Tradeoffs Among Predator Control, Moose Harvests, and Trophy Antlers: Principles Pertinent to Managing Alaska's Wildlife.

Predation from large carnivores and human harvest are the two main mortality factors affecting the dynamics of many ungulate populations. We examined long-term moose (Alces alces) harvest data from two countries that share cross-border populations of wolves (Canis lupus) and their main prey moose. We tested how a spatial gradient of increasing wolf territory density affected moose harvest density and age and sex composition of the harvested animals (n = 549,310), along a latitudinal gradient during 1995–2017. In areas containing average-sized wolf territories, harvest density was on average 37% (Norway) and 51% (Sweden) lower than in areas without wolves. In Sweden, calves made up a higher proportion of the moose harvest than in Norway, and this proportion was reduced with increased wolf territory density, while it increased in Norway. The proportion of females in the adult harvest was more strongly reduced in Sweden than in Norway as a response to increased wolf territory density. Moose management in both countries performed actions aimed to increase productivity in the moose population, in order to compensate for the increased mortality caused by wolves. These management actions are empirical examples of an adaptive management in response to the return of large carnivores.

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.

A comparison of the dynamics of pairs of adjacent moose (Alces alces) populations, one lightly harvested by controlled hunts and the other heavily harvested by open hunts, was conducted in 3 areas in southwestern Quebec. Computer simulations were used to evaluate effects of different hunting regimes on populations, and to generate management predictions. The model assumed that predation was the major mortality factor in the absence of hunting, and that natural mortality was density dependent. The model gave acceptable predictions. Results revealed that density was inversely related to hunting pressure. Field data suggested that unbalanced sex ratios diminished productivity; when this relationship was included in simulations, it appeared to predict observed differences. To optimize moose harvest in southwestern Quebec, we recommend retaining at least 40% males among adults, increasing calf cropping, and limiting hunting to the level of maximum sustained yield. J. WILDL. MANAGE. 45(3):598-611 Computer models for simulating responses of wildlife populations to harvest schemes are being used with increasing frequency. Workers seldom lack ingenuity for simulating population mechanisms; a major limitation, however, arises in applying existing data to calibrate and test these mechanisms. Without a basic understanding of population responses, quantitative predictions having the accuracy required by managers cannot be made. The study reported here, involving 2 levels of hunting pressure on moose, appears to demonstrate that simulated relationships based on data can provide a reasonable basis for management predictions. Moose management in North America has been strongly influenced by the concepts of net and gross productivity (Pimlott 1959), but these do not account for variations in density or nonhunting mortality as functions of hunting impact (Caughley 1974). Caughley (1976) preented a model for ungulates that was based on optimum yield, as was the more general model of Clark (1976); we have applied this approach to moose. In Quebec, the number of moose hunters increased from 43,000 to 98,000 between 1964 and 1977, while the legal take remained relatively stable at 8,000 moose, resulting in declining hunter success (Bouchard and Gauthier 1978). It was essential to determine whether the decline was due to a declining moose population, an inability of new hunters to find suitable hunting sites, or increasing wariness of moose. We designed a study to document effects of different hunting pressures on moose populations, and constructed models to incorporate field data for predicting an optimum-yield management strategy. We thank D. St.-Hilaire, who was in charge of the aerial surveys for most of the project, and J. Boivin, L. Breton, G. Michaud, D. Pare, R. Pariseau, and M. Poulin, who participated in the surveys. We appreciate the skill of pilots B. BenI Present address: Department of Zoology, Clemson University, Clemson, SC 29631. 598 J. Wildl. Manage. 45(3):1981 This content downloaded from 157.55.39.163 on Wed, 21 Sep 2016 05:39:53 UTC All use subject to http://about.jstor.org/terms MOOSE HARVEST IN QUEBEC* Crete et al. 599 bridge, P. Langlois, J.-G. Lejeune, F. Lemieux, Z. Lemieux, C. Richard, and B. Samson and mechanics P. Saint-George and T. To. The computer center of the University of Minnesota provided computer time for simulations.

In many areas of the boreal forests and temperate mountains of Canada, resource extraction activities have created forage conditions that are favourable to the growth of moose (Alces alces) populations. In turn, these increased moose populations buoy the abundance of wolves (Canis lupus), which then have negative impacts on caribou (Rangifer tarandus) populations. Consequently, caribou have been declining where increased resource extraction, moose, and wolves occur. To abate unsustainable predation pressure on caribou by wolves, the moose hunting quota was expanded for 17 years to reduce and then stabilize the moose population in the Revelstoke Valley, British Columbia, Canada. However, a reduction in forestry activity paired with habitat protections slowed the early seral conditions that favour moose. Consequently, both hunter‐caused mortality and habitat loss may have been contributing to observed moose declines that occurred during this period. Within this changing regulatory and biophysical landscape, we sought to address two research objectives. First, we evaluated how increasing the moose hunting quota influenced the total yield of harvested animals. We expected that density‐dependent responses by the moose population would bolster the number of harvestable animals on the landscape. Second, we tested how different forest harvest scenarios might influence moose habitat, wolf densities, and thus caribou population growth rates into future decades. We used data from moose GPS collars (39 individuals), eight aerial population surveys, hunter harvest statistics, estimates of carrying capacity thresholds, and forest harvest records. The latter data series spanned 1961–2020 and informed the resource selection function and calculations for our first research objective as well as the predictive modelling for our second research objective. Between 2003 and 2020, we found that the habitat amounts for moose declined by 44.8%. There were 42% more moose harvested under increased moose hunting quotas than were projected to be harvested under a simulated status quo quota. As the moose population declined and stabilized, we observed higher recruitment rates (e.g. calf:cow ratios) that further contributed to the number of harvested moose. Our simulations indicated that the only forest harvesting scenario where moose carrying capacity would be low enough to stabilize caribou population growth rates by 2040 was to cease forest harvesting entirely in 2020. Practical implication: an increased observed moose harvest quota mitigated the negative effects of forestry on caribou, aided in caribou recovery, and struck a balance that also provided food security and recreational opportunities for moose harvesters.

There are ongoing debates among different stakeholders about which forest and ungulate management strategies will sustain high levels of timber and animal harvest and maintain important ecosystem functions under climate change. Ungulate-forest interactions are complex, including periods where forest regeneration is sensitive to browsing pressure, making it difficult to predict the consequences of a given strategy over time. To aid decision-making, we simulated the impacts of moose browsing on forest succession under 18 different combinations of moose (Alces alces) harvest rate levels and forest management scenarios in a boreal forest landscape in southern Sweden given projected changes in forest growth due to climate change. We found that the current management practices are important for sustaining a moose-forest system. Increasing moose harvest rates led to slightly smaller moose populations, larger estimates of landscape carrying capacity, and less biomass removal of Scots pine (Pinus sylvestris), a commercially valuable species. However, minor changes in the moose harvest were hardly affecting timber production. Increasing the timber harvest rotation time led to the highest estimates of Scots pine biomass, while thinning younger cohorts lead to the highest estimates of Norway spruce (Picea abies) biomass. These changes came without much effect to moose population dynamics. However, the increased broadleaf production scenario had a very large positive effect on total aboveground live biomass of deciduous species and on landscape carrying capacity and moose density. This scenario subsequently resulted in the greatest estimates of biomass removal of Scots pine, highlighting the tradeoffs associated with increased moose production.

• Understanding the population dynamics of a high-density ungulate is challenging. • We built an age/sex-structured population model for a moose population. • Our model successfully reproduced changes in abundance and demographic drivers. • We offered a way to build and calibrate complex population models with scarce data. • We identified mechanisms at play in a protected area without hunting and wolves. High-density populations can threaten the ecological integrity of ecosystems through cascading effects. In such cases, management practices must be guided by sufficient knowledge of the biological mechanisms at play. Simulation models are powerful tools for acquiring such knowledge. The moose ( Alces alces americana ) is a species that recently became overabundant in some areas of eastern North America, sometimes requiring specific management measures. While numerous models exist for moose population dynamics, few are adapted to high density populations like the one in Forillon National Park (Quebec, Canada), a protected area in which the moose's apical predator (grey wolf Canis lupus ) is absent. We developed a sex- and age-structured population model respecting these conditions that we parameterized using pattern-oriented modelling to help explain the changes in moose density observed over nearly 4 decades. The most plausible sequence of vital rates identified exhibited negative density dependence in survival, reproduction and dispersal. Predation by alternative predators, black bears ( Ursus americanus ) and coyotes ( Canis latrans ), caused substantial mortality of calves each year. Unlike elsewhere in northeastern North America, winter tick only had a slight effect on calf survival. Variations in the population’s sex ratio were mainly explained by sex-biased dispersal. Our study provides new insights concerning the dynamics of high-density ungulate populations in the absence of their apical predator, and our modelling approach helped reveal new methodological challenges and opportunities. We also present a comprehensive process to build and parameterize a complex population model using scarce data.

Given recent actions to increase sustained yield of moose (Alces alces) in Alaska, USA, we examined factors affecting yield and moose demographics and discussed related management. Prior studies concluded that yield and density of moose remain low in much of Interior Alaska and Yukon, Canada, despite high moose reproductive rates, because of predation from lightly harvested grizzly (Ursus arctos) and black bear (U. americanus) and wolf (Canis lupus) populations. Our study area, Game Management Unit (GMU) 20A, was also in Interior Alaska, but we describe elevated yield and density of moose. Prior to our study, a wolf control program (1976–1982) helped reverse a decline in the moose population. Subsequent to 1975, moose numbers continued a 28‐year, 7‐fold increase through the initial 8 years of our study (λB1 = 1.05 during 1996–2004, peak density = 1,299 moose/1,000 km2). During these initial 8 hunting seasons, reported harvest was composed primarily of males ( = 88%). Total harvest averaged 5% of the prehunt population and 57 moose/1,000 km2, the highest sustained harvest‐density recorded in Interior Alaska for similar‐sized areas. In contrast, sustained total harvests of <10 moose/1,000 km2 existed among low‐density, predator‐limited moose populations in Interior Alaska (≤417 moose/1,000 km2). During the final 3 years of our study (2004–2006), moose numbers declined (λB2 = 0.96) as intended using liberal harvests of female and male moose ( = 47%) that averaged 7% of the prehunt population and 97 moose/1,000 km2. We intentionally reduced high densities in the central half of GMU 20A (up to 1,741 moose/1,000 km2 in Nov) because moose were reproducing at the lowest rate measured among wild, noninsular North American populations. Calf survival was uniquely high in GMU 20A compared with 7 similar radiocollaring studies in Alaska and Yukon. Low predation was the proximate factor that allowed moose in GMU 20A to increase in density and sustain elevated yields. Bears killed only 9% of the modeled postcalving moose population annually in GMU 20A during 1996–2004, in contrast to 18–27% in 3 studies of low‐density moose populations. Thus, outside GMU 20A, higher bear predation rates can create challenges for those desiring rapid increases in sustained yield of moose. Wolves killed 8–15% of the 4 postcalving moose populations annually (10% in GMU 20A), hunters killed 2–6%, and other factors killed 1–6%. Annually during the increase phase in GMU 20A, calf moose constituted 75% of the predator‐killed moose and predators killed 4 times more moose than hunters killed. Wolf predation on calves remained largely additive at the high moose densities studied in GMU 20A. Sustainable harvest‐densities of moose can be increased several‐fold in most areas of Interior Alaska where moose density and moose: predator ratios are lower than in GMU 20A and nutritional status is higher. Steps include 1) reducing predation sufficient to allow the moose population to grow, and 2) initiating harvest of female moose to halt population growth and maximize harvest after density‐dependent moose nutritional indices reach or approach the thresholds we previously published.

Moose (Alces alces) management strategies in Lithuania, East Europe, were analyzed. The study was intended to show the (un)sustainability of the current management approach in relation to changes in hunting rules, hunting organization and development of the responsible administrative bodies. Moose population and bag dynamics were analyzed using I index in connected scatterplots and compound annual growth rates (CAGR). In 1962–2020, the CAGR of the moose population was 3.84%, resulting in a population size increase of nearly 10 times. The seesaw principle in moose management was confirmed, showing three periods of population decrease (1973–1977, 1989–1995, 2000–2005), and two periods of hunting bag decrease (1976–1978 and 1990–1993). All decline phases were related to legal and administrative issues in the country. Since 2006, population growth has not been controlled. Lithuania has no long-term strategy of the moose population management at any administrative level. The current management approach is not sustainable, as it has not ensured long-term stability of the moose population. The current continuous growth of population, followed by only a moderate increase in the hunting bag, is related to the possibility for owners to adopt long-term planning of the hunting plot units.

Abstract: Age ratios (e.g., calf:cow for elk and fawn:doe for deer) are used regularly to monitor ungulate populations. However, it remains unclear what inferences are appropriate from this index because multiple vital rate changes can influence the observed ratio. We used modeling based on elk ( Cervus elaphus ) life‐history to evaluate both how age ratios are influenced by stage‐specific fecundity and survival and how well age ratios track population dynamics. Although all vital rates have the potential to influence calf:adult female ratios (i.e., calf:cow ratios), calf survival explained the vast majority of variation in calf:adult female ratios due to its temporal variation compared to other vital rates. Calf:adult female ratios were positively correlated with population growth rate (Λ) and often successfully indicated population trajectories. However, calf:adult female ratios performed poorly at detecting imposed declines in calf survival, suggesting that only the most severe declines would be rapidly detected. Our analyses clarify that managers can use accurate, unbiased age ratios to monitor arguably the most important components contributing to sustainable ungulate populations, survival rate of young and Λ. However, age ratios are not useful for detecting gradual declines in survival of young or making inferences about fecundity or adult survival in ungulate populations. Therefore, age ratios coupled with independent estimates of population growth or population size are necessary to monitor ungulate population demography and dynamics closely through time.

Abundant ungulate populations are considered ecosystem drivers that affect forestry and agriculture. Their management is ecologically and economically based on game density regulations, considering the balance between density and carrying capacity of the territory, population status and dynamics, as well as hunting needs.
 Although the population status and dynamics are not new questions, it is still unclear how to manage populations properly depending on the hunting intensity. We aimed to analyse and compare the ungulate population status like moose, red deer, roe deer and wild boar in the Punia pine forest, where the commercial hunting is conducted, and in the hunting grounds managed by hunter clubs in Prienai forest. We performed the study during four hunting seasons of 2009-2010, 2010-2011, 2011-2012, and 2012-2013. The data on harvesting and abundance were obtained from the field works and using the official statistics of the Ministry of Environment.
 The moose local populations are not abundant, or animals occur occasionally, and their density does not reach the minimum permissible rate. At the same time within the study area, moose hunting is remained to be insufficient. The red deer population is rather stable in the hunting grounds used by hunter clubs while hardly reaches the minimum density rate. On the commercial hunting area, the population density 2-3 times exceeds the permissible density rate. The red deer population should be harvested more intensively. The wild boar is used intensively in the grounds of hunter clubs, while animal density is close to the permissible rate. Unfortunately, on the areas of commercial hunting, wild boar is not actively managed that is why their density exceeds permissible rate even four times.
 The main harvested species are red deer and wild boar in both hunting grounds. Their abundant populations stay close to permissible density rate. However, gamekeepers keep the larger animal numbers on the areas of commercial hunting. As the main aim is the trophy hunting, the stags and boars are most used when compared to females and young. Therefore, on the areas of commercial hunting, use of wild boar and red deer is unreasonable and their density exceeds permissible rate several times. Keywords: commercial hunting, hunting clubs, population, status, ungulate

Abundant ungulate populations are considered ecosystem drivers that affect forestry and agriculture. Their management is ecologically and economically based on game density regulations, considering the balance between density and carrying capacity of the territory, population status and dynamics, and hunting needs.
 Although the population status and dynamics are not a new question, it is still unclear how to manage populations properly depending on the hunting intensity. We aimed to analyse and compare the ungulate population status like moose, red deer, roe deer and wild boar in the Punia pine forest, where the commercial hunting is conducted, and in the hunting grounds managed by hunter clubs in Prienai forest. We performed the study during four hunting seasons 2009 - 2013. The data on harvesting and abundance were obtained from the field works and using the official statistics of the Ministry of Environment.
 The moose local populations are not abundant, or animals occur occasionally, and their density does not reach the minimum permissible level. The control of moose is non-purposeful and insufficient on both of study territories. The red deer population is rather stable on areas of hunting clubs while hardly reaches the minimum density level. On the commercial hunting area, the population density 2-3 times exceeds the permissible density level. Red deer population should be used more intensively. Wild boar is used intensively on the areas of hunter clubs, and animal density is close to the permissible level. Unfortunately, on the areas of commercial hunting, wild boar is used passively that is why their density exceeds permissible level even four times.
 The main harvested species are red deer and wild boar in both hunting grounds. Their numerous populations stay close to permissible density level. However, gamekeepers keep the larger animal numbers on the areas of commercial hunting. As the main aim is the trophy hunting, the stags and boars are most used when compared to females and young. Therefore, on the areas of commercial hunting, use of wild boar and red deer is unreasonable and their density exceeds permissible level several times.
 Keywords: commercial hunting, hunting clubs, population, status, ungulate

We analyzed harvest data to test hypotheses that nearly 4 decades of effort to reduce abundance of brown bears (Ursus arctos), black bears (U. americanus) and gray wolves (Canis lupus) in an 60,542 km2 area in south-central Alaska (Game Management Unit [GMU] 13) was positively correlated with moose (Alces alces) harvests in some time-lagged fashion. Predator-reduction efforts were progressively more aggressive over decades (both de facto and officially designated predator control) and did not have clear starting points which complicated our post hoc analyses. We documented no positive correlations (p > 0.05) between harvests of brown and black bears and subsequent moose harvests for any time lag. Moose harvest was negatively correlated with the previous years’ wolf harvest, but the relationship was weak (correlation = −0.33, p < 0.05). Consequently, we reject our hypotheses that harvest of predators was positively correlated with moose harvests. We also observed no differences in mean moose harvests during periods of officially designated wolf control (2005–2020) and a previous period (p > 0.50). We recommend that predator reductions designed to improve hunter harvests of moose be conducted within a research framework that will permit improved interpretations of results and the implementation of an adaptive-management approach to achieve management objectives.

BackgroundClimate is an important driver of ungulate life-histories, population dynamics, and migratory behaviors, and can affect the growth, development, fecundity, dispersal, and demographic trends of populations. Changes in temperature and precipitation, and resulting shifts in plant phenology, winter severity, drought and wildfire conditions, invasive species distribution and abundance, predation, and disease have the potential to directly or indirectly affect ungulates. However, ungulate responses to climate variability and change are not uniform and vary by species and geography. Here, we present a systematic map protocol aiming to describe the abundance and distribution of evidence on the effects of climate variability and change on ungulate life-histories, population dynamics, and migration in North America. This map will help to identify knowledge gaps and clusters of evidence, and can be used to inform future research directions and adaptive management strategies.MethodsWe will catalogue evidence on how climate variability and change affect the life-histories, population dynamics, and migration patterns of the fifteen ungulate species native to North America. We will search both academic and grey literature, using academic journal databases and specialist websites. Articles will be screened for inclusion at the title/abstract and full-text levels, and data will be extracted from articles that pass the full-text review. These data will be summarized quantitatively, visually, and with a narrative review to describe the distribution and abundance of evidence on the effects of climate variability and change on ungulates in North America.

There is increasing pressure to manage forests for multiple objectives, including ecosystem services and biodiversity, alongside timber production. However, few forests are currently co-managed for timber and wildlife, despite potential economic and conservation benefits. We present empirical data from a commercial Norway spruce (Picea abies) and Scots pine (Pinus sylvestris) production system in southern Norway in which moose (Alces alces) are an important secondary product. Combining long-term hunting and forestry records, we identified temporal variation in clear-felling over the past five decades, peaking in the 1970s. Herbicide treatment of regenerating stands and a fivefold increase in moose harvest has lead to a reduction in availability of successional forest per moose of >90 % since the 1960s. Field estimates showed that spraying with the herbicide glyphosate reduced forage availability by 60 and 96 % in summer and winter, respectively, 4 years after treatment. It also reduced moose use and habitat selection of young spruce stands compared with unsprayed stands. Together these lines of evidence suggest that forest management led to an increase in moose carrying capacity during the 1970s and a subsequent decline thereafter. This is likely to have contributed to observed reductions in moose population productivity in southern Norway and is counter to sustainable resource management. We therefore call for better integration and long-term planning between forestry and wildlife management to minimise forest damage and the development of large fluctuations in ungulate populations.

mation has been gained in neighboring areas from studies by McDowell and Moy (1942) in the Absaroka Mountains of Montana; and McMillan (1953) in Yellowstone National Park; and Bassett (1951), Rudersdorf (1952) and Harry (1957) in Jackson Hole, Wyoming. None of these studies are completely applicable to, or adequate for, moose management in the Gravelly Mountain area. This study was conducted on a full time basis during the summer and winter and on a part time basis during the fall, between May 26, 1958, and March 19, 1959. The aim of the study was to provide data for management of moose by investigating plant species and range areas that are indicative of carrying capacity of the range, the rate of moose production, and possible conflicts with other forms of land use. Food habits, movements and population structure were emphasized. Moose management in this area must be integrated with other land uses including production of livestock and other big game species. The writer expresses his sincere appreciation to Don C. Quimby for directing the study and giving aid in preparation of the manuscript; Joseph Townsend, Ralph Rouse and other members of the Montana Fish and Game Department for aid in planning the study and in the field; James Peek and David Spaulding for assistance in conducting field work; and Walter Sperry for hospitality and aid. The writer was employed during the study by the Montana Fish and Game Department under Federal Aid Project W-73-R-4.