THE STATUS AND MANAGEMENT OF MOOSE IN NORTH AMERICA – CIRCA 2015 H. R. Timmermann1 and Arthur R. Rodgers2 1RR #2 Nolalu, Ontario, Canada POT 2K0; 2Ontario Ministry of Natural Resources and Forestry, Centre for Northern Ecosystem Research, 103–421 James Street South, Thunder Bay, Onatario, Canada P7E 2V6 ABSTRACT: Both declining and increasing moose (Alces alces) populations have been reported across North America over the last decade. We surveyed all jurisdictions with extant moose popula- tions to determine the extent of these population trends. In 2014–2015, the North American moose population was estimated at ~1,000,000 animals distributed in 30 jurisdictions, which is unchanged since the turn of the century. Populations occurred in 12 Canadian provinces or territories, and in at least 18 states. In the past 5 years, moose density is believed to be increasing in 9, relatively stable in 8, and declining in 11 jurisdictions; estimates of change were unavailable in 2 jurisdictions. In 2014–2015, an estimated 425,537 licensed moose hunters harvested 82,096 moose in 23 jurisdictions. Hunter numbers increased by 39,118, whereas total harvest remained virtually unchanged from a dec- ade earlier. Harvests by Indigenous and subsistence users, although largely unquantified, are believed substantial and important to quantify in certain jurisdictions. A variety of active and passive harvest strategies used to manage moose are discussed. ALCES VOL. 53: 1–22 (2017) Key words: Alces alces, distribution, harvest, hunter numbers, Indigenous hunters, licensed qualifications, moose population status, National Parks, seasons, subsistence Over the last decade there have been sev- eral reports of declining moose (Alces alces) populations across North America (Lenarz et al. 2010, Smith et al. 2011, DeCesare et al. 2014), but there have also been accounts of increasing numbers in other areas (Wattles and DeStefano 2011, Harris et al. 2015, LaForge et al. 2016, Tape et al. 2016). In this paper, we update the status and manage- ment of North American moose circa 2014–2015 from that reported in 2000–2001 (Timmermann 2003) to determine the extent of these population trends across the contin- ent. A comprehensive 9-page questionnaire (located at http://alcesjournal.org/index.php/ alces) similar to that employed previously (Timmermann 1987, Timmermann and Buss 1995, Timmermann 2003), and a literature review were used to update the status, popula- tion estimates, and harvest and non-harvest management strategies used in 23 jurisdic- tions with an annual licensed moose harvest. An additional 7 jurisdictions where hunting is currently prohibited were contacted to deter- mine population status. Tabulated data were returned for final perusal, edits, or corrections solicited. This paper reports on current (year 2014–2015) population status and strategies used to manage hunting harvest and non- harvest management of moose across North America. Affiliations of those providing infor- mation through personal communication (pers. comm.) are provided in Acknowledgements. 1 http://alcesjournal.org/index.php/alces http://alcesjournal.org/index.php/alces HISTORICAL DISTRIBUTION AND CURRENT STATUS The distribution of moose in North America during the 20th century has been described by several authors including Peterson (1955), Telfer (1984), Kelsall (1987), Karns (1998), Franzmann (2000), and Rodgers (2001); 4 subspecies are recog- nized, namely A. a. gigas, andersoni, ameri- cana, and shirasi (Peterson 1955). In the past 40+ years many have detailed expanding distributions of moose in both western and eastern states, provinces, and territories (Kelsall and Telfer 1974, Compton and Oldenberg 1994, Karns 1998, Peek and Morris 1998, Brimeyer and Thomas 2004, Toweill and Vecellio 2004, Base et al. 2006, Thomas 2008, Wolfe et al. 2010, Matthews 2012, Labonte et al. 2013, Wattles and DeStefano 2011, 2013, DeCesare et al. 2014). Periodic winter aerial surveys based on the Gasaway method are used by most agen- cies to estimate moose populations and trends (Gasaway et al. 1986, Peterson and Page 1993, Timmermann 1993, Smits et al. 1994, Lynch and Shumaker 1995, Bisset 1996, Lenarz 1998, Timmermann and Buss 1998, Bisset and McLaren 1999, Bontaities et al. 2000, Ward et al. 2000, Gosse et al. 2002, Heard et al. 2008, Larter 2009, Moen et al. 2011a, Cumberland 2012, Fieberg and Lenarz 2012, DelGiudice 2013, Kantar and Cumberland 2013, Millette et al. 2014, Seaton 2014, Harris et al. 2015). Moose are considered among the more difficult ungulates to survey (Harris et al. 2015) and estimating either abun- dance or population trends from raw counts obtained by aerial survey can be challenging. Most agencies estimate total jurisdictional populations based on the cumulative total of specific management areas sampled every 3 or more years. Such jurisdictional estimates are often considered relatively crude and are primarily used to assess population trends, re- cruitment, and distribution over time. Real changes in population estimates are indicated by changes of ~20% or more between surveys (Gasaway and Dubois 1987). New Hampshire, Maine, and Vermont rely heavily on surveys of moose observations by deer (Odocoileus virgi- nianus) hunters and vehicle collision rates to estimate population trends. New Hampshire and Vermont use these annual deer hunter sur- veys in a related regression formula developed from concurrent infrared aerial surveys in a 3-year New Hampshire study (Bontaites et al. 2000, Millette et al. 2014). Jurisdictions not employing formal methods of population as- sessment base their estimates on professional opinion. Consequently, population estimates are not necessarily comparable across jurisdic- tions or years because of the high variation in methodology and quality of data. As with all survey data, absolute counts are not achievable and the data herein should be treated as provid- ing an indication of trends rather than absolute population estimates; the direction of popula- tion change (decreasing, increasing, or stable) is more important than the magnitude of change since the last jurisdictional survey (Timmermann 2003). Eastern North America Currently, moose (A. a. americana) appear to be still expanding and/or occupying former range in the states of Maine, Massachusetts, New York, and Connecticut (Kilpatrick et al. 2003, Hickey 2008, Labonte et al. 2013, Wattles and DeStefano 2011, 2013, S. Heer- kens, L. Kantar, A. LaBonte, and D. Scarpitti, pers. comm. 2015; Fig. 1, Table 1). Moose in Vermont and New Hampshire have re- occupied all suitable habitat and are currently considered to be in slow decline (Musante et al. 2010, C. Alexander and K. Rines, pers. comm. 2015; Fig. 1, Table 1). Factors believed responsible for lower densities in Vermont include purposeful harvest to reduce specificregionalpopulations(Andreozzi et al. 2014). In New Hampshire, high abundance of winter ticks (Dermacentor albipictus) due to shorter winters and possible increased 2 STATUS OF MOOSE IN NA CIRCA 2015 – TIMMERMANN ALCES VOL. 53, 2017 AND RODGERS F ig . 1 . E st im at es o f 2 0 1 4 -2 0 1 5 p o st -h u n t m o o se p o p u la ti o n s in 3 0 N o rt h A m er ic an ju ri sd ic ti o n s. ALCES VOL. 53, 2017 TIMMERMANN AND RODGERS – STATUS OF MOOSE IN NA CIRCA 2015 3 T ab le 1 . N u m b er s o f sp o rt h u n te rs , h ar v es t, an d p o st -h u n t p o p u la ti o n es ti m at es fo r 2 4 N o rt h A m er ic an ju ri sd ic ti o n s su rv ey ed in 2 0 0 0 –0 1 (2 0 0 1 ) an d 2 0 1 3 –1 4 (2 0 1 4 ). 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Current moose populations in Maine, however, appear to have more than doubled since 2001 (Timmermann 2003, Wattles and DeStefano 2011) and are second to only Alaska in the United States (Lichtenwalner et al. 2014). Populations in the Canadian Provinces of New Brunswick and Quebec have increased, while those on Cape Breton, Nova Scotia are believed rela- tively stable or in slight decline since 2001 (Beazley et al. 2008, Smith et al. 2010, S. Lefort, P. MacDonald, and D. Sabine, pers. comm. 2016; Fig. 1, Table 1). On mainland Nova Scotia, the current population esti- mated at 500 is likely still in decline, and is designated “Endangered” under the Nova Scotia Endangered Species Act (P. MacDonald, pers. comm. 2016). Moose numbers on the is- land of Newfoundland have been decreasing since 2001, whereas moose in the Labrador portion of the Province appear to be increas- ing in recent years (J. Neville, pers. comm. 2015). Overabundant moose populations on the island of Newfoundland, where densities remained higher than elsewhere in North America at the turn of the century, have led to habitat deterioration and localized popula- tion decline (McLaren et al. 2004). Conse- quently, harvest quotas were adjusted to modify population size in an effort to reduce and sustain specific populations (McLaren and Mercer 2005), and more recently to help address moose-human conflicts in select management units (J. Neville, pers. comm. 2016). Western North America Moose populations (Alces a. shirasi) are believed to have doubled in Washington State since 2000–2001 (R. Harris, pers. comm. 2015) and have dispersed into Oregon (P. Matthews, pers. comm. 2015; Fig. 1, Table 1). Density has declined in Idaho and Wyoming, but is stable in Utah (D. Brimeyer, K. Hersey,Ta b le 1 co n ti n u ed T o ta l h u n te rs N o n -r es id en t h u n te rs T o ta l es ti m at ed h ar v es t E st im at ed m o o se p o p u la ti o n A g en cy 2 0 0 1 2 0 1 4 2 0 0 1 2 0 1 4 2 0 0 1 2 0 1 4 2 0 0 1 2 0 1 4 V er m o n t 2 1 5 3 4 2 2 2 3 4 1 5 5 1 7 1 3 ,5 0 0 + 2 ,2 0 0 � N ew H am p sh ir e 5 8 5 1 2 7 7 6 1 9 4 1 9 9 1 5 ,0 0 0 + 3 ,8 0 0 � T O T A L 3 8 6 ,4 1 9 4 2 5 ,5 3 7 1 6 ,1 5 8 1 4 ,1 8 3 8 3 ,2 4 6 8 2 ,0 9 6 9 3 5 ,6 3 5 –1 ,0 5 0 ,6 3 5 1 ,0 8 2 ,0 2 0 –1 ,0 8 9 ,0 2 0 1 Y K In d ig en o u s h ar v es t is n o t in cl u d ed . 2 N W T In d ig en o u s h ar v es t is n o t in cl u d ed . 3 N U es ta b li sh ed 1 9 9 9 , fo rm er ly p ar t o f N W T , p o p u la ti o n 3 2 ,0 0 0 (2 0 11 ), 9 ,9 8 4 ,6 7 0 k m 2 . 4 M B 2 0 1 4 h ar v es t es ti m at e p en d in g su rv ey co m p le ti o n . 5 M N cl o se d m o o se se as o n b eg in n in g 2 0 1 3 . ALCES VOL. 53, 2017 TIMMERMANN AND RODGERS – STATUS OF MOOSE IN NA CIRCA 2015 5 and S. Nadeau, pers. comm. 2015). Wyoming populations declined from an estimated 13,865 in 2001 to 7,700 in 2008, and to 4,650 currently (Timmermann 2003, Brimeyer and Thomas 2004, Smith et al. 2011, D. Brimeyer, pers. comm. 2015). Populations have grown in Colorado and remain relative- ly stable compared to a declining trend in Montana (Tyers 2006, DeCesare et al. 2014, N. DeCesare and A. Holland, pers. comm. 2015). Periodic dispersal into the central United States, primarily from North Dakota and Minnesota, is reported as far south as Kansas and Missouri (Hoffman et al. 2006). Moose (Alces a. andersoni) populations in central British Columbia have declined, but overall, the Provincial population has remained relatively stable since 2000–2001 (Kuzyk and Heard 2014, Kuzyk 2016). Moose (Alces a. gigas) in Alaska have increased and those in the Yukon Territories have remained stable (B. Dale, R. Florkie- wicz, and K. Titus, pers. comm. 2015; Fig. 1, Table 1). Moose on the Arctic coastal plain in Alaska have expanded and contracted their numbers and range twice in the past 25 years (B. Dale and K. Titus, pers. comm. 2015). Recent research has linked range expansion in Arctic Alaska to warming and the asso- ciated increase in shrub habitat (Tape et al. 2016). Moose have also expanded their range in northern Southeast Alaska where first observed in Haines in 1924, and now inhabit the Gustavus Forelands (1966; Glacier Bay National Park). Populations of moose now occur on all the major islands of the central Southeast Panhandle of Alaska (B. Dale and K. Titus, pers. comm. 2015). Central North America Moose have expanded northward in Nunavut and Labrador, and are found as far north as 67˚ 31ʹ′ near Kugluktuk in Nunavut and Richards Island in the Northwest Terri- tories Mackenzie Delta (V. Crichton, pers. comm. 2015). Population estimates for vast portions of the Northwest Territories and Nunavut (formerly part of the NWT) are not currently available (M. Dumond and A. Smith, pers. comm. 2015; Fig. 1, Table 1). A further 6 of 8 jurisdictions in the mid- continent report recent, declining trends in moose populations (Alces a. andersoni/ americana) including the adjacent provinces of Alberta, Saskatchewan, Manitoba, and Ontario, as well as the neighboring states of North Dakota and Minnesota (Lenarz et al. 2010, R. Corrigan, G. DelGiudice, H. Hristienko, G. Lucking, L. McInenly, J. Smith, R. Tether, pers. comm. 2015; Fig. 1, Table 1). LaForge et al. (2016) report increasing moose populations in the farm- lands of southern Saskatchewan. Similarly, Manitoba populations have increased in the last 20 years in the southwest farmlands of the province where access is controlled and few predators exist (H. Hristienko and K. Rebizant, pers. comm.). These expansions appear linked to the reduction of small, pri- vately owned farms being replaced by larger corporate farms, and a corresponding decline in undocumented harvest. However, the ad- jacent jurisdictions of Ontario, Manitoba, and Minnesota give lower overall estimates than in 2001 (Table 1). Minnesota closed their harvest in the northwestern region in 1997 due to dramatic population decline from unknown causes (M. Schrage, pers. comm. 2001, Wünsch- mann et al. 2015). The estimated decline was from 4,264 in 1983 to 1,486 in 1995, to ~900 animals in 2001; essential collapse of this population occurred by the early 2000s. Murray et al. (2006) concluded that the giant liver fluke (Fascioloides magna) was largely responsible for this decline. A concurrent de- cline in adjacent northeastern North Dakota was investigated by Maskey (2011) who sug- gested other factors such as brainworm play a larger role in moose mortalities. Minnesota closed its moose hunting season in 6 STATUS OF MOOSE IN NA CIRCA 2015 – TIMMERMANN ALCES VOL. 53, 2017 AND RODGERS northeastern Minnesota in 2013 after num- bers dropped from ~8,500 in 2006 to 3,500 in 2014 (G. DelGiudice and L. McInenly, pers. comm. 2015). Manitoba’s moose popu- lation is believed to have dropped from a his- torical high of 45,000 several decades ago to 27,000 in 2015 (H. Hristienko, pers. comm. 2015). Disease, over-harvest, and human de- velopment of landscapes are the primary fac- tors thought responsible for the decline (Crichton et al. 2004). Recent surveys in northwestern Ontario indicate a correspond- ing decline in certain moose populations (OMNRF 2015, Table 1). Current populations are increasing in Michigan, largely due to higher density esti- mates on Isle Royale (Vucetich and Peterson 2015). Abundance in 2015 was estimated as 323 in the reintroduced population in the western Upper Peninsula, but low productiv- ity and calf:cow ratios suggest population decline (Dodge et al. 2004, D. Beyer, pers. comm. 2015). Populations in neighboring Wisconsin, where moose regularly move in and out of northern Michigan and Minnesota, are currently estimated at <50 (K. Wallenfang, pers. comm. 2015; Fig. 1). The United States Fish and Wildlife Service (USFWS) is consid- ering a listing under the Endangered Species Act of the northwestern subspecies of moose (Alces a. andersoni) that is purported inhabiting upper Michigan, Isle Royale, Minnesota, North Dakota, and Wisconsin (https://www.fws.gov/midwest/es/soc/pdf/ FRBatch90DayFndngs03June2016PIversion. pdf). In response, Michigan and Wisconsin submitted letters to the USFWS indicating that moose in their jurisdictions originated from eastern moose populations (Alces a. americana) (K. Wallenfang, pers. comm. 2016). To summarize in 30 jurisdictions, current moose density is believed stable in 8, increasing in 9, decreasing in 11, with data unavailable in 2 (Fig.1, Table 1). In 22 jurisdictions (circa 2014–2015) for which population estimates are available, and in which an annual licensed harvest occurred in 2014, the total population estimate is 1,082,020 to 1,089,020 animals which is collectively similar to that reported in 2001 (Table 1). Remarkably little overall change has occurred despite the majority of jurisdic- tions reporting either increasing or decreas- ing populations. Population estimates in 12 Canadian jurisdictions totaled 790,845 in 2014 compared to a range of 734,000 to 849,000 in 11 jurisdictions in 2001 (Table 1, Fig.1, Timmermann 2003). The total population increased in 17 states from 204,150–205,130 in 2001, to 274,768–302,268 in 18 states in 2014 (Table 1, Fig.1, Timmermann 2003). Of the 7 states where hunting is prohibited, 3 report expanding populations (Oregon, Michigan, New York), stable popu- lations exist in Wisconsin, Massachusetts, and Connecticut, whereas Minnesota closed their season in 2013 due to significant popu- lation decline (Moen et al. 2011b; Fig. 1). Factors affecting decreasing densities A host of factors are believed responsible for moose population declines including climate change, illegal harvest, habitat loss or degradation, parasites and disease, disturb- ance, moose-vehicular collisions, predators, and unregulated recreational and Indigenous and subsistence harvests (West 2009). In the 11 of 30 (37%) jurisdictions that indi- cated a declining population trend, the “most important factors” were: Parasites and Disease (8 jurisdictions), Predators (7), Natural Habitat Loss (5), Unregulated Har- vest (3), Warmer Summers/Winters (2), Increased Access and Vehicle Technology (1), Higher Deer Densities (1), Over harvests by Licensed Hunters (1), Increased Hunting Pressure (1), and Variable Factors (1). Minne- sota initiated a $1.2 M moose mortality study in 2013 to help determine factors responsible for the recent dramatic population decline. Preliminary results provide evidence of the ALCES VOL. 53, 2017 TIMMERMANN AND RODGERS – STATUS OF MOOSE IN NA CIRCA 2015 7 https://www.fws.gov/midwest/es/soc/pdf/FRBatch90DayFndngs03June2016PIversion.pdf https://www.fws.gov/midwest/es/soc/pdf/FRBatch90DayFndngs03June2016PIversion.pdf https://www.fws.gov/midwest/es/soc/pdf/FRBatch90DayFndngs03June2016PIversion.pdf importance of parasites and disease and pre- dators as mortality factors (Wünschmann et al. 2015). In Maine and New Hampshire, similar research initiated in 2014 indicates that winter ticks remain a primary influence on calf mortality and adult cow productivity (L. Kantar and K. Rines, pers. comm. 2016). HARVEST MANAGEMENT Economic impact Moose, a symbol of wilderness, are much valued by Indigenous hunters, Metis People, recreational hunters, and a host of non- consumptive users (Timmermann and Rodgers 2005). Licensed recreational hunting promotes substantial benefits to local economies valued in the $100s of millions annually. In the early 1990s, for example, Legg (1995) estimated CAN $134.7 M in Ontario for all hunter-related activities in 1993. More re- cently, Maine estimated the economic impact of 3,095 resident and 310 non-resident hun- ters to represent US $11.9 M and $3.9 M in 2014 (L. Kantar, pers. comm. 2015), and Alaska valued its non-resident hunt at $11M in 2014 (B. Dale, pers. comm. 2015). Similar- ly, Quebec estimated 176,710 residents and 2,707 non-residents generated CAN $204 M and $8.0 M in 2014 (S. Lefort, pers. comm. 2015). Harvest control objectives Three territories and 9 provinces in Canada, and 11 states in the United States ad- ministered a moose hunt in 2014 (Table 1). Collectively, 425,537 licensed hunters harvested an estimated 82,096 moose in 2014–2015; a decade earlier, the harvest was 83,246 moose by 386,419 licensed hun- ters (Table 1). Hunting regulations continue to become more restrictive and complex as the demand on moose populations and corre- sponding harvest success rates increase, due in part, to increased road access and use of mechanized equipment (Timmermann and Buss 1998). Specific and strategic management of hunting is required to affect the desired allocation of moose harvest among licensed hunters, secure the sustain- ability of moose populations, and achieve other specified management objectives for a particular area. Specific moose management plans, guidelines, or statements existed in 13 jurisdictions in 2000–2001 (Maine, Vermont, New Hampshire, Utah, Colorado, Wyoming, Idaho, Yukon Territory, British Columbia, Alberta, Saskatchewan, Ontario and Quebec; Timmerman 2003). Specific harvest policy is currently guided by an approved or draft management policy including goals and objectives in 13 jurisdictions; 3 employ un- written or a generalized wildlife policy. For example, Alaska’s constitution, statutes, and regulations direct management activities and objectives through a public process (B. Dale, pers. comm. 2015), Minnesota’s policy is guided by a research and management plan (McGraw et al. 2010, Minnesota DNRC 2011, Moen et al. 2011b), and Colorado uses a specific management plan for each of 10 herds (A. Holland, pers. comm. 2015). British Columbia has recently developed a provincial guidance and direction frame- work for sustainable moose management (British Columbia Fish &Wildlife Branch 2015). Beginning in 2007, Ontario con- ducted a 2-year broad review and wide consultation of their moose management program that produced a new set of policies and guidelines with objectives and strategies to address the declining population and har- vest (OMNRF 2008). Two options to control calf harvests included a shorter calf season within the regular season and a draw for calf tags (Bottan et al. 2002, Timmermann et al. 2002, OMNRF 2009a, b). A moose management plan has been developed in Newfoundland and Labrador that will help address human-wildlife conflicts (J. Neville, pers. comm. 2015), and Quebec currently employs a fourth iteration of a manage- ment plan spanning the period 2012–2019 8 STATUS OF MOOSE IN NA CIRCA 2015 – TIMMERMANN ALCES VOL. 53, 2017 AND RODGERS (S. Lefort, pers. comm. 2015). Saskatchewan and Manitoba are developing specific man- agement plans (H. Hristienko and R. Tether, pers. comm. 2015). Allocation of Hunting Opportunities Moose are publicly owned and held in trust by provincial, territorial, and state wild- life agencies. The first priority of most agen- cies is to ensure the long-term conservation of moose populations and their habitats. Harvest allocation is given prime consider- ation to subsistence use by Indigenous people under Treaty or other legal agreements in at least 20 of 23 jurisdictions that currently manage a harvest. Resident hunters in 20 of 23 jurisdictions are typically favored over non-residents and non-resident foreigners (10 of 23) in allocating harvest opportunities. In 2014–2015, non-residents were eligible to hunt in 20 of 23 jurisdictions (Table 1). Add- itional controls such as increased license fees, resident-only seasons, guide requirements, and limited permits are commonly placed on non-resident hunters giving residents priority in allocation of hunting opportunities. A guide was required by 10 of 23 agencies, and at least 6 agencies required non-residents to register with a licensed tourist outfitter, and 8 required foreigners to do so to enhance safety and success, as well as provide local economic benefit. Some agencies restrict or limit moose hunting opportunities including all states ex- cept Alaska. Washington and North Dakota offer a single moose hunt per lifetime, and Colorado, Utah, and Idaho limit hunters to one antlered animal per lifetime. Others require a waiting period between hunts: 2 years in Idaho, 3 years in New Hampshire, Maine, Saskatchewan, and Manitoba, 5 years in Wyoming, Vermont, and Nova Scotia, and 7 years in Montana (if successful). Hunters in Alaska and 8 Canadian jurisdictions may hunt annually within quotas regardless of previous harvest success. Minnesota closed the entire state to moose hunting in 2013 after the northeastern population declined by half since 2006. Manitoba legislated 3 Conserva- tion Closure Game Hunting Areas and 1 partial closure in 2011, coupled with a wolf reduction initiative to promote moose recovery (H. Hristienko, pers. comm. 2015, V. Crichton, pers. comm. 2016). Ontario continues to offer moose hunting opportunities for physically- challenged hunters in one Wildlife Manage- ment Unit (Armstrong and Simons 1999). Control Concepts Agencies employ a variety of strategies to regulate harvests and distribute hunting pressure (Timmermann 1987, 2003). Passive strategies include season length and timing, access restrictions, weapon requirements, and license qualification prerequisites; active measures include limiting license sales or specifying the sex, age, or number of animals taken by specific area. Objectives often in- clude the harvest of pre-determined numbers to sustain, increase, or reduce populations. In Alberta, Xu and Boyce (2010) developed an age-sex matrix model for harvest quota management of moose populations that allows easy application by managers respon- sible for setting harvest quotas. Antler-based hunting regulations in British Columbia may have resulted in disrupted reproductive patterns and a consequent over-harvest of large bulls (Child et al. 2010). In interior Al- aska, harvest restrictions on bull moose based on antler architecture allowed the recovery of bull:cow ratios from 26:100 to 32:100 after only 2 years of use (Young and Boertje 2008). Conversely, liberal antlerless hunts in Alaska are considered vital to control moose populations from reaching unsustainable densities in specific management units - a general season harvest ticket is available to all residents/non-residents (Young and Boertje 2004, Young et al. 2006, Boertje et al. 2007, 2009, Young and Boertje 2011). ALCES VOL. 53, 2017 TIMMERMANN AND RODGERS – STATUS OF MOOSE IN NA CIRCA 2015 9 A specific controlled hunt to reduce a local moose population and impacts on cole crops was implemented in Maine in 2009 (Kantar 2011). New Hampshire and Vermont have increased regional/local harvest rates with higher antlerless quotas to alleviate browsing impacts on regenerating forests and vehicular collisions (C. Alexander, pers. comm. 2002, Andreozzi et al. 2014). In 2014–2015, 9 agencies offered unlimited se- lective or non-selective harvest opportunities, and all 23 jurisdictions restricted or limited harvests on a selective or non-selective basis in certain management areas (Fig. 2). In addition, closed seasons were employed to prevent licensed harvest in specific areas, including certain provinces, territories, states, National Parks, and Wildlife Refuges. License qualifications and fees In 2014, proof of hunting proficiency, ei- ther a previous license or completing a hunter safety education course, was required to ob- tain a moose hunting license in all jurisdic- tions. In 2015, Canadian resident license fees averaged CAN $52.57 (range $5.00 in the Yukon Territory to $81.30 in New Bruns- wick), and non-resident licenses averaged $374.45 (range $48.00 in Quebec to $619.24 in New Brunswick). Resident fees in the states averaged US $162.00, (range = $25.00 in Alaska to $ 413.00 in Utah), and non-resident fees averaged $1,168.03 (range = $350.00 in Vermont to $2,271.25 in Idaho). Some agencies, including Alaska and Maine, charged higher fees to non-resident foreign- ers. Export permits or trophy fees are required, in addition to the license fee, to transport an animal from the Yukon Territory, Northwest Territories, British Columbia, Alberta, and Ontario. Currently, only New Hampshire requires moose hunters to demon- strate shooting proficiency using convention- al fire-arms prior to purchasing a hunting license, as described by Buss et al. (1989). Previously, New Brunswick and Newfoundland required hunters to pass a shooting and written test before qualifying for a big game hunting license (Timmermann and Buss 1995). Al- aska requires all archery and black powder hunters to pass a proficiency test (W. Regelin, pers. comm. 2002). Seasons Season length and timing are used to manage the availability of hunting opportun- ity, hunter success relative to vulnerability based on moose behavior, and seasonal access. Seasons are generally specific to weapon type (e.g., conventional firearms, black powder, or archery), and seasons tend to be longer in more remote areas and shorter closer to population centers. The most liberal season length (365 days, 1 July-30 June) occurs in 3 Game Management Areas in Nunavut (Table 2). Season lengths for all hunts in parts of Alaska, Idaho, Wyoming, Montana, North Dakota, Yukon, Northwest Territories, British Columbia, Alberta, Saskatchewan, Manitoba, Ontario, Quebec, and Newfoundland equal or exceed 3 months; New Brunswick, Nova Scotia, Vermont, and New Hampshire restrict season length to 5–9 days. Early archery seasons are typically in addition to firearm seasons, and are offered in 17 jurisdictions (Table 2). Most firearm seasons begin during the latter portion of the rut period (Wilton 1995) and many extend into November or December. Split seasons (early vs. late fall) occur in at least 11 jurisdic- tions. Minnesota closed their northeast moose hunt in 2013 after a continuous 41-year period of offering a limited non-selective hunt requir- ing all eligible hunters to apply in groups of up to 4 individuals (Judd 1972). Management areas and harvest strategies All agencies have subdivided their moose range into various sized management areas (Wildlife, Game, or Moose Management Units) to facilitate specific harvest control 10 STATUS OF MOOSE IN NA CIRCA 2015 – TIMMERMANN ALCES VOL. 53, 2017 AND RODGERS F ig . 2 . M o o se h ar v es t st ra te g ie s em p lo y ed b y 3 0 N o rt h A m er ic an ju ri sd ic ti o n s (c ir ca 2 0 1 4 – 2 0 1 5 ). N u m b er s in d ic at e m an ag em en t ar ea s o r su b d iv is io n s u n d er ea ch h ar v es t st ra te g y in ea ch ju ri sd ic ti o n . ALCES VOL. 53, 2017 TIMMERMANN AND RODGERS – STATUS OF MOOSE IN NA CIRCA 2015 11 strategies. The number of management areas ranges from 3 in Nunavut to 443 in the Yukon, and vary in size from 47 km2 in North Dakota to 278,183 km2 in the Northwest Ter- ritories (Table 2). All jurisdictions except Nunavut continue to employ either a selective or non-selective limited hunter participation strategy, or a combination of both (Fig. 2). Most favor some form of limited selective or limited non-selective strategy to control sex- and age-related harvests. Alaska alone continues to employ registration hunts which require mandatory kill registration and sea- son termination once a prescribed harvest is achieved. Several agencies have developed harvest strategies that maximize hunter participation. For example, sharing a moose between 2 or more hunters optimizes hunting opportunities and accommodates hunters who wish to hunt with friends. Perhaps the most liberal ap- proach is in Ontario which previously allowed all eligible hunters to hunt calves in any of 64 Wildlife Management Units (Timmermann Table 2. Characteristics of moose hunting seasons in North America, 2014–2015. Number of management areas Season length/timing Size (km2) Agency With moose Min. Max. With open Season Max days Earliest Latest Yukon Territory 443 64 2,919 360 92 Aug. 01 Oct. 31 Northwest Territories 6 56,270 278,183 61 123 Sept. 01 Jan. 31 Nunavut 3 N/A N/A 1 365 July 01 June 30 British Columbia 193 465 18,982 1771 107 Aug. 15 Nov. 30 Alberta 177 99 26,079 1631 91 Aug. 25 Nov. 30 Saskatchewan 82 232 82,443 801 92 Sept. 01 Nov. 30 Manitoba 60 222 139,204 291 117 Aug. 31 Jan. 24 Ontario 76 832 122,397 691 93 Sept. 14 Dec. 15 Quebec 30 1,471 204,142 281 92 Aug. 27 Dec. 01 New Brunswick 25 826 6,402 25 10 Sept. 20 Sept. 30 Nova Scotia 5 204 2,700 51 6 Sept. 29 Dec. 11 Newfoundland 54 217 4,533 541 121 Aug. 29 Jan. 24 Alaska 22 9,117 217,559 221 243 July 01 April 15 Washington 11 747 2,857 111 60 Oct. 01 Nov. 30 Idaho 70 220 7,843 461 86 Aug. 30 Dec. 01 Utah 15 809 5,394 12 34 Sept. 12 Oct. 15 Wyoming 38 100 10,000 311 81 Sept. 01 Nov. 20 Montana 88 95 60,926 821 87 Sept. 15 Nov. 30 North Dakota 7 47 36,514 51 101 Sept. 04 Dec. 13 Colorado 65 130 1,540 571 25 Sept. 12 Oct. 14 Minnesota 30 200 875 — season closed - 2013 Maine 28 1,424 5,320 25 24 Sept. 28 Nov. 28 Vermont 21 639 2,036 161 6 Oct. 1 Oct. 26 New Hampshire 22 391 1,365 20 9 Oct. 18 Oct. 26 1 Offers early archery season. 12 STATUS OF MOOSE IN NA CIRCA 2015 – TIMMERMANN ALCES VOL. 53, 2017 AND RODGERS 2003, Fig. 2). In 2015 calf hunting was reduced to a 2-week period, and beginning in 2016, opening seasons will be delayed by 1 week (OMNRF 2016). In addition, Ontario hunters may apply in groups of up to 15 hunters for the chance of obtaining an adult tag that is area-specific, allowing a limited number of tags to be spread more evenly among hunter groups (OMNRF 2015, 2016). Sharing a moose is currently required in at least 2 jurisdictions including Quebec (mini- mum 2 hunters/moose) and Manitoba which provide an option of purchasing a conserva- tion license (2 hunters using a single tag; S. Lefort and V. Crichton, pers. comm. 2015). Timmermann (2003) detailed additional moose-sharing mechanisms including a “Group Hunt” and a “Limited Entry Shared Hunt” in British Columbia, a “Special Antlered Moose Partner License” in Alberta, and a “Companion Moose Hunting Stamp” in Nova Scotia. Each successful permittee in Maine, New Hampshire, and Vermont may select a sub-permittee to hunt together and harvest a single moose. To help maximize hunter oppor- tunity, Newfoundland gives preference to party applications (restricted to 2 individuals) over individuals, and to those unsuccessful in previ- ous years (Timmermann 2003). Yukon has “Special Guide Licenses” that annually allow 100 Yukon residents to guide a non-resident hunter, in part to accommodate family mem- bers/friends from outside the Territory (S. Czetwertynski, pers. comm. 2016). Harvest assessment All sources of mortality must be assessed to monitor the effectiveness of various har- vest strategies. Hunters, whether successful or not, are required to report their hunting ac- tivity in 11 of 23 jurisdictions, and harvest registration is compulsory in 13 (Table 3). Twelve jurisdictions apply a non-compliance penalty to hunters failing to report, although enforcement of these requirements varies among agencies. Timmermann (2003) provided a more detailed description of this subject, including the use of interactive voice response technology, use of a telephone ques- tionnaire, and modeling to predict population changes resulting from various harvest strategies. Moose hunter education and engagement All first time hunters are required to suc- cessfully complete a hunter safety/education course in 23 jurisdictions that managed a moose hunt in 2014. Most (19 of 23) charged fees ranging in Canada from no fee in Yukon to CAN $160 in Ontario, and in the United States from no fee in Washington and Vermont, US $5 in New Hampshire, and $10 in Utah and Wyoming. Four states (Idaho, Utah, Vermont, and New Hampshire) incorporate a practical shooting test in their Hunter Safety/Education Course. All jurisdictions (22 of 23) except Nunavut provide moose hunters with infor- mation on their official websites, and 9 use social media. Printed hunting regulations were available in 22 of 23 jurisdictions, and 15 provided printed pamphlets, brochures, and/or fact sheets. Television and/or radio was used to provide information in 4 jurisdic- tions, 10 used newspapers and/or magazines, and 17 used email and/or traditional mail. Harvest by native and subsistence users Currently, most North American moose management agencies give primary consider- ation to subsistence use by Canadian Indigen- ous peoples and Native American peoples in recognition of obligations made under histor- ical treaties signed by both federal govern- ments (Crichton et al. 1998, Lynch 2006). Currently, nine 9 of 24 jurisdictions (Yukon, Northwest Territories, British Columbia, Manitoba, Ontario, Quebec, Nova Scotia, Idaho, Montana) report primary allocation of the moose resource to subsistence use by Indigenous People under Treaty or other legal agreements. In many areas unrestricted ALCES VOL. 53, 2017 TIMMERMANN AND RODGERS – STATUS OF MOOSE IN NA CIRCA 2015 13 access to moose exists year-round, and cur- rent regulations are considered liberal given the widespread use of modern technology (Courtois and Beaumont 1999). Because con- flicts often occur between licensed sport hun- ters and Indigenous People, moose managers must consider the annual harvest by both groups in formulating hunting regulations (Lynch 2006). The harvest by Indigenous hunters is difficult to quantify and unfortu- nately, little effort has been made to measure the magnitude of this harvest which some managers believe approaches or exceeds the licensed harvest in certain jurisdictions. Metis, who are considered people of mixed Indian and White ancestry (Swail 1996), are testing their perceived rights in court in Al- berta, Manitoba, and Nova Scotia where they claim the right to hunt and fish on trad- itional territory both within and outside their current harvesting zone (V. Crichton, pers. comm. 2016). The Supreme Court of Canada Table 3. Moose harvest assessment strategies used in North America, 2014–2015. Hunt activity report Kill registration Non-compliance penalty Agency Compulsory Voluntary Compulsory Voluntary Yukon Territory X X1 Fine - CDN$100.00 Northwest Territories X X1 none Nunavut X X Fine - CDN$200.00 British Columbia X X Fine - CDN$230.00 Alberta none X N/A Saskatchewan X none N/A Manitoba X none N/A Ontario2 2 X 2 none1 N/A2 Quebec none X Fine- CDN$250-750.00 New Brunswick X X Fine- CDN $100-500.00 Nova Scotia X X N/A Newfoundland X X none Alaska X X Loss of future eligibility Washington X X Fine - US$25.00 Idaho X X Fine - US$25.00 - 1,000 & jail Utah X none Ineligible to apply next year Wyoming X X none Montana X X N/A North Dakota X X Ineligible to apply next year Colorado X X Loss of future eligibility Minnesota Season Closed 2013 Maine X X Fine - US$100-1,000 / Lic. loss Vermont X X Fine- US$262 + & Lic. loss New Hampshire X X Fine- US$248-1,000 & loss of future eligibility 1 Export permit/trophy fee. 2 Compulsory hunt activity/harvest report in 5 Wildlife Management Units; $150.00 fine for non-compliance and inability to receive a tag in subsequent year. 14 STATUS OF MOOSE IN NA CIRCA 2015 – TIMMERMANN ALCES VOL. 53, 2017 AND RODGERS has refused to hear an appeal involving Metis hunting and fishing rights in Alberta follow- ing a Supreme Court ruling 10 years previous that granted hunting rights to Ontario Metis. However, in April 2016 the Supreme Court declared that the federal government has con- stitutional responsibility for Métis and non- status Indians, which could have important implications for their hunting and fishing rights. Timmermann (2003) provided esti- mates of the annual moose harvest by Indigen- ous and Metis peoples in British Columbia, Alberta, Saskatchewan, Manitoba, Ontario, Quebec, New Brunswick, Nova Scotia, North- west Territories, and the Yukon Territory. A moose monitoring program was estab- lished in the Northwest Territories between the Dehcho First Nations and government biologists that yielded valuable biological in- formation to identify changes in moose popu- lations (Larter 2009). In a similar effort to involve Indigenous people in northwestern Ontario, LeBlanc et al. (2011) suggested that provincial calculations may underesti- mate total harvests by up to 40%, and concluded that developing a working rela- tionship with Indigenous communities is necessary to effectively manage moose in Ontario. A moose management plan was drafted by the Nova Scotia Mi’Kmaq in 2005 to aid in tripartite negotiations between the band and the provincial and federal gov- ernments (Bridgland et al. 2007). The Nova Scotia DNR is currently working with the Nova Scotia Mi'Kmaq to develop a collab- orative management plan for Cape Breton moose (P. MacDonald, pers. comm. 2016). It is hoped such collaborative management will sustain moose populations long-term on Cape Breton Island. Cooperative manage- ment of moose between state and 3 tribal bands in northeastern Minnesota has led to increased levels of trust since 1988 (Edwards et al. 2004). In Colorado, moose harvested by Native and subsistence hunters is monitored by the Brunot Agreement with the Southern Ute and Ute Mountain Tribes (A. Holland, pers. comm. 2015). Future sustainable har- vests and population goals will largely remain elusive until the total harvest, includ- ing harvests by Indigenous and Metis peoples and subsistence users, are both agreed to and verifiable. Under federal regulations in Alaska, all rural residents are subsistence users which allows certain communities or households to harvest moose. In addition, special regula- tions allow moose harvesting outside of nor- mal hunting seasons by Alaskan natives for ceremonial and cultural purposes. All agen- cies suggested such harvests were “substan- tial” in specific local areas during the period 2000–2001. Oregon has 2 native tribes which hunt ungulates in areas where moose occur; one tribe hunts moose and the other is consid- ering a moose season (P. Matthews, pers. comm., 2016). Illegal hunting losses appear to be signifi- cant in some jurisdictions including Colorado, Utah, and Ontario (Timmermann 2003). Most agencies encourage all hunters to report illegal infractions using a toll-free telephone number. Ontario introduced a “Moose Watch” Program in 2001 to help reduce poaching (Todesco 2004). Conservation Officers in Ontario’s Northeast Region found 1,741 illegally killed moose from 1997–2014 (Todesco 2004, C. Todesco, pers. comm. 2016). During this per- iod, >238,000 hunters were contacted, 7,328 warnings issued, and 5,514 charges were laid while conducting moose hunt enforcement duties. MANAGING UNHUNTED POPULATIONS Parks, refuges, and special areas Most North American jurisdictions where moose occur provide for areas where hunting is not a primary management object- ive. Currently, 7 states have no open moose hunting season and 11 of 23 jurisdictions pro- vide closed seasons in 2–33 management ALCES VOL. 53, 2017 TIMMERMANN AND RODGERS – STATUS OF MOOSE IN NA CIRCA 2015 15 areas (Fig. 2). The assumed common object- ive of closed areas that include game or wild- life reserves, national, provincial, territorial, and state parks, and nature reserves is the preservation of moose in representative nat- ural habitats for education and recreational enjoyment. Further maintenance of biodiver- sity and ecosystem function is often a stated objective. A review of moose management objectives and programs in parks, refuges, and special areas was detailed by Timmermann and Buss (1995) and Timmermann (2003). Moose are native to at least 30 North American national parks in 18 jurisdictions with Isle Royale perhaps the most famous, boasting a 58-year continuous ecological study of wolves and moose beginning in 1959 (Vucetich and Peterson 2015). Timmermann (2003) detailed moose-related studies in several National Parks including Isle Royale in Michigan, Elk Island in Alberta, Voyageurs in Minnesota, and Gros Morne in Newfoundland. West (2009) reported approximately 39,000 moose inhabited 35 National Wildlife Refuges in the United States, with ~38,000 in Alaska alone; 9 refuges used management practices to specifically benefit moose (i.e., prescribed wildland fire). DISCUSSION Telfer (1984) found a close correspond- ence between the southern limit of moose distribution worldwide and the 20 °C July isotherm. Since then, several studies have suggested that moose numbers and the south- ern limit of their distribution may be affected by climate change (Thompson et al. 1998, Murray et al. 2006, Lenarz et al. 2010) but others have documented increasing numbers and expanding moose populations at the southern edges of their jurisdictional bound- ary (Wattles and DeStefano 2011, Harris et al. 2015, LaForge et al. 2016). Our survey across North America further indicated that 10 of 15 moose populations at the southern limit of their distribution are stable or increasing. Although we acknowledge that many of the reported population estimates and trends were based on professional opinion rather than systematic surveys, the weight of evi- dence suggests that moose are not immediate- ly at risk of disappearing from southern regions of their distribution. The more inter- esting enquiry at this time would be to deter- mine how and why moose continue to thrive and even expand their range southward in certain areas. Moving forward in the face of climate change and ever-increasing human develop- ment, it will be vitally important to maintain systematic aerial surveys to monitor moose population trends, and to implement these wherever they are not in use. This will be a challenge not only because of the financial and human resources required, but also be- cause climate change may hinder the collec- tion of long-term data due to lack of snow required to efficiently conduct aerial surveys. Thus, further research into new technologies that are not hindered as much by environ- mental conditions will be highly beneficial, such as forward-looking infrared radiometer systems (FLIR; Millette et al. 2014) and sen- sor-equipped drones. Our survey revealed variation in moose population trends across North America, and local variation expected within jurisdic- tions. As indicated by survey respondents, numerous factors can affect moose popula- tion trends both locally and regionally. Although research might be undertaken to identify the most important factors in a par- ticular area, the responses of local moose managers are limited because many cannot be controlled directly (e.g., parasites and disease, weather). Instead, most moose managers can only use harvest and habitat management to mitigate declines in moose population numbers. These options will be- come ever more important as climate change and human development gradually increase their influence on moose numbers and distri- bution across North America. 16 STATUS OF MOOSE IN NA CIRCA 2015 – TIMMERMANN ALCES VOL. 53, 2017 AND RODGERS ACKNOWLEDGEMENTS Appreciation is extended to the following individuals in the United States who provided unpublished information in response to a 9-page questionnaire survey: Bruce Dale and Kim Titus, Alaska Department of Fish and Game, Juneau, Alaska; Rich Harris, Washington Department of Fish and Wildlife, Olympia, Washington; Pat Matthews, Oregon Department of Fish and Wildlife, Enterprise, Oregon; Steve Nadeau, Idaho Fish and Game, Boise, Idaho; Kent Hersey, Division of Wildlife Resources, Salt Lake City, Utah; Andy Holland, Colorado Parks and Wildlife, Fort Collins, Colorado; Doug Brimeyer, Wyoming Game and Fish Depart- ment, Jackson, Wyoming; Nick DeCesare, Montana Fish, Wildlife and Parks, Missoula, Montana; Jason Smith, North Dakota Game and Fish Department, Jamestown, North Dakota; Glenn DelGiudice and Leslie McInenly, Minnesota Department of Natural Resources, Forest Lake, Minnesota; Dan Storm and Kevin Wallenfang, Department of Natural Resources, Rhinelander, Wisconsin; Dean Beyer, Michigan Department of Natural Resources and Northern Michigan University, Marquette, Michigan; Lee Kantar, Maine Department of Inland Fisheries and Wildlife, Bangor, Maine; Kristine Rines, New Hamp- shire Fish and Game Department, New Hampton, New Hampshire; Cedric Alexander, Vermont Fish and Wildlife Department, St. Johnsbury, Vermont; Andrew LaBonte, Depart- ment of Energy and Environmental Protection, North Franklin, Connecticut; David Scarpitti, Massachusetts Division of Fisheries and Wild- life, West Boylston, Massachusetts; Steven Heerkens, Department of Environmental Con- servation, Utica, New York. Appreciation is extended to the following individuals in Canada who provided unpub- lished information in response to a 9-page questionnaire survey: Sophie Czetwertynski and Rob Florkiewicz, Environment Yukon, Whitehorse, Yukon; Angus Smith and Jan Adamczewski, Department of Environment and Natural Resources, Government of North- west Territories, Yellowknife, Northwest Territories; Mathieu Dumond, Department of Environment, Arviat, Nunavut; Gerry Kuzyk, Ministry of Forests, Lands and Natural Resources Operations, Victoria, British Columbia; Rob Corrigan, , Environ- ment and Parks, Edmonton, Alberta; Rob Tether and Mike Gollop, Saskatchewan Ministry of Environment, Saskatoon, Saskatchewan; Hank Hristienko and Ken Rebizant, Manitoba Conservation, Winnipeg, Manitoba; Vince Crichton, retired Manitoba Conservation, Winnipeg, Manitoba; Greg Lucking and Patrick Hubert, Ontario Ministry of Natural Resources and Forestry, Peterborough, Ontario; Charlie Todesco, Ontario Ministry of Natural Resources and Forestry, Wawa, Ontario; Sébastien Lefort, Quebec Ministère des Forêts, de la Faune et des Parks, Quebec City, Quebec; Dwayne Sabine, New Brunswick Department of Natural Resources, Fredericton, New Bruns- wick; Peter MacDonald, Nova Scotia Depart- ment of Natural Resources, Kentville, Nova Scotia; John Neville and Conor Edwards, Newfoundland and Labrador Department of Environment and Conservation, Corner Brook, Newfoundland. 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Alces 46: 15–35. 22 STATUS OF MOOSE IN NA CIRCA 2015 – TIMMERMANN ALCES VOL. 53, 2017 AND RODGERS THE STATUS AND MANAGEMENT OF MOOSE IN NORTH AMERICA -CIRCA 2015 HISTORICAL DISTRIBUTION AND CURRENT STATUS Eastern North America Western North America Central North America Factors affecting decreasing densities HARVEST MANAGEMENT Economic impact Harvest control objectives Allocation of Hunting Opportunities Control Concepts License qualifications and fees Seasons Management areas and harvest strategies Harvest assessment Moose hunter education and engagement Harvest by native and subsistence users MANAGING UNHUNTED POPULATIONS Parks, refuges, and special areas DISCUSSION ACKNOWLEDGEMENTS REFERENCES