PROVINCIAL POPULATION AND HARVEST ESTIMATES OF MOOSE IN BRITISH COLUMBIA Gerald W. Kuzyk Ministry of Forests, Lands and Natural Resource Operations, P.O. Box 9391, Victoria, British Columbia V8W 9M8, Canada ABSTRACT: Provincial population and harvest estimates of moose in British Columbia, Canada were assessed over a 28-year period from 1987 to 2014. The population generally remained stable, whereas the licensed hunter harvest declined gradually by about half despite constant hunter effort. The annual population estimate ranged from a low of 157,000 moose in 1994 to a high of 190,000 in 2011, with an overall mean of 172,000 ± 9900 (SD). In 2014, the relative status of hunted popula- tions within 7 wildlife administrative units was 1 increasing, 3 stable, and 3 in decline. The mean an- nual licensed harvest was 10,038 ± 2137 (SD) moose, and the mean harvest rate was 6 ± 1.3% (SD). In December 2013, British Columbia initiated a 5-year (2013–2018) research project to identify factors contributing to the decline of the moose population and licensed harvest. ALCES VOL. 52: 1–11 (2016) Key words: Alces alces, British Columbia, harvest, moose, population Periodic updates of moose (Alces alces) abundance are necessary to assess manage- ment objectives (Brown 2011), evaluate sus- tainable harvest (Timmerman and Buss 2007), and to provide information to the public. Assessing licensed harvest concur- rent with population estimates should pro- vide better understanding and explanation of population fluctuations over time. Moose population estimates are also used for com- parison among jurisdictions to assess pat- terns of broad-scale population trends. In North America, there is current concern for declining populations in southern parts of moose range (Murray et al. 2006, Lenarz et al. 2009), whereas populations remain stable in other areas (Murray et al. 2012). Explanations for population change include human-caused habitat alterations (Rempel et al. 1997), climate change (Rempel 2011), and a combination of natural and human- influenced variables (Murray et al. 2006, Brown 2011). Moose in British Columbia are highly valued for food, social, and ceremonial pur- poses by First Nations, for recreational and commercial harvest opportunities by licensed hunters, and for wildlife viewing. Specific management objectives for moose harvest are to manage for First Nations use, support a sustainable licensed hunter harvest, and pro- vide for diverse hunter opportunities (BC FLNRO 2015). Assessment of abundance and licensed harvest estimates is required to ensure that harvest levels are sustainable (Hatter 1999), objective information is avail- able for management decisions, and to pro- vide accurate information on the status of moose to stakeholders and the public (BC FLNRO 2015). The purpose of this paper is to provide an overview of the population abundance and licensed harvest of moose in British Columbia from 1987 to 2014. STUDY AREA British Columbia is an ecologically di- verse province (Meidinger and Pojar 1991) 1 where moose are widely distributed (Fig. 1) and occupy a range of landscapes including wet coastal habitats, dry interior forests, cold northern forests, and montane habitats (Eastman and Ritcey 1987). At the provincial scale, moose co-exist with several ungulate species including bison (Bison bison), mule deer (Odocoileus hemionus), white-tailed deer (Odocoileus virginianus), elk (Cervus elaphus), and caribou (Rangifer tarandus) (Shackleton 1999). The main predators of moose are wolves (Canis lupus), grizzly bears (Ursus arctos), and black bears (U. americanus), with cougars (Puma concolor) important in southern British Columbia (Spalding and Lesowski 1971). Bull hunts were mostly open seasons, with antler restric- tions or limited entry hunts occurring between 15 August and 30 November. Antlerless harvest was largely restricted to limited entry hunts with some general open seasons for calves in select areas. Seasons for antlerless moose occurred between 1 Oc- tober and 10 December (BC MOE 2010). Fig. 1. Distribution and population status (i.e., stable, increasing, decreasing) of moose in 7 wildlife administrative units in British Columbia, Canada, 2014. 2 POPULATIONS AND HARVEST OF MOOSE IN BC – KUZYK ALCES VOL. 52, 2016 Hunting seasons were generally available throughout the distribution of moose with the exception of Regions 1 and 2 which have few moose (i.e., <130 combined) and National Parks (<1% of land area) where licensed hunting is prohibited. METHODS Moose population estimates were pro- duced by regional biologists in 7 wildlife administration units (Regions; Fig. 1) from 1987 to 2014, and then combined for a pro- vincial total. There were 3–5 year intervals between estimates to provide time to assess potential changes in moose abundance at the provincial scale. Minimum and maximum estimates were derived from 2000 to 2014 be- cause of the need to convey uncertainty when comparing estimates between years. These estimates were developed using the best available information from a combination of sources including aerial surveys, big game stock assessments, and expert opinion. A third degree polynomial was used to fit a long-term population trend line to the abundance estimates from 1987 to 2014. The polynomial was preferred to a linear or log-linear trend line because the polynomial was sensitive to fluctuations in population size. In the 7 regions where moose were hunted, the trend (stable, declining, increas- ing) was determined from the change in abundance estimates and the slope of the trend line from 2011 to 2014. Aerial surveys were the most important source of information because they provided data for estimation of population size, dens- ity, and composition. All surveys were required to follow provincial standards that are based on defensible scientific methods (RISC 2002). Stratified random block sur- veys were used (Gasaway et al. 1986) or modified to include habitat-based stratifica- tion (Heard et al. 2008). A standard sightabil- ity correction factor was applied to account for detection probability based on research with radio-marked moose in central British Columbia (Quayle et al. 2001). Aerial sur- veys were required to conform to standards for accuracy and precision (1-α) and to produce a 90% CI with allowable error (±15–25%). The frequency of stratified ran- dom block surveys was based on available funds and prioritization criteria which in‐ cluded time since last survey, First Nations concerns, impact to hunter opportunity, population objectives, and if the survey was part of an ongoing monitoring program (BC FLNRO 2015). Aerial composition surveys were also conducted to determine bull:cow and calf:cow ratios; ground-based surveys following provincial standards were used oc- casionally (RISC 1998, D’ Eon et al. 2006). Big game stock assessments were used to help estimate population size and sustain- able harvest levels as outlined in the provin- cial moose harvest management procedure (BC MOE 2010). These assessments helped maximize information from aerial surveys and hunter harvest (Griffiths and Hatter 2011), and incorporated uncertainty asso- ciated with extrapolating area-based survey results to regional population estimates. They helped determine the maximum al- lowable mortality and accounted for First Nations harvest and road/rail mortality where available. Population models were one component of big game stock assess- ments and were occasionally used in the regional population estimates by fitting an- nual licensed harvest data to periodic survey data (White and Lubow 2002). Population variables used in the models generally included annual licensed harvest data, post- hunt population size and composition, over- winter survival, and recruitment rates (Griffiths and Hatter 2011). If empirical in- formation was lacking about a population, re- gional biologists used a broad spectrum of expert opinion including field information gathered from resident hunters and trappers, guide-outfitters, First Nations, and other ALCES VOL. 52, 2016 KUZYK – POPULATIONS AND HARVEST OF MOOSE IN BC 3 resource professionals. This information was gathered during a variety of forums and loca- tions including formal stakeholder meetings and informal discussions. Licensed harvest of moose was moni- tored annually from 1987 to 2014 with a pro- vincial resident hunter survey, and guide declarations for non-resident hunters. Har- vest information from First Nations was not part of the provincial hunter survey and was largely unknown (BC FLNRO 2015), with the exception of certain First Nations com- munities that voluntarily provided informa- tion. Estimates of licensed hunter harvest (resident and non-resident combined), hunter days, and hunter numbers were available, all with 95% confidence intervals (CI). These estimates were produced from mail-out ques- tionnaires sent to a random sample of resident hunters; from 2008 to 2014 an average of 13,003 questionnaires were mailed annually with an average response rate of 61%. Licensed harvest rates were calculated from the provincial population estimate for a given year and the average of the 3 nearest harvest estimates; 2014 was an exception when the average of the 2 nearest harvest estimates were used because of delay in the 2015 estimate. Combined resident and non- resident hunting license sales from 1989 to 2014 were used to further measure hunter interest. RESULTS The mean annual population estimate of moose in British Columbia was 172,000 ± 9900 (SD) from 1987 to 2014. Annual esti- mates were relatively stable ranging from a low of 157,000 moose in 1994 to a high of 190,000 in 2011 (Fig. 2). The minimum and maximum estimates (i.e, from 2000 to 2014) reflected varied levels of uncertainty (Fig. 2). The 2014 estimates varied among the 7 regions with hunted populations: 3 were considered stable (Regions 3, 6, and 7B; Figs. 1 and 3), 3 were declining (Regions 4, 5, and 7A; Figs. 1 and 4), and one region was increasing (Region 8; Figs. 1 and 3). The mean annual licensed harvest from 1987 to 2014 was estimated as 10,038 ± 2137 (SD). Total harvest declined gradually by about one-half during this period, yet hunter effort (average days hunted) remained stable (Fig. 5). The mean annual licensed harvest rate from 1987 to 2014 was 6 ± 1.3% (SD), ranging two-fold from a high of 8% in 1987 to a low of 4% in 2011. From 1987 to 2014, the mean number of licensed hunters (resident and non-resident combined) was 33,721 ± 4292 (SD) that spent 273,622 ± 32,521 (SD) days of hunter effort (Table 1). The mean annual hunting license sales was 39,815 ± 4158 (SD) from 1989 to 2014 and varied minimally from 1993 to 2014 (Table 1). DISCUSSION The annual moose population in British Columbia during 1987–2014 was relatively stable, averaging 172,000. In 2014 hunted populations were stable in 3 regions, decreas- ing in 3 regions, and increasing in one. Although both provincial and regional popula- tion estimates had varied levels of uncertainty, they remain important for resource mana- gers to address management objectives (BC FLNRO 2015), and to inform First Nations, stakeholders, and the general public about the status of moose in British Columbia. The estimation error was partially re- sponsible for the uncertainty reported in the abundance estimates. The variation in the population estimates may reflect the varied abundance and composition of local and regional predators (Ballard and Van Ballenberghe 2007), human-altered land- scape change (Rempel et al. 1997) which may enhance forage quality and quantity while facilitating predator and hunter access to moose, and variation in licensed and un- licensed harvest levels (Timmerman and 4 POPULATIONS AND HARVEST OF MOOSE IN BC – KUZYK ALCES VOL. 52, 2016 Buss 2007). Other factors such as weather, disease, parasites, and accidents including road and rail mortality also influence local moose abundance. The quality of data used to develop the population estimates could also be improved with increased financial and logistical support that would provide more aerial surveys over a broader geograph- ical area. Of most concern to stakeholders were re- cent (2008–2014) population declines in Regions 4, 5, and 7A (Fig. 4). In two regions (Region 5 and 7A) the moose declines coin- cided with a mountain pine beetle (Dendroc- tonus ponderosae) epidemic (Chan-McLeod 2006) which led to increased salvage logging and associated road building. This type of landscape change can presumably alter the spatial dynamics of moose, predators, and hunters, ultimately influencing moose abun- dance and harvest rate. Although moose should benefit from salvage logging through increased forage production (Janz 2006), those benefits are not immediate and may be offset by higher harvest and predation due to easier access afforded by high density of roads and cutblocks (Ritchie 2008). To address the recent moose population declines, British Columbia initiated a provin- cially-coordinated research project in 2013 to evaluate the landscape change hypothesis (Kuzyk and Heard 2014) and to increase science-based information for moose man- agement. To date, unpublished data from this research has provided no evidence that low pregnancy rates, infectious disease, or parasites are influencing the moose popula- tion (H. Schwantje, BC FLNRO, personal communication). Similarly, preliminary adult survival rates are within the limits of a stable moose population (92 ± 8% in 2013–2014 and 92 ± 5% in 2014–2015; Kuzyk et al. 2015). In southeastern British Columbia (Re- gion 4), declining forage production in older burns and wolf predation are believed limit- ing to moose population growth (Stent 2009, 2012). Further, in an attempt to reduce predation of an endangered caribou popula- tion, the local moose density was reduced which lowered wolf abundance in a small Fig. 2. Provincial population estimates of moose and trend line derived from inventories, population modeling, and expert opinion from 1987 to 2014 in British Columbia, Canada. Minimum and maximum ranges in population estimates are presented from 2000 to 2014. ALCES VOL. 52, 2016 KUZYK – POPULATIONS AND HARVEST OF MOOSE IN BC 5 portion of the region (~6,375 km2) (Serrouya et al. 2011, Serrouya 2013). Given stakeholder and public concern for declining moose populations, it is import- ant to maintain a balanced, provincial-level assessment and approach that also addresses regions with stable or increasing populations. The large northwestern (Region 6) and north- eastern (Region 7B) regions with stable Fig. 4. Regional moose population estimates and declining trend lines in Regions 4, 5, and 7A as derived from inventories, population mod- eling, and expert opinion, 1987–2014, British Columbia, Canada. Minimum and maximum ranges in population estimates are presented for 2000–2014. Fig. 3. Regional moose population estimates and trend lines in Regions 3, 6, 7B, and 8 as derived from inventories, population modeling and expert opinion, 1987–2014, British Co- lumbia, Canada. Minimum and maximum ranges in population estimates are presented for 2000–2014. 6 POPULATIONS AND HARVEST OF MOOSE IN BC – KUZYK ALCES VOL. 52, 2016 moose populations are more remote than those in the southern half of the province and have not undergone landscape change that presumably facilitates hunter and preda- tor access. These regions also experienced little impact from the mountain pine beetle outbreak compared to the central interior regions (Region 5 and 7A). The one stable population in the south was largely affected by the mountain pine beetle and salvage logging, but had lower wolf density com- pared to northern regions (BC FLNRO 2014, Kuzyk and Hatter 2014). The increas- ing population in the southern region (Re- gion 8) overlapped with a recolonizing wolf population (BC FLNRO 2014). Further, this regional estimate was revised in 2013 with a habitat-based model (Gyug 2013) that may have amplified the estimated increase in abundance between 2011 and 2014. The average (6%) and range (4–8%) of the provincial licensed harvest rate were mid-range of values reported throughout North America (2–16%; Crête 1987). More conservative harvest rates of 5% are recommended for northern systems where predation is believed to limit moose density (e.g., Yukon; Hayes et al. 2003), and may be appropriate in northern regions of British Columbia (Hatter 1999). First Nations harvest of moose is thought to be broadly distributed province-wide (BC FLNRO 2015), but because no formal method exists to quantify First Nations harvest, the total harvest and rates reported here are under- estimated and conservative. For example, local harvest may have been underestimated by up to 40% in Ontario by not accounting for First Nations harvest (Leblanc et al. 2011). Harvest information from First Nations in British Columbia would benefit future man- agement efforts to ensure sustainable harvests for all users including First Nations, recre- ational hunters, and the guide-outfitting indus- try (BC FLNRO 2015). An important outcome from this assess- ment was documentation of the gradual de- cline in licensed harvest by approximately half over 28 years from 1987 to 2014, despite constant hunter effort, indicating that the kill Fig. 5. Annual estimates of provincial moose harvest and hunter effort (average days hunted) by licensed hunters, British Columbia, 1987–2014. ALCES VOL. 52, 2016 KUZYK – POPULATIONS AND HARVEST OF MOOSE IN BC 7 per unit of effort (kills/hunter days) had declined. The disparity between these two trends may be related to difficulties produ- cing accurate provincial population estimates that are driven by wide regional variation. Further, changes in the hunting season struc- ture in the early 1990s reduced harvest levels in some regions (Hatter 1999), and similarly, a regulatory change allowing shared limited entry hunts in the early 2000s raised hunter effort through increased opportunity to hunt moose, without increasing harvest. Finally, although hunters maintained constant hunt- ing effort as harvest declined, lower hunter success often reflects inclement weather and human disturbance that influence moose dis- tribution. Given the number, frequency, and variable proportional influence of these fac- tors, kill per unit of effort is probably not a reliable measurement to assess moose Table 1. A summary of annual moose license sales and annual estimates of licensed hunters, hunter days, and moose harvest in British Columbia, Canada, 1987–2014. Year Licensed hunters Licensed hunter days Licensed harvest License sales 1987 42,526 338,482 13,463 N/A 1988 42,679 334,246 13,539 N/A 1989 41,979 332,852 14,070 51,520 1990 42,104 334,718 13,457 50,367 1991 39,400 304,852 12,251 46,010 1992 38,973 314,613 11,557 45,289 1993 33,236 252,647 10,025 38,538 1994 31,423 247,039 9944 37,714 1995 31,778 248,281 11,047 38,018 1996 30,923 245,617 9701 35,948 1997 32,085 251,582 10,494 37,243 1998 35,617 276,206 11,438 41,089 1999 29,840 250,287 7459 35,612 2000 31,106 255,569 9182 36,221 2001 30,988 272,771 10,290 36,145 2002 31,829 256,975 10,803 37,010 2003 31,493 238,983 11,309 36,608 2004 27,293 214,743 9571 40,438 2005 31,498 253,619 9980 37,175 2006 32,010 247,409 9939 38,374 2007 31,719 260,126 8000 38,069 2008 31,368 267,654 8730 37,125 2009 32,880 291,920 8074 40,371 2010 32,242 270,781 8836 39,733 2011 32,324 280,931 7660 40,503 2012 32,277 276,699 7576 40,236 2013 32,420 280,133 6890 40,109 2014 30,172 261,677 5773 39,723 Mean 33,721 ± 4292 273,622 ± 32,521 10,038 ± 2137 39,815 ± 4158 8 POPULATIONS AND HARVEST OF MOOSE IN BC – KUZYK ALCES VOL. 52, 2016 abundance in British Columbia (Hatter 2001). 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