Deer populations up, hunter populations down ... - Shawn J. Riley

ECoSCIENCE

10 (4): 455-461 (2003)

Deer populations up, hunter populations down: Implications of interdependence of deer and hunter population dynamics on management1 Shawn J. RILEY2,

Department of Fisheries and Wildlife, Michigan State University, East Lansing, Michigan 48824, U.S.A., e-mail: [email protected]

Daniel J. DECKER, Jody W. ENCK, Paul D. CURTIS, T. Bruce LAUBER & Tommy L. BROWN, Human Dimensions Research Unit, Department of Natural Resources, Cornell University, Ithaca, New York 14853, U.S.A. Abstract: White-tailed deer (Odocoileus virginianus) are managed to yield diverse impacts, including effects to ecosystems. Many conventional hunting systems manage deer abundance through rules that strive to produce recreation opportunities and an equitable distribution of antlered bucks among hunters. To protect against excessive harvests, antlerless deer harvests often are regulated through quotas. This approach is effective when deer productivity does not outstrip capacity of the hunter population to harvest required numbers of antlerless deer. In many areas of North America, abundance of white-tailed deer has increased dramatically in the past two decades, which has caused many wildlife managers to ask whether deer populations can be controlled with conventional harvest strategies. We used population reconstruction modeling to simulate deer populations from mixed hardwood forests in southern New York, determined antlerless deer harvests needed to stabilize or reduce populations, and evaluated whether current hunting systems can effectively achieve potential ecosystem objectives. Current hunter willingness to seek or use antlerless deer permits likely is inadequate to stabilize or reduce deer densities. This situation may be exacerbated in the future with occurrence of diseases in deer or other factors that diminish hunter participation. We discuss implications for effectiveness of ecosystem management. Keywords: harvest, hunter, New York, Odocoileus virginianus, population, white-tailed deer, wildlife management. Résumé : Les populations de cerfs de Virginie (Odocoileus virginianus) sont gérées de façon à réduire les impacts négatifs associés à la prolifération de cet animal, notamment le broutement excessif dans certains écosystèmes. En général, l’abondance des cerfs est contrôlée par une chasse récréative qui assure une répartition équitable des mâles avec bois entre les chasseurs. Pour éviter des récoltes excessives, la chasse aux cerfs sans bois est souvent réglementée par des quotas. Cette approche fonctionne lorsque la productivité des cerfs ne dépasse pas un certain niveau et que les chasseurs sont en mesure de récolter le nombre désiré de cerfs sans bois. Toutefois, dans plusieurs régions de l’Amérique du Nord, l’abondance du cerf de Virginie s’est accrue de façon telle au cours des deux dernières décennies que plusieurs gestionnaires de la faune se demandent si les populations peuvent être contrôlées par les stratégies de récolte habituelles. À l’aide de la modélisation, nous avons déterminé quelle doit être la récolte de cerfs sans bois pour stabiliser ou réduire les populations de cerfs. Nous avons également évalué si les programmes actuels de chasse répondent aux objectifs de protection des écosystèmes. Le modèle s’applique aux populations de cerfs des forêts feuillues du sud de l’état de New York. Dans le système de chasse actuel, la bonne volonté des chasseurs pour prélever des cerfs sans bois n’est pas suffisante pour stabiliser ou réduire les densités de cerfs. Ce problème risque de s'aggraver avec l’apparition de maladies chez les cerfs ou d’autres facteurs pouvant diminuer la participation des chasseurs. Nous terminons cet article par une discussion sur l’efficacité de la gestion des écosystèmes. Mots-clés : cerf de Virginie, chasseur, gestion de la faune, New York, Odocoileus virginianus, population, récolte. Nomenclature: Corbet, 1984.

Introduction Abundant white-tailed deer populations represent one of the greatest challenges in natural resource management early in the 21st century. In much of North America, numbers of white-tailed deer (Odocoileus virginianus) and numbers of deer hunters, the principal tool for management of deer populations (Carpenter, 2000), are changing in opposite directions. White-tailed deer populations 1Rec.

2002-09-12; acc. 2003-05-09. for correspondence.

2Author

recently exceeded record levels (Warren, 1997), while hunter numbers declined. A declining population of big game hunters is occurring in the U.S.A. (Enck, Decker & Brown, 2000). The decline of big game hunters has been less severe in Canada, although a decline is still evident (DuWors et al., 1999). An aging hunter population and low recruitment of new hunters suggest further declines in the hunter population can be anticipated (Enck, Decker & Brown 2000). These divergent trends may have serious implications for society and wildlife managers.

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Deer are implicated as a causal factor in prevention of forest regeneration (Tilghman, 1989), reduced songbird numbers (deCalesta, 1994), decreased aesthetic attributes of ecosystems (Underwood & Porter, 1991), increased damage to agricultural crops (Conover, 1997), increased frequency of motor vehicle and aviation crashes (Romin & Bissonette, 1996; Dolbeer, Wright & Cleary, 2000), and perpetuation of disease (Ostfield, Jones, & Wolff, 1996). Complex relationships exist between deer density and impacts to ecosystems from browsing. However, deer densities > 8 deer·km-2 are thought to impede sustainability of ecosystem form and function in northern hardwood forests (deCalesta & Stout, 1997). Mortality from hunting of antlerless deer has been a conventional means to manage abundance of white-tailed deer (Woolf & Roseberry, 1998). Regulated hunting, historically rooted in a time when deer were scarce (McCabe & McCabe, 1997), protected deer populations from excessive harvest because growth rates of a deer population primarily are dependent on the female mortality rates (i.e., harvest). In a closely regulated hunting system, if hunters are abundant and deer are scarce, there is little cause for concern about excessive harvest, and antlerless licence/permit systems work well to distribute hunters and minimally affect deer populations (Denney, 1978). However, if deer are abundant and hunters are scarce, the ability to control populations through recreational harvest may be reduced (Brown et al., 2000). Declining numbers of hunters, limited hunter access to deer on private lands and developed areas, and insufficient willingness among hunters to kill antlerless deer are factors prompting agencies to carefully examine the interdependence of deer and hunter population dynamics (McShea, Underwood & Rappole, 1997; Wright, Kaiser & Emerald, 2001). An assumption in most conventional deer harvest strategies is that adequate demand for and successful use of antlerless deer permits exists to achieve desired deer harvest. This assumption warrants careful examination. The issue of matching hunter numbers and willingness to kill antlerless animals is not confined to white-tailed deer in North America. States such as Colorado and Montana are experiencing a new phenomenon of not being able to achieve adequate antlerless elk (Cervus elaphus) harvest to control populations (R. Kahn, pers. comm.; K. L. Hamlin, pers. comm.). In a pilot study, Brown et al. (2000) asserted that harvests needed to stabilize deer populations under existing hunting regulations in New York exceeded reported antlerless harvests. We report findings from more thorough analyses in the Northern Glaciated Allegheny Plateau (NGAP), a dominant type of forested ecosystem comparable to many other hardwood types in northeastern North America. The question we address is, can adequate harvests of antlerless deer be achieved when numbers of deer are up and numbers of deer hunters are down?

(Severinghaus & Moen, 1983), our 25,519-km2 study area encompassed 13 counties where NGAP comprised more than 80% of the land classification type (Figure 1). Counties were used as base units of analysis because deer harvest data are most accurately collected and reported by New York’s Wildlife Management Agency at this scale (Kautz, 1995). Study area boundaries included Lake Erie on the west, Pennsylvania (a continuation of NGAP) to the south, gentler terrain and open, agricultural habitats of the Erie and Ontario Lake Plain ecoregion to the north, and the Catskill Plateau to the east. The NGAP, characterized by numerous moraines, drumlins, kettles, and other glacial features, is a subsection of the Laurentian Mixed Forest (McNab & Avers, 1994). Predominant land features are moderately dissected plateaus, broadly rolling hills, and narrow valleys. Elevation ranges from 200 m to 610 m. Nearly 70% of the area is in second- or third-generation forested uplands (Alerich & Drake, 1995). Dominant forest types include oak-hickory, mesic beech-maple, oak-pine, and hemlocknorthern hardwoods. Active and idle farmland covers much of the landscape on lower slopes and valleys. Cities and towns are scattered adjacent to major waterways associated with the Susquehanna, Delaware, and Allegheny watersheds. To assess what might happen under status quo management policies and to determine whether deer populations could be reduced with hunter harvests, we compared estimates of needed antlerless deer harvest, derived by modelling deer population dynamics, with reported antlerless and potential antlerless deer harvests. Needed antlerless deer harvest was defined as harvest required to reduce total post-hunt deer population density to ≤ 8 deer·km-2 in the 13 NGAP counties within 5 y. This density objective was selected because previous studies in northern hardwoods (deCalesta, 1994) suggested impacts from herbivory at deer densities > 8 deer·km-2 prevented regeneration of desirable trees such as sugar maple and diminished other ecosystem functions. A 5-y time span was chosen because it is a common time specified to reach objectives in wildlife planning efforts and it often is used by citizen

Adirondack Mountains Lake Ontario

Lake Erie

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Methods STUDY AREA The NGAP ecoregion covers portions of 23 counties in southern New York. To include only counties with similar deer population dynamics as related to habitat

Syracuse

Buffalo

78o55'W

42o40'N N

40o44'N

73o47'W

FIGURE 1. Map of the 13-county (25,519 km2) study area in southern New York used for assessing effectiveness of hunting to control white-tailed deer populations.

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task forces when setting management objectives in New York (Stout et al., 1996). Reported antlerless deer harvest was the number of antlerless deer killed during the 1997-1998 deer season as determined by a hunter survey (Curtis et al., 2000). Potential antlerless deer harvest was defined as the highest total number of antlerless deer hunters reportedly would kill in a hunting season if they could obtain an unlimited number of deer management permits (DMPs). HUNTER BEHAVIOUR New York’s deer management system, at the time of this analysis, allowed hunters to purchase a big-game licence valid for one antlered deer. Antlerless deer harvest was regulated on a deer management unit (DMU) basis by quotas of DMPs that could be obtained prior to hunting season through a lottery-type drawing. Hunters could apply for up to two DMPs in most DMUs or for a single DMP in each of two different DMUs. Each DMP entitled a hunter to harvest one antlerless deer, or a deer with antlers < 10 cm in length. Application for these permits was voluntary and DMPs were free, but there was onetime US$2 application fee that had to be mailed with the application to the wildlife agency by a certain deadline. We determined reported and potential antlerless deer harvest from a mail survey of 5,323 deer hunters, stratified proportionately by county of sale, drawn from the 513,310 persons who bought a big-game licence in New York State for the 1997-1998 hunting season. We began the survey on 5 January 1999 using a conventional fourwave mailing procedure (Dillman, 1978). Nonresponse bias was assessed via telephone interviews with 50 nonrespondents to the mail survey. We adjusted the survey results to account for nonresponse bias only for statewide variables. Because we based reported and potential harvests on respondents who actually went hunting, the number of active NGAP hunters was estimated by multiplying the total number of persons who bought a big-game licence in New York State in 1997 by the proportion of survey respondents (adjusted for nonresponse bias) who indicated they hunted during 1997 (89.5%). This product was multiplied by the proportion of respondents who hunted deer ≥ 1 d in the NGAP during 1997 to estimate Allegheny Plateau hunters (APHUNTR). Reported antlerless deer harvest was estimated by multiplying APHUNTR by the mean number of DMPs for which APHUNTR applied (APPLIED) and in turn multiplying this product by the mean per hunter permit fill rate (FILL) for NGAP hunters. These procedures likely overestimated harvest because hunters could apply for and fill up to two antlerless permits, but some of those permits may have been for other parts of New York (i.e., outside the NGAP area). Thus, we had estimates of APPLIED and FILL, but did not know whether hunters applied for or used all their permits in the NGAP. In the questionnaire, we asked how many deer in total (TOTDEER) hunters would want to harvest if they could harvest an unlimited quantity and what was the minimum number of those deer they would want to be antlered bucks (MINBUCKS). From these data, we estimated the maximum number of antlerless deer (MAXDOES) they

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would want to harvest by subtracting MINBUCKS from TOTDEER. We assumed this to be a reasonable estimate for New York hunters because other regulations, which otherwise might limit hunter access to bucks, were not in place at the time of the survey. To estimate potential antlerless deer harvest, we multiplied APHUNTR × MAXDOES × FILL. By using survey data to estimate both reported and potential harvest (rather than an agency estimate of reported harvest), we could compare reported and potential harvests using the same dataset. We were confident about comparing needed harvest (calculated from harvest data) to reported and potential harvest because our estimate of reported harvest was similar to agency estimates obtained through other methods. We determined a point estimate for reported harvest and low and high bounds on that estimate. For the two averaged variables (APPLIED, FILL) used in the equation for reported harvest, a mean value was calculated and bounded 2 SE below (low estimate) and above (high estimate) the mean. Three estimates for reported harvest were then calculated: (1) using the mean value for all variables in the equation, (2) using the low estimates for all variables, and (3) using the high estimates for all variables. Finally, we assessed whether agency estimates of reported harvest were within the high and low bounds of our estimate for reported harvest using survey data. Our models for needed and potential harvest include two assumptions important for result interpretation. The first assumption was that hunter bias against filling an antlerless permit with a fawn compared to an older female deer (0.75) represents hunter behaviour accurately. This assumption was based on observed hunter take under New York hunter regulations (Kautz, 1995). The second assumption was that hunters' rate of success in filling antlerless permits remains constant above two permits. The latter assumption likely results in a higher estimate of potential harvest than would be observed because success rates per permit probably decline as individual hunters obtain greater numbers of permits. Because actual experience in New York has been limited to a maximum of two DMPs, we had no data that could provide guidance on this relationship. In Michigan, where hunters could purchase one license per day for antlerless deer in most hunting units, 22% of hunters killed at least one deer, but only 14% killed two or more deer (Frawley, 2002). DEER POPULATIONS Deer population dynamics were calculated with a population reconstruction process (Downing, 1980) assisted by an updated version of Deer CAMP (Computer-Assisted Management Program) (Moen, Severinghaus & Moen, 1986). This version of Deer CAMP determines the population needed to support observed harvest levels and estimated crippling loss, illegal kill, road kill, predator kill, and summer and winter fawn mortality over a set period of years while reaching a specified ratio of population change between two given years. A change ratio of 1.064, which matched observed changes in numbers of antlered bucks harvested between 1991 and 1998, was used to provide the model with a target for population change. Agestructure data were derived from hunter checks in New 457

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York (New York Department of Environmental Conservation, unpubl. data). Relatively conservative values for mortality factors were used because no empirical estimates existed. Mortality factors included (1) crippling loss (10% of reported harvest), (2) illegal kill (2% of the pre-hunt population), and (3) road and predator kills not accounted for in fawn mortality (1% of population). Summer-fall fawn mortality (mean, 42%; range, 37-57%) was calculated each year in proportion to fawn:doe ratios in the harvest. Winter fawn mortality (mean, 4%; range, 0.5-15%) was calculated each year in proportion to the yearling fraction of antlered buck harvest the following fall. Values were conservative in that if wounding rates were higher, the summer-fall fawn mortality rates were lower, or the number of predator kills was higher, a greater total deer population was needed to achieve the observed hunter harvest. Lower mortality rates were used for yearling and prime-aged adult deer, proportional to an age-related vulnerability curve that reached a maximum at age 13. This non-random portion of summer and winter mortality accounted for physiological stress and included age-related predation. Rates were applied to reconstructed populations as dynamic calculations throughout the biological year rather than to a specific number of deer for the entire year. No attempt was made to estimate deer abundance in areas closed to hunting (e.g., suburban parks, preserves, etc.); therefore, density estimates should be considered minimum values. Two management scenarios were tested. The first scenario modelled what would occur if the status quo persisted (mean 1994-1998 harvest rates for antlered and antlerless deer). To reveal the harvest needed to achieve 8.0 deer·km-2, the second scenario held harvest rates for bucks constant (mean 1994-1998 rates) and varied harvest rates for antlerless deer until the management objective was achieved in 5 y. Harvests from the two scenarios were then compared with potential harvest to determine whether hunting could be used as an effective mechanism for population control of deer.

Results Model estimates of pre-hunt total deer in the study area between 1989 and 1998 ranged from 308,661 to 378,929 (Figure 2). These estimates represent densities of 12.1 and 14.8 deer·km-2. The model deer population at the start of the 1997 hunting season comprised 378,929 deer, of which 33.8% were fawns, 47.5% were does (≥ 1.5 y), and 18.7% were bucks (≥ 1.5 y) (Figure 3). Total reported harvest was 81,273 deer, which included 16,601 fawns, 22,314 does, and 42,358 bucks. Based on the reconstructed population, harvest rates for the 1997 hunting season were 13.0% for fawns, 12.4% for does, and 58.2% for bucks. If harvest rates for antlered and antlerless deer stayed at 1997 levels, our model predicted the 2002 pre-hunt deer population would increase to 407,000 deer, or about 15.9 deer·km -2 . To achieve a reduction in density to 8.0 deer·km-2, antlerless harvest rates would need to be about 26%, or the harvest level would have to start at 81,000 antlerless deer and end at about 46,000 antlerless deer. 458

400 350 300

Total deer Adult females Fawns Adult males

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19 89 19 90 19 91 19 92 19 93 19 94 19 95 19 96 19 97 19 98

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FIGURE 2. Estimates of the 1989-1998 pre-hunting season whitetailed deer population, based on population reconstruction, by sex and age class in a 13-county (25,519 km2) region of southern New York.

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140 ×103) Number of deer (×

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Male fawn

Female Yearling Adult Yearling Adult male female female male fawn Age class and gender

FIGURE 3. Estimates of 1997 pre-hunting season white-tailed deer population by gender and age class, from population reconstruction, and harvest, based on hunter surveys, in a 13-county (25,519 km2) region of southern New York.

We estimated that 204,390 people hunted ≥ 1 d in the study area during the 1997-1998 hunting season. These hunters applied for and received an average of 1.082 ± 0.058 DMPs (mean ± SE), and used an average of 20.2% ± 2.4% of the DMPs they received. Based on these findings, we estimated reported antlerless harvest to be 44,642 ± 7,387 deer. Bounds on our estimate (37,255-52,659) encompass the wildlife agency’s point estimate of 38,915 harvested antlerless deer developed through hunter mail-in report cards. Under a scenario of unlimited availability, hunters stated they would apply for an average of 1.13 ± 0.09 antlerless permits. The potential antlerless harvest that would result from this stated willingness was estimated to be 46,654 ± 8,817 deer, or 4.5% higher than the 1997-1998 reported harvest. This indicates a possible discrepancy of more than 34,000 antlerless deer between potential and needed harvest.

Discussion What does the dilemma of deer numbers up and hunter numbers down portend for wildlife management? In New York, a potentially severe problem has developed

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in the NGAP region. Ecosystem-oriented objectives have not been established in our NGAP study area, and the cascade effect of herbivory by deer remains uncertain. However, deer density in the NGAP currently is far in excess of levels needed to achieve regeneration of many favoured hardwood forest species (Alverson, Waller & Solheim, 1988; deCalesta & Stout, 1997; Healy, 1997), and cascade effects from these densities likely are changing composition of species in the ecosystem (deCalesta, 1994). The relevance of our observations likely is not restricted to southern New York, because hunter numbers are down across much of the range of white-tailed deer in eastern North America (Enck, Decker & Brown, 2000), while deer numbers are mostly up (Warren, 1997). By concentrating efforts, enough hunter harvest may be achieved to control deer on small scales such as individual management units, preserves, or refuges (Kilpatrick, Spohr & Chasko, 1997), but the ability to affect whitetailed deer populations on large scales (i.e., ecosystems) is diminishing (Brown et al., 2000). Our data indicate that hunter willingness to kill antlerless deer may not be great enough to control deer populations, a problem that is compounded by declining hunter numbers and diminished recruitment of new hunters (Enck, Decker & Brown, 2000). Even in states, such as Michigan, that offer essentially unlimited numbers of licences for antlerless deer, declines in hunter number and participation are observed (Frawley, 2002). This phenomenon may be exacerbated by several phenomena looming on the horizon, such as chronic wasting disease (Bartelt, Pardee & Thiede, 2003) and other factors that may increase perception of risk to hunters or other consumers of hunting or venison. For example, if consumption of venison becomes a public health concern associated with chronic wasting disease, programs like “Hunters for the Hungry”, which might normally encourage some hunters to kill more deer than they can personally use, may be less effective. These types of programs greatly decreased operations following the onset of chronic wasting disease in Wisconsin (Bartelt, Pardee & Thiede, 2003). Many factors affecting recruitment and retention of new hunters result from macro socioeconomic (e.g., economic, social and cultural) phenomena that are mostly outside the purview of traditional wildlife management activities (Dann & Peyton, 1996). Ongoing loss of experienced hunters will worsen the effects of declining hunter recruitment. Antecedents to recruitment and retention of hunters include social support, mentoring, and apprenticeship; however, wildlife agencies have little experience in influencing these factors to gain more hunters (Enck, Decker & Brown, 2000). Additional research is needed to understand these relationships, but such knowledge may have little practical importance for agency programs since it is unlikely any agency initiative would have sufficient effect to reverse the trends and make a meaningful difference for deer management (Decker, Enck & Brown, 1993). Citizen task forces in the NGAP study area, in cooperation with wildlife managers, have set objectives for reducing deer populations (Stout et al., 1996). Mechanisms to harvest more antlerless deer include multiple DMPs offered in most DMUs and new incentive programs to

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encourage landowners to harvest antlerless deer. In 2001, these efforts resulted in a total harvest of 115,513 deer, of which 47,238 were antlerless. This antlerless harvest was very similar (584 deer greater) to our projected antlerless deer harvest under a scenario of unlimited DMPs. Yet, this harvest represents only 58% of the estimated 1998 harvest needed to initiate a reduction of deer to 8·km-2. Because 4 y elapsed between 2001 and when our estimates were made, deer harvests now needed to reduce deer density probably are even greater. Without an appreciable gain in hunter numbers, participation, harvest effectiveness, or willingness to kill more antlerless deer, a greater investment is needed to develop alternatives to hunting as the sole population control mechanism. Access to hunting is limited or not feasible in many areas. In these situations, deer are not susceptible to conventional management regardless of whether hunters are present and willing to harvest antlerless deer, unless non-hunting techniques are used. These refuges likely replenish deer into hunted areas and magnify difficulties of achieving herd reduction (Brown et al., 2000). A decline in rural human populations likely will lead to fewer hunters recruited in the future (Stedman & Heberlein, 2001). Hunting eventually may become less a recreation and more a community service or civic duty should the impacts of deer be broadly recognized. Culling may be a more appropriate term for the kind and purpose of hunting under such circumstances. This situation currently is being tested in Wisconsin, where preliminary results from that state’s attempts to eradicate all whitetailed deer within a region infected with chronic wasting disease suggest hunter participation declines as deer populations are reduced (T. Van Deelen, pers. comm.; Van Deelen & Etter, 2003). No easy solutions exist, although several conclusions are apparent from our experience in New York. We believe a paradigm shift, already underway in some states, is needed in public white-tailed deer management. The shift needed is from one of protection and distribution of a scarce resource to one of managing impacts of deer (Riley et al., 2002). More focus on education and engagement of non-hunting stakeholders is needed to ensure that decisions about hunting and population control arise from community deliberation and not merely from agencies. To be effective, any population-control mechanism will require acceptance by society (Curtis et al., 1997). Professionals from many fields concerned with ecosystem management will need to get involved in deer management. No other area of wildlife management will require more integrative thinking than that involved with crafting solutions to the dilemma of rising deer numbers and declining hunter numbers. Solving the dilemma likely will require bringing together an array of disciplines such as sociology, social psychology, economics, education, city and regional planning, communication, and wildlife ecology. Acknowledgements E. Kautz provided deer harvest data from New York State Department of Environmental Conservation (NYSDEC) records. K. Sullivan compiled and entered harvest data, and A. Moen 459

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conducted reconstruction modeling of white-tailed deer populations. Members of the NYSDEC deer management team contributed to the early discussions that were the genesis of this paper, and team members provided valuable critiques of methods and interpretation of results. Staff in the Human Dimensions Research Unit at Cornell University provided support in administration of hunter surveys. W. Moritz and two anonymous reviewers improved the manuscript with their suggestions. Preparation of this paper was supported in part by the Cornell University Agricultural Experiment Station through Hatch Project NYC 147-403 and by the Michigan State University Agricultural Experiment Station through state project MICL02031.

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