Status and Trends of Moose Populations and Hunting Opportunity in the Western United States
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STATUS AND TRENDS OF MOOSE POPULATIONS AND HUNTING OPPORTUNITY IN THE WESTERN UNITED STATES M. Steven Nadeau1, Nicholas J. DeCesare2, Douglas G. Brimeyer3, Eric J. Bergman4, Richard B. Harris5, Kent R. Hersey6, Kari K. Huebner7, Patrick E. Matthews8, and Timothy P. Thomas9 1Idaho Department of Fish and Game, 600 S. Walnut, Boise, Idaho 83709, USA; 2Montana Fish, Wildlife and Parks, 3201 Spurgin Road, Missoula, Montana 59804, USA; 3Wyoming Game and Fish Department, Box 67, Jackson, Wyoming 83001, USA; 4Colorado Parks and Wildlife, 317 W. Prospect Avenue, Fort Collins, Colorado 80526, USA; 5Washington Department of Fish and Wildlife, 600 Capital Way North, Olympia, Washington 98504, USA; 6Utah Division of Wildlife Resources, Box 146301, Salt Lake City, Utah 84114, USA; 7Nevada Department of Wildlife, 60 Youth Center Road, Elko, Nevada, 89801; 8Oregon Department of Fish and Wildlife, 65495 Alder Slope Road, Enterprise, Oregon 97828, USA; 9Wyoming Game and Fish Department, Box 6249, Sheridan, Wyoming 82801, USA. ABSTRACT: We review the state of knowledge of moose (Alces alces shirasi) in the western US with respect to the species’ range, population monitoring and management, vegetative associations, licensed hunting opportunity and hunter harvest success, and hypothesized limiting factors. Most moose monitoring programs in this region rely on a mixture of aerial surveys of various formats and hunter harvest statistics. However, given the many challenges of funding and collecting rigorous aerial survey data for small and widespread moose populations, biologists in many western states are currently exploring other potential avenues for future population monitoring. In 2015, a total of 2,263 hunting permits were offered among 6 states, with 1,811 moose harvested and an average success rate per permit-holder of 80%. The spatial distribution of permits across the region shows an uneven gradient of hunting opportunity, with some local concentrations of opportunity appearing consistent across state boundaries. On average, hunting opportunity has decreased across 56% of the western US, remained stable across 17%, and increased across 27% during 2005–2015. Generally, declines in hunting opportunity for moose are evident across large portions (62–89%) of the “stronghold” states where moose have been hunted for the longest period of time (e.g., Idaho, Montana, Utah, and Wyoming). In contrast, increases in opportunity appear more common at peripheries of the range where popula- tions have expanded, including most of Colorado, northeastern Washington, southern Idaho, and east- ern Montana. There are many factors of potential importance to moose in this region, including parasites, predators, climate, forage quality, forage quantity, and humans. State wildlife agencies are currently conducting a variety of research focused on population vital rates, the development of monitoring techniques, forage quality, trace mineral levels, and evaluation of relative impacts among potential limiting factors. ALCES VOL. 53: 99–112 (2017) Key Words: Alces alces shirasi, Colorado, hunter harvest, Idaho, Montana, Nevada, Oregon, population trends, range, Shiras moose, Utah, Washington, Wyoming The occupied range of moose (Alces knowledge of moose in the western US with alces) extends southward into the western respect to range, population monitoring and United States along the Rocky Mountains management, vegetative associations, licen‐ and the Western Cordillera ecoregion (CEC sed hunting opportunity and hunter harvest 1997; Fig. 1). Here, we review the state of success, and hypothesized limiting factors. 99 STATUS OF MOOSE IN WESTERN US – NADEAU ET AL. ALCES VOL. 53, 2017 Fig. 1. Predicted range of moose in western USA circa 2015, based on compilation of state distribution data characterizing known occupancy by resident moose and predicted occupancy through species distribution modeling of Beauvais et al. (2013). In particular, we use hunter opportunity and as a distinct subspecies during the early part harvest data to address spatio-temporal trends of the 20th century (A. a. shirasi; Nelson in regional moose populations. We focus 1914, Peterson 1952), though historical ac- specifically on trends in hunting opportunity counts suggest they were rare throughout the over the past decade (2005–2015) because US Rocky Mountains until the mid-1800s they facilitate a simple assessment of local (Karns 2007). After periods of population trends with comparable data across the expansion and subsequent declines due to 8-state region. overharvest during the late 1800s, it is large- ly believed that moose populations increased RANGE AND HABITAT to new highs during the early to mid-1900s Moose in the Rocky Mountains of the within portions of Idaho, Montana, Utah, USA and southern Canada were described and Wyoming (Brimeyer and Thomas 2004, 100 ALCES VOL. 53, 2017 NADEAU ET AL. – STATUS OF MOOSE IN WESTERN US Toweill and Vecellio 2004, Wolfe et al. populations in Saskatchewan and North Da- 2010, DeCesare et al. 2014). During the kota, would be better described as A. a. shir- latter half of the 20th century, moose also asi or as belonging to the northwestern naturally colonized eastern portions of subspecies A. a. andersoni. Washington and Oregon, and translocations Habitats occupied by moose vary were used to introduce moose to Colorado throughout the western US with respect to and unoccupied portions of Wyoming and gradients in abiotic conditions (e.g., eleva- Idaho, as well as to augment populations in tion, temperature, and precipitation), vege‐ Utah (Kufeld 1994, Olterman et al. 1994, tative associations, and large mammal Base et al. 2006, Wolfe et al. 2010, Matthews predator-prey communities. We summarized 2012). Sightings of moose occurred in moose management units across each state Nevada as early as the late 1980s, but have within which licensed moose hunting is of- increased in recent years, including multiple fered, in terms of mean values of elevation, verified sightings of cows with calves in annual precipitation (cm), and minimum 2016. While numbers in outlying states January and maximum July temperatures (°C) such as Colorado and Washington appear in a GIS using a digital elevation model and to be stable or increasing, declines have 30-year climate averages during 1981–2010 been noted recently in previous stronghold (PRISM Climate Group 2016). Though portions of Idaho, Montana, Utah, and moose management units in Colorado and Wyoming (Harris et al. 2015, Monteith et al. Utah contain the southern-most introduced 2015, DeCesare et al. 2016). and naturally occurring moose populations This geographic area is primarily occu- in the world, the relatively high elevations pied by the Shiras subspecies of moose of occupied habitats in these regions result (A. a. shirasi) which exists throughout the in climates similar to neighboring states Rocky Mountains from Colorado and Utah further north (Table 1). Average elevation northward to southern Alberta and British ranged from 911 m in Washington to 2712 Columbia. Recent range expansion of moose m in Colorado, average annual precipitation into eastern Montana has come with some ranged from 62–84 cm annually, and mini- uncertainty regarding subspecies identity. It mum January and maximum July tempera- is unclear whether animals east of the Rocky tures averaged 14 to 6 °C and 24 to Mountain chain in the eastern portions 27 °C, respectively, among states during of Montana, with neighboring moose 1981–2010 (Table 1). Table 1. Descriptive statistics characterizing means per moose management unit in elevation, precipitation, minimum January temperature, maximum July temperature, summarized across units within each western state where licensed moose harvest is allowed, 1981–2010. Minimum January Maximum July Elevation (m) Precipitation (cm) Temperature (°C) Temperature (°C) State Mean Range Mean Range Mean Range Mean Range Colorado 2712 (2075–3304) 63 (35–111) 13 (16 to 9) 24 (20–30) Idaho 1656 (804–2353) 84 (24–150) 9(14 to 4) 26 (23–31) Montana 1849 (814–3071) 71 (32–119) 11 (15 to 6) 24 (18–29) Utah 2226 (1896–2715) 64 (48–83) 11 (14 to 8) 26 (23–28) Washington 911 (723–1169) 66 (46–101) 6(8to5) 27 (24–28) Wyoming 2387 (1945–2999) 62 (20–123) 14 (18 to 11) 24 (19–29) 101 STATUS OF MOOSE IN WESTERN US – NADEAU ET AL. ALCES VOL. 53, 2017 Studies of vegetation associations inhab- red-osier dogwood, serviceberry, and menzie- ited by moose in the western US include a sia (Menziesia ferruginea) (Matchett 1985). preponderance of evidence for selection of In north-central Idaho moose select conifer willow (Salix spp.) communities and plants forest including old growth and mature for space use and food habits, particularly mixed-age stands of grand fir (Abies during winter (McMillan 1953, Knowlton grandis) and subalpine fir (Pierce and Peek 1960, Dorn 1970, Wilson 1971, Pierce and 1984) where they forage primarily on Pacific Peek 1984, Van Dyke et al. 1995, Kufeld yew (Taxus brevifolia), Sitka alder (Alnus and Bowden 1996, Dungan and Wright viridis), and menziesia (Pierce 1984). Moose 2005, Baigas et al. 2010, Vartanian 2011, in Washington occupy habitats typically Burkholder et al. 2017). The importance of characterized by dense conifer forest, includ- willow has been documented primarily in rela- ing mixed stands of western red cedar (Thuja tively colder and drier portions of the range plicata) and western hemlock (Tsuga hetero- (e.g., in portions of Colorado, Utah, Wyoming, phylla) and consume diets of willow, Ceano- southeast