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.
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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,
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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( 8to 5) 27 (24–28) Wyoming 2387 (1945–2999) 62 (20–123) 14 ( 18 to 11) 24 (19–29)
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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 Idaho, and southwest Montana), thus spp., and other shrubs and forbs during though exceptions to the importance of wil- summer, and conifers such as cedar and hem- low exist in local populations in these areas. lock during winter (J. Goerz, University of For example, in certain portions of western Montana, pers. comm.). Colorado and Utah, moose have also colo- nized upland shrub communities including POPULATION MONITORING oakbrush (Quercus spp.), serviceberry (Ame- Moose are widespread across broad lanchier spp.), and mountain mahogany (Cer- regions of the western USA but densities are cocarpus spp). In portions of southwest low relative to populations further north in Montana moose occupy forested stands of Canada, Alaska, and the northeastern USA. aspen (Populus spp.) and Douglas-fir (Pseudo‐ Recent population estimates (2014) in our tsuga menziesii) and feed on a range of other study area jurisdictions were 2,400 in Color- shrubs and saplings, including but not limited ado, 10,000 in Idaho, 4,000 in Montana, 20 to serviceberry, huckleberry (Vaccinium in Nevada, 70 in Oregon, 2,625 in Utah, spp.), red-osier dogwood (Cornus sericea), 3,200 in Washington, and 4,650 in Wyoming and subalpine fir (Abies lasiocarpa) (Stevens (Timmermann and Rodgers 2017; Nevada es- 1970). In southeast Idaho moose forage pri- timate unpublished). These abundance esti- marily on bitterbrush (Purshia tridentata), mates are not necessarily comparable as willow, serviceberry, chokecherry (Prunus they were each derived with different virginiana), and aspen (Populus tremuloides) methods and generally, with uncertainty. (Ritchie 1978). Resources to monitor moose are relatively West of the continental divide where oc- sparse when compared with those devoted cupied portions of the range include more to more abundant elk (Cervus canadensis), maritime-influenced climates, moose associ- deer (Odocoileus spp.), and pronghorn ate with various forest communities where (Antilocapra americana) populations. Thus, willow-dominated lowlands are less preva- population monitoring of moose in many lent. In wetter climates such as in northwest areas is met with challenges of limited data Montana, moose show strong associations and low statistical power (Harris et al. with conifer forests, including but not limited 2015, DeCesare et al. 2016). to regenerating vegetation following timber To date, most monitoring programs rely harvest (Matchett 1985, Langley 1993). For- primarily on a mixture of aerial surveys of age species in these areas have included various formats and hunter harvest statistics.
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Colorado Parks and Wildlife biologists use exploration of forward-looking infrared spreadsheet population models that are based (FLIR) technology in comparison to aerial on estimates of survival and recruitment surveys in northern Idaho (sensu Storm largely obtained opportunistically during et al. 2011), patch occupancy modeling of surveys for other species (White and Lubow hunter sightings data in Montana (sensu 2002). In Idaho biologists have similarly col- Rich et al. 2013), and development of a lected survey data incidental to elk surveys smartphone app for collecting public sight- for several decades, but have not developed ings of moose in Washington (sensu Teacher a statewide trend index or population estima- et al. 2013). tion technique. Montana Fish, Wildlife and Parks biologists conduct minimum count HUNTING OPPORTUNITY, SUCCESS aerial surveys in a subset of moose hunting RATES, AND TRENDS units, and rely primarily on hunter harvest Licensed hunting of moose began in the statistics elsewhere for assessment of trends late 19th century in Idaho, Montana, and Wyo- (DeCesare et al. 2016). Moose have more re- ming, but these seasons were subsequently cently colonized portions of Oregon and closed in 1897–1899 due to concerns of over- Nevada where state agency biologists pri- harvest. Hunting seasons were (re)instated marily monitor them via public sighting within 6 Rocky Mountain states in the reports and opportunistic ground and aerial following chronological order: 1912 survey detections. The Utah Division of (Wyoming), 1945 (Montana), 1946 (Idaho), Wildlife Resources conducts aerial surveys 1958 (Utah), 1977 (Washington), and 1985 specifically targeting moose every 3 years (Colorado). The number of harvested moose within hunted units, in addition to collecting has declined in the past decade in 4 of 6 incidental observations during elk surveys. western states (Idaho, Montana, Utah, and Washington Department of Fish and Wildlife Wyoming) that allow hunting (Fig. 2). Con- biologists conduct annual helicopter surveys versely, harvest numbers have continued to construct an annual index that is useful to to increase in Colorado and Washington track general trends and recruitment rates through added opportunities in traditional within most of the core moose range in and newly opened hunting units (Fig. 2). To northeastern Washington; however, despite date, moose hunting has not been initiated incorporating covariates affecting detection in Nevada or Oregon, though a bull moose probabilities, these data remain imprecise was incidentally harvested in Nevada in the (Harris et al. 2015). Wyoming Fish and 1950s. In 2015, a total of 2,263 hunting per- Game biologists conduct annual moose- mits were offered by the 6 states, resulting in specific aerial surveys in the more abundant 1,811 harvested moose and an average harvest herd units and use these data in a modified success rate of 80% (Table 2). Since 1990, spreadsheet simulation model (similar to the highest single-year state harvest was that used in Colorado); incidental observa- 1,215 moose in Wyoming in 2001 (Fig. 2). tions also occur in surveys of other big In contrast, the highest harvest in 2015 was game populations (Monteith et al. 2015). 666 moose in Idaho. Proportionate harvest Given the many challenges of funding of antlerless (cows and calves) moose ran‐ and collecting rigorous aerial survey data ged from 0 to 17% in states with generally for small and widespread moose popula- declining opportunity, compared to 35% tions, many western state agencies are cur- and 49% antlerless harvest in Washington rently exploring other potential avenues for and Colorado, respectively, where hunter op- future population monitoring. These include portunity continued to increase through 2015
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Fig. 2. Moose harvest trends in the western USA from 2000 to 2015.
(Table 2). For the states focal to this sum- Rockies and Columbia Mountains region of mary, moose hunting is managed through lot- Washington, northern Idaho, and northwest tery permit systems that produce high harvest Montana; 2) the Middle Rockies region of success rates averaging 75-95% annually southwest Montana, southeast Idaho, and (Table 2). There is a tendency for somewhat western Wyoming; 3) the Southern Rockies lower rates of success on antlerless permits region of north-central Colorado; and 4) versus antlered-only or either-sex permits the Wasatch and Uinta ranges in north-central (Table 2). Utah (Fig. 3a). Spatial distribution of permits across the We summarized trends in hunter op‐ region shows an uneven gradient of hunting portunity across the 11-year period of opportunity (i.e., number of moose hunting 2005–2015 for all hunting management units permits) across the states (Fig. 3a). Although or districts (hereafter “units”) across the en- moose occupy areas beyond hunting units tire region. Given differences in popula‐ alone, multi-state concentrations of moose tion monitoring techniques across states, hunting opportunity appear in approximately hunter opportunity data provide the most 4 portions of the range: 1) the Northern standardized means of assessing and
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Table 2. Numbers of moose hunting permits issued and moose harvested, percentage of antlerless moose (cows and calves) in the harvest, and success rates of hunters holding antlered (including either-sex) and antlerless permitsamong states in the western USA during the 2015 hunting season. % Antlerless in % Hunter success, % Hunter success, State Permits Harvest harvest Antlered1,2 Antlerless1 Colorado 313 233 48.9% 85.6% 65.5% Idaho 873 666 17.4% 77.6% 73.9% Montana 362 268 13.6% 73.8% 76.1% Nevada 0 0 –– – Oregon 0 0 –– – Utah 143 137 0% 95.8% – Washington 168 142 35.2% 91.6% 72.1% Wyoming 411 365 16.4% 90.8% 80.0%
1Hunter success measured as harvested moose per permit allocated, not accounting for number of permit-holders that actually hunted. 2Hunter success for antlered moose includes the combined success rates of both antlered-only and either-sex permit-holders, given either-sex permit-holders harvest predominately antlered moose (e.g., 94% of hunters in Washington).
Fig. 3. Regional patterns in a) the availability of moose hunting permits in 2015 and b) changes in permit availability per hunting unit between 2005 and 2015 across the western USA. Note some hunting units were merged together for one or both years to facilitate comparisons of equal areas among years.
105 STATUS OF MOOSE IN WESTERN US – NADEAU ET AL. ALCES VOL. 53, 2017 comparing trends among states. In the ab- abundant enough to support hunting for the sence of consistent sampling-based surveys, longest period of time (i.e., Idaho, Montana, we treat hunting opportunity as an index to Utah, and Wyoming). We note that this trend population status, assuming that changes in was evident prior to 2005 in Wyoming opportunity within units reflect relative where loss of 527 permits occurred between trends in abundance. In order to allow com- 2000 and 2005. parison of equal areas across years, we In contrast, increases in hunting oppor- merged some units together for either the tunity were more common at peripheries of 2005 or 2015 seasons to accommodate the range where populations expanded, in- changes in unit boundaries or regulations cluding most of Colorado, central Utah, among years. southern Idaho, northeastern Washington, In total, we assessed changes in oppor- and eastern Montana. The increases in east- tunity across 273 units within 6 states from ern Montana mirror those in similar prairie 2005 to 2015 (Table 3). Overall, moose hunt- habitats of neighboring jurisdictions in ing opportunity declined by 23% across the southeast Alberta, western North Dakota, entire region, from 2,970 permits in 2005 and southwest Saskatchewan where moose to 2,279 permits in 2015, but trends varied populations have generally increased in re- among local units. On average, hunting op- cent years (Laforge et al. 2016). Increases portunity decreased across 56% of the units, in hunting opportunity in Colorado were par- remained stable across 17%, and increased tially in response to continued introductions across 27% (Table 3). Visual display of of moose into new areas including the Grand trends per hunting unit shows a diversity of Mesa National Forest (2005–2007) and the dynamics, from areas that were closed to White River National Forest (2009–2010). hunting to areas newly opened to hunting Not coincidentally, the few units in Colorado (Fig. 3b). Generally, declines in hunting op- where hunting opportunity declined were portunity for moose are evident across also those serving as source populations for much of their occupied area in the western translocations. The Wasatch Unit has the US, including large portions (62–89%) of most recently established population and “stronghold” states where moose have been offers the most hunting opportunity in Utah.
Table 3. Trends in moose hunting opportunity (i.e., number of moose hunting permits) from 2005–2015, summarized per hunting unit across 6 western states. Trends in moose permits per unit during 2005–2015 2 State Nunits Median unit area (km ) Decreased Stable Increased Colorado 42 1127 5% 5% 90% Idaho 92 834 62% 21% 17% Montana 85 1490 67% 20% 13% Utah 9 2431 89% 0% 11% Washington 7 2468 14% 0% 86% Wyoming 38 1810 66% 24% 11% Area-weighted 56% 17% 27% average1
1Area-weighted averages were estimated by weighting states according to the product of the number of hunting units and median area per unit for each state.
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POTENTIAL LIMITING FACTORS been linked to expansion of moose in Alaska There are many factors of potential im- through habitat change (Tape et al. portance to moose population dynamics 2016), and some southern populations re- across the western USA, and data concern- main stable or increasing (Murray et al. ing their presence, prevalence, or effects on 2012), we share the speculative concern of moose vary across populations. Multiple Lenarz et al. (2010) over the long-term via- parasites and diseases including the arterial bility of moose populations along their worm (Elaeophora schneideri), winter tick southern range edge. Current climate projec- (Dermacentor albipictus), giant liver fluke tions suggest that temperature will increase (Fascioloides magna), chronic wasting dis- during all seasons in the Rocky Mountain ease, hydatid worm (Echinococcus granulo- region, and precipitation may increase in sus), and other tapeworms (Taenia spp.) northern portions (Rocca et al. 2014). Warmer have been documented in the region from temperatures may possibly induce heat stress- examination of hunter-killed, live-captured, related impacts in free-ranging moose as or opportunistically collected specimens of measured in captive moose (Renecker and moose or other ungulate species (e.g., Worley Hudson 1986, McCann et al. 2013). Indirect et al. 1972, Samuel et al. 1991, Dunkel et al. effects of warmer climate may also impact 1996, Pessier et al. 1998, Henningsen et al. moose, as mediated by changes in parasite- 2012, LeVan et al. 2013). Large predator host communities or plant communities communities may have up to 4 species - black and/or phenology (Rempel 2011, Monteith bears (Ursus americanus), grizzly bears et al. 2015). (Ursus arctos), cougars (Puma concolor), Interestingly, we find that the southern- and wolves (Canis lupus) - that vary in dens- most global populations of moose found in ity and potential to affect moose population Colorado appear to be the most stable in dynamics within individual states (sensu the western USA as evidenced by population Griffin et al. 2011, Brodie et al. 2013). growth and increased hunting opportunity Presence of wolves, grizzly bears, and (Fig. 3b). However, latitude alone may not chronic wasting disease vary most across sufficiently characterize variation in climate states (Table 4). across this region, as portions of Colorado The potential importance of nutritional are similar in temperature regime to more limitations has been documented in certain northern areas (Table 1), and increasing western moose populations. For example, populations such as those in Colorado or Ruprecht et al. (2016) reported low fat levels Washington are also relatively young and ex- in moose in Utah relative to northern popula- panding into unoccupied habitats. It is pos‐ tions, as well as lower pregnancy and twin- sible that the dynamics of these populations ning rates than in other North American are still in accordance with the earlier stages populations at lower latitudes. Both are sug- of the eruptive cycle of introduced ungulate gestive of nutritional limitation, although populations identified by Caughley (1970). nutritional condition is affected by factors The complexities of time since establish- other than forage including disease, para- ment and density dependence may confound sites, and combined influences related to comparisons among populations with respect climate change. to climate- or habitat-related conditions; how- Climate change may have particularly ever, direct physiological impacts of heat pronounced effects on populations at the stress should manifest regardless. In short, periphery of a species’ range (Hampe and multiple factors other than latitude alone are Petit 2005). Although climate change has influential on the short- and long-term
107 TTSO OS NWSENUS WESTERN IN MOOSE OF STATUS
Table 4. Documented presence, absence or uncertainty regarding parasites and predators of potential importance to moose dynamics across states of the western USA, circa 2015.
Parasites and disease Predators Elaeophora Dermacentor Echinococcus Chronic Fascioloides Parelaphostrongylus State schneideri albipictus granulosus wasting disease magna tenuis Black bear Grizzly bear Cougar Wolf –
Colorado ++ ++ + + ––++ – 2017 53, VOL. ALCES ++ – AL. ET NADEAU Idaho ++ ++ ++ – ++ – ++ + ++ ++
108 Montana ++ ++ ++ – ++ – ++ ++ ++ ++ Nevada + + ––– – + – ++ – Oregon ++ ++ ––++ ++ Utah ++ ++ + + – ++ ++ Washington + ++ ––++ + ++ ++ Wyoming ++ ++ + + – ++ ++ ++ ++
++ = Documented as commonly present in moose or areas occupied by moose. + = Documented as present among ungulate populations, but rare in moose or areas occupied by moose. – = Not documented and presence seen as unlikely in moose or areas occupied by moose. [blank] = Not documented and presence unknown in moose or areas occupied by moose. ALCES VOL. 53, 2017 NADEAU ET AL. – STATUS OF MOOSE IN WESTERN US dynamics of moose populations in North ACKNOWLEDGEMENTS America. We thank M. Atamian, D. Base, J. State-regulated hunter harvest, tribal Goerz, H. Ferguson, S. Hansen, A. Holland, harvest, and illegal harvest of moose are M. Lloyd, J. Newby, J. Oyster, K. Podruzny, expected to play some role in regulating A. Prince, T. Smucker, and P. Wolff for their moose populations across the western contributions to the collection and compil- USA. Hunting permits are allocated consid- ation of moose monitoring data that went ering local objectives throughout this region, into this manuscript. J. Gude, J. Maskey, and range from conservative permit numbers M. Mitchell, and J. Smith contributed to to minimize impacts of hunting, to liberal meetings and discussions that led to the numbers of permits to reduce populations development of this manuscript. We thank and their impacts on humans and the envir- A. Apa, K. Logan, the associate editor and onment (Table 2; DeCesare et al. 2014). 1 anonymous reviewer for helpful edits and Tribal harvest of moose is permitted in feedback. most western states through treaty rights, but the availability of specific data varies REFERENCES by jurisdiction. In Montana, tribal harvest BAIGAS, P., R. A. OLSON,R.M.NIELSON, was estimated to increase the total annual S. N. MILLER, and F. G. LINDZEY. 2010. harvest by 7–16% during 1986–2012. Illegal Modeling seasonal distribution and spa- harvest may also have measurable impact on tial range capacity approximations of moose; for example, the result of multiple moose in southeastern Wyoming. Alces studies in Idaho suggested that 31–50% of 46: 89–112. known mortality was associated with illegal BASE,D.L.,S.ZENDER,andD.MARTORELLO. harvest (Ritchie 1978, Pierce et al. 1985, 2006. History, status, and hunter harvest Toweill and Vecellio 2004). of moose in Washington state. Alces 42: 111–114. FUTURE RESEARCH BEAUVAIS, G., M. ANDERSEN,D.KEINATH, State wildlife agencies are conducting J. AYCRIGG,andJ.LONNEKER. 2013. Pre- research in conjunction with universities and dicted vertebrate species habitat distribu- – coordinating research among states to leverage tions and species richness. Pages 58 110 resources across jurisdictions. Research objec‐ in J. Aycrigg, M. Andersen, G. Beauvais, tives include work focused on population M. Croft, A. Davidson, L. Duarte, J. vital rates (e.g., adult female survival, fecun‐ Kagan, D. Keinath, S. Lennartz, J. Lonneker, T. Miewald, and J. Ohmann, dity, and calf survival), movements and editors. Ecoregional Gap Analysis of spatial ecology, resource selection, nutritional the Northwestern United States: North- ecology and forage monitoring, baseline west Gap Analysis Project. Draft report. disease and mineral monitoring, the devel- University of Idaho, Moscow, Idaho, USA. opment of monitoring techniques, and iden- BRIMEYER, D. G., and T. P. THOMAS. 2004. tification of limiting factors. Research History of moose management in Wyo- objectives regarding limiting factors include ming and recent trends in Jackson Hole. the assessment of relative impacts of preda- Alces 40: 133–144. tion, parasites, climate change (direct effects BRODIE, J., H. JOHNSON,M.MITCHELL,P. of heat stress and indirect effects on moose ZAGER,K.PROFFITT,M.HEBBLEWHITE, foraging behavior and parasite loads), and M. KAUFFMAN,B.JOHNSON,J.BISSON- habitat changes (e.g., decline of early seral ETTE,C.BISHOP,J.GUDE,J.HERBERT, forests) on moose vital rates. K. HERSEY,M.HURLEY,P.M.LUKACS,
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