We reviewed 10 moose mortality studies, 36 case histories, 10 manipulative studies, 15 moose nutrition studies, and 3 recent successful uses of nutrition-based management to harvest excess female moose. Results of these studies support application of long-term, substantial predator control for increasing yield of moose in these simple systems where moose are a primary prey of 3 effective predators. We found no substantive, contradictory results in these systems. However, to identify and administer feasible moose population objectives, recently established moose nutritional indices must be monitored, and regulatory bodies must accept nutrition-based management. In addition, the efficacy of techniques to reduce bear predation requires further study. Predicting precise results of predator control on subsequent harvest of moose will continue to be problematic because of a diversity of changing interactions among biological, environmental, and practical factors. In Alaska, the governor has the prerogative to influence regulations on predator control by appointing members to the Board of Game. At least annually, the Board of Game hears a wide spectrum of public opinions opposing and favoring predator control. We summarized these opinions as well as the societal and cultural values and expectations that are often the primary basis for debates. Advocates on both sides of the debate suggest they hold the higher conservation ethic, and both sides provide biased science. We recommend a more constructive and credible dialogue that focuses openly on values rather than on biased science and fabricated conspiracies. To be credible and to add substance in this divisive political arena, biologists must be well informed and provide complete information in an unbiased and respectful manner without exaggeration.     

Predation on Moose Calves by European Brown Bears

Abstract: In North America, brown bears (Ursus arctos) can be a significant predator on moose (Alces alces) calves. Our study in Sweden is the first in which brown bears are the only predator on moose calves. Bears and moose occurred at densities of about 30/1,000 km² and 920/1,000 km², respectively, and bears killed about 26% of the calves. Ninety-two percent of the predation took place when calves were <1 month old. Bear predation was probably additive to other natural mortality, which was about 10% in areas both with and without bears. Females that lost their calves in spring produced more calves the following year (1.54 calves/F) than females that kept their calves (1.11 calves/F), which reduced the net loss of calves due to predation to about 22%.     

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 (x = 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 population's 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.     

Bayesian spatial modeling of moose count data: increasing estimator efficiency and exploring ecological hypotheses

Moose management throughout much of Alaska and Canada relies on aerial count data, which are commonly collected using the geospatial population estimator (GSPE) protocol. The GSPE uses a model-based analytical approach and finite-population block kriging to estimate abundance from a collection of sampled survey units. Widespread implementation and well-documented analytical software have resulted in reliable estimates of moose abundance, density, and composition across a large proportion of their range. Analysis is conducted almost exclusively using the GSPE software, which fits a fixed model structure to data collected within a single year. The downside of this approach to analysis is that the fixed model structure is inefficient for estimation, leading to more field effort than would otherwise be necessary to achieve a desired level of estimator precision. We developed a more easily modified and flexible Bayesian spatial general additive model approach (BSG) that accommodates spatial and temporal covariates (e.g., habitat characteristics, trend), multiple survey events, prior information, and incomplete detection. Using a series of 6 GSPE surveys conducted in Yukon-Charley Rivers National Preserve, Alaska, USA, from 2003–2019, we established the equivalence of the 2 approaches under similar model structures. We then extended the BSG to demonstrate how a more comprehensive approach to analysis can affect estimator precision and be used to assess ecological relationships. The precision of annual abundance estimators from the BSG were improved by an average of 43% over those based on the standard GSPE analysis, highlighting the very real costs of assuming a fixed (i.e., suboptimal) model structure. The population increased at a rate of 2.3%/year (95% CrI = 0.8–3.8%), and the increase was largely explained by a parallel increase in wildfire extent (i.e., high quality moose habitat). These results suggest that our approach could be used to increase estimator efficiency or decrease future survey costs without any modifications to the basic protocol. While modification of the GSPE software is possible, practitioners may find the BSG approach more convenient for quickly developing model structures for a particular application, thereby allowing them to extract more information from existing and future datasets.     

Effects of willow nutrition and morphology on calving success of moose

Across much of North America, populations of moose (Alces alces) are declining because of disease, predation, climate change, and anthropogenic-driven habitat loss. Contrary to this trend, populations of moose in Colorado, USA, have continued to grow. Studying successful (i.e., persistent or growing) populations of moose can facilitate continued conservation by identifying habitat features critical to persistence of moose. We hypothesized that moose using habitat with higher quality willow (Salix spp.) would have a higher probability of having a calf-at-heel (i.e., calving success). We evaluated moose calving success using repeated ground observations of collared individuals with calves in an occupancy model framework to account for detection probability. We then evaluated the impact of willow habitat quality and nutrition on moose calving success by studying 2 spatially segregated populations of moose in Colorado. Last, we evaluated correlations between willow characteristics (browse intensity, height, cover, leaf length, and species) and willow nutrition (dry matter digestibility [DMD]) to assess the utility of using those characteristics to assess willow nutrition. We found willow height and cover had a high probability of being positively associated with higher individual-level calving success. Willow DMD, browse intensity, and leaf length were not predictive of individual moose calving success; however, the site with higher mean DMD consistently had higher mean estimates of calving success for the same year. Our results suggest surveying DMD is likely not a useful metric for assessing differences in calving success of individual moose but may be of use at population levels. Further, the assessment of willow morphology and density may be used to identify areas that support higher levels of moose calving success.