Population characteristics of' a ten-trall Appalachian whi-tetailed deer h.erd
Abstract Reliable estimates o i white-tailed deer (Odocoileus virginianusi population parameters
are needed for effective population management. W e used radiotelemetrv to compare survival and cause-specific rnortality rates between male and iemale white-tailed deer and present reproductive data for a high-density deer herd i n the central Appalachians of West Virginia during February 1999-May 2002. W e recorded 343 winter deer captures. Our capture rate of 78 fawns/100 adult females was similar to visual observations of known fawns and adult iemale deer during winter. In-utero reproductive rates of adults 121 year old) was 138 fawns:100 females, based on a sample of 44 female deer. We radiomonitored 138 iemale and 43 male deer during the 3-year studv. Yearling male annual rnortality rates ior human-induced and natural mortality were 0.63 (SD=0.09) and 0.12 (SD=0.12), respectivel!,. Conversel); among yearling females, annual mortality rates for h u m a n - i n d ~ c c dand natural nortality were 0.09 (St?= 0.06) and G.G5 (SD= 9.031, respectively. Adult male annual mortality rates for human-induced and natural mortality were 0.73 (SD=0.16) and 0.00, respectively, whereas adult female annual mortality rates for human-induced and natural rnortality were 0.04 (SD=O.01) and 0.08 (SD=O.O?), respectively O u r observcld survival, mortality, rapture! and reproductive data are characteristic of a population shaped by moderate fawn recruitment and excessive harvest ot yearling male deer. W e recommend a reduction in female deer to promote torest regeneration and protect biodiversity in this region. W e believe this can best he achieved by liberalizing antlerless harvest regulations, through hunter education and corporate landowner incentive programs.
Key words Appalachians, mortality, Odocoileus virginianus, recruitment, reproduction, survival,
white-tailed deer
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Current annual forest removals in =St Virginia are >12;000,000 rnj of sawtimber and pulpwood (IJnited States Department of Agriculture Forest Service 3004), a level which approximates that of the large, widespread harvests at the turn of the twentieth century (Stephenson 199317). Forest management and wood production are 2 vnluable mainsray of the state's and region's cconomy (Spencer 19', r\dams et 21. 2000). Likewise. this
portion of the central Appalachians. particularly the eastern one-third of the state, is regarded as an important biodiversity hotspot within the Appalachians and among all temperate forested Chaplin et a1.2000. ecosystems (Stephenson 1 9 9 3 ~ . Roberston and Hull 3003. Ford er 31.2005). From a natural resource management perspective. the conservation of biodiversiw m d managrment o f economically valuable and ecologically sustainable
forests can be complement;~r!. activities within the Appalachians (Fredericksen 1998 1. 1-1c~n7e\-er. sustaining these outputs either singl!- or ! in concerr is complicated b. forest system stressors such as atmospheric acid deposition (Adams 1999j. exotic insect and disease pathogens (Stipes 199'). Souto and Shields 2000, Davidson et al. 2001). ancl white-tailed deer (.Orfocoilezrs oirgininizrrs) herbivury (deCalesta 1994,Fredericksen 1998,Horsle)er al. 200.3) that are occurring in the central Appalachirms. For wildlife biologists. whitetailed deer herbivory jmpacts to ecosystem,processesand forest management activities . are o f ,paramount . -. . interest and concern in this region. After the nea; extirpation .of white-tailed deer populations from West Virginia during the early 1900s, populations increased due to buck-only hunting regulations, increases in favorable habitat, and restocking efforts (Allen and Cromer 1977). Throughout the last half of the twentieth century. West Virginia had liberal buck harvests with comparatively conservative doe harvests. Populations have increased in some locations to a point where forest damage is apparent (Michael 198-, 1992 ) . On corporate forest lands. altered or failed wood?regeneration is common (Tilghman 1989, Horsley et al. 2003). To promote healthier ecosystems and balanced deer populations, harvest regulations have been altered t o increase the harvest of female deer. However, in comparison with other southeastern states, antlerless harvest in West Virginia is low (iT% of total, Annual Meeting of the Southeast Deer Study Group 2003) and likely insufficient to stabilize deer herds in the near future (J. L. Crum. West Virginia Division of Natural Resources, personal communication). In West Virginia this is due partly to decreasing resident and nonresident hunter recruitment, coupled with a stable to decreasing human population. No sex-. age-, or cause-specific estimates of sunrival and mortality rates exist for white-tailed deer in West Virginia. hindering biologists' cfforts. Reliable estimates of survi.ia1. mortalit!: and reproductive rates are requisite to understanding :~nd managing deer populations c Caughley 19-0, Eberhardt 1985. Dusel; rt al. 1989). Radiotelemetry prov~des in-depth evaluation of tllr timing, causan t.s. and degree of mortalin. (Heisey and Fulier 1985. Nelson 2nd Mech 1986). Hercin. we compared sur1.ival m t l cause-speclf'ic mortality rates between male .md female white-tailed $leer using iadiutclrmcrry ;ind present reproducrive :md
recruitment datn for a higll-densit!- clerr herd that i~ in conflicr with land rnanapmenr in the centrai Appalachians of &est Virginia.
Study area
O ~ i study was conducted on the ?vleaci\);.estv:lc r Corporation's Wildlife and Ecosystem Research Forest (MWWERF) in Kandolph County, West n;hich was established Virginia (ZW-i2'N. 8O0?'iV], in 1994 ;to assess the influence of industrial silvicultuyr on ecological and ecosystem processes in the central Appalachians. lie 3,360-ha ICIVOVERF occurrkci -in the Unglaciated Allegheny blountain and ~ l & e k Physipgraphic province within the c&tral ~ ~ ~ a i a c h j awhere precipitation averaged ns; from 170-190 cm/vear and elevations ranged from 7401,200 rn (Smith 1995). Forest cover was primarily an Allegheny hardwood-northern hardwood type (Ford and Rodrigue 2001). Silviculturally,the WMhIERF was intensely managed, with even-aged regeneration methods dominating. Preharvest white-tailed deer densities during our study- were estimated as 12-20 clrcr/iin2 iLanpdon 2001). Abomasal parasite counts suggested the deer population was at nutritional carrying capacity (Fischer 1996). The iMWWERF was open to hunting under West Virginia Divisicn of Natura! Resources harvest regulations. with access via foot-travel. Recent and ongoing research have determined that white-tailed deer were serving as a primary ecological agent shaping plant species diversity abundance, and structure on the iMWWERF (B. Collins. Savannah River Ecology Laboratory,personal communication, Collins 2004). Indirect impacts to biota. such as ground-nesting songbirds, have been noted (Weakland 2000).
Methods
We captured deer from Januan through April (1999-2001j using modified Clover traps (Clover 19i.i) and rocker nets (Hawkins et al. 1968) baited with corn. We immobilized. ear-tagged. and radiocollared (Advanced Telrmetry Systems. Isanti. blinn.) deer upon capture. We used both physical restraint and chemical (2.7 mg ~ ~ l a z i liydrochlone ride/kg body weight) immobilization teclmiquea. For chemicall?- immobilized deer. we used ' 2 intravenous and ' 2 intramuscular injections o f yohimbine h!-drochloride (0.5 mg/kg body weight) as n re~~ersal agenf. Radiocollars wrre equipped w ~ t h 3-
IIOLI?- nlort:llit\ sensor> \%,k aged clrcr b. tooth ! eruptio~:. replacemenr. and \vear (brveringhaus 1949). XI1 capture antl Iia~ictling procedures mere approved b ! the I - i t o f Georgia' Instirutiorial Aninlal (Arc and IIsc Coninlittee (Permii No.X2002-101 19-0). We monitorecl each deer 2 3 times/--cek from its release. until it died. the srud!- terminated (1 J u n e 2002). o r it was censored. \Vc censored deer w h e n the! dropped their collars or the collar batcen expired. W i e n mortal~ty signals were dctecred, w e located t h e carcass via homing and conducted a necrops). to determine cause. Predation was identified following procedures in Wade and Bowns (1984). We determined starvation from visual inspections of femur marrow (Verrne and Ullre>. 1984). Disease was inferred following Davidson and Nettles (1997). Collars from harvested animals commonly were left at the property gates with a note, turned in to the field station, left with the gut pile, o r taken home with hunters. We recorded all visual observations of deer during vehicular travel on the A W E noting age M Wm (fawn. adult [21 year old], or unknown). sex (male. fernair. o r unknownj, group size, and tag numbers when possible. We calculated monthly ratio of hwns:100 adult females for known age and sex observations only We determined age-specific inurero reproductive rates (feiuses/dot.) from 3 does necropsied in 2000.2 does necropsied in 2001. and 39 does necropsied in 2002. Median and mode conception dates were both 16 November (range= 4 November-25 December). We acquired and examined all female reproductive tracts from January-March. We determined sunrival and cause-specfic mortality rates from telemetn data for fawns, y e a r h g s , and adults (22 years old) with lMICROMORT (Heisey and Fuller 1985). For all analyses w e assumed n 1 June birthday of each cohort. We divided the vear into 3 seasons: summer (1 June-30 September).autumn (1 October-3 1 December), and winter il January-31 .May). We grouped mortality sources as either natural (stanration, predation. o r disease) o r human-induced (legal harvest. mounding loss. illegal harvest, or vehic~~lar trauma). Our analyses considered the exposure days of censored deer tliro~yh the cia!- of censoring Orangilder and Sherlff 1990'). We did not include deer that died of natural causes 131 days ;&er caprure in the analyses. i o void inclusion of deer wlnerable to predation due to capture rnyopath~ (Beringer et al. 19% ).
\k poolec! c l : m from dl .4 >-earst o increase sanlplc sizes oi' mnies, tllereb! fxcilitating rnc;uningfui between-sex comparisons ( m I k e l e n e t al. lqc)-) L 7 0 validatr pooling of data from the 3 years. wc. used Ztailed %-tests to compare mortalin- rate5 u-itl~in females among > w r s . Sinlilarl!-. wr used 2tailed Z-tests to compnrr mortality rates among ages. between sexes. antl among s e a s o n (Heise!and Fuller 1985) on pooled data. To maintain experiment-wise error rate of a = 0 . 0 5 . wc compared related tests following Ronkrroni correction (Sokal and Rohlf 1995). Wt-estimated survival of fawns from birth to capture (i.e.. recruitment ape) using change-in-ratio (CIR) estimators (Paulik and Kobson 1969). We defined a deer capture as any capture of an indiviclual deer not previously captured within t h e current trapping vear (i.e.. recaptures within a trapping year were not included). Time 1 ratios were estimated from capture data. survival estimates, and inutero reproductive estimates. Specifically. w e estimated the total number of fawns born to female deer by 1) adding the estimated number of deaths (via yearling and adult survival rates from June-December) to the age-specific number captured (this yielded an estimated number of yearling and adult females alive at birth), 2) multiplying the age-specific number of females alive at birth and csiimatccl age-specific in-il~eroi c p r ~ d u c t i v trates (this yielded an age-specific estimate of fawns born'). and 3) summing the age-specific estimates of fawns born across ages. We obtainedTime 2 ratios directly from capture data and validated them with observational data.
Results
We recorded 3;l-3 captures of white-tailed deer. Fawns were the dominant age class. representing 4 1 . X ( J Z = 143) of a 1 captures. Females were the 1 dominant sex class, comprising '5.8% (72 = 260) of ail captures. Adult male deer were notably scarce. representing only 1.5% ( 7 7 = 5 ) of all captures (Figure 1). The ratio of fawns:100 adult ( 2 1 year old) females at capture was ' 8 . Overall in-utero reproductive rates of yearling and adult females were 138 fawns: 100 females ( 5 1 Pawns per 3- yearling and :~clult females). Ikproducrive rates wrre greater for adults rhan no ! ye;urlings. \Ve obsen~ed e\-idencc o f I,reeding b. female fawns ( I Z = - ) , m d 1 adult had 5 fetuses in(item.
Deer.popuiation characteristics
Campbell et al.
215
From c a p t i n d a t a . age-specific rcproductivc rates. and estimates o f female survival probabilitie> (see below). xvc estimated fawns and breeding females alive a: Timr 1 as 17-1 and 199. respectivel!-. Fawns and b r e e d i n g females at Time 2 werr I t i and 183. respectivel!.. From these data. w e estimated the probability of fawns surviving from birth Females Maies to capture as 0.57. This Individual captures estimate suggested that fawns experienced greater Figure 1. Population protile irom white-tailed deer captures (Jan-Apr, n = 3 4 3 i on the mortality from J u n e MeadWestvaco Wildlife anc! Ecosystem Research Forest, West Virginia iron1 1999-2002. Slashed areas plus respective shaded areas are the estimated nurnher of fawns horn to iernale December than from Janudeer captured (from in utero reprocluctive rates, assuming a 50:50sex ratio st b ~ r t l i . ar>,-May(0.15 and 0.20 for female and male fawns. respective1y;Table 1). We radiomonitored 148 female and 43 male deer W e made 8:497 visual observations of known fsvns and adu!t female deer Ratios of fmns:100 from 1 June 15199 through 31 May 3003. adult females were low in June, increased during Throughout the study 16 females (14 adulrs, 2 yearwinter. and then declined in April and May (Figure lings, 0 fawns) and 11 males (0 adults, ? yearlings. 2). During January-April (i.e., during captures), the 3 fawns) were censored. VC;e observed 66 m o r r d l i ~ ratio of fawns:lOO adult females observed did not events, 35 within females (21 adults, 6 yearlings. differ from the ratio obtained from capture data 6 fawns) and 31 within males (5 adult. 19 yearlings, (Figure 2 ) . thereby supporting use of capture ratios 7 fawns). Predation was the primary mortality source for females. Coyotes (Canis lntmns) killed in CIR analyses. 10 femaies, black bears (Ursus nmericmzzls) killed 3 females. and unidentified predators killed 3 females. Three females died of unknown natural causes. 2 of starvation, and 2 of disease. Human-induced morralip accounted for 12 female deaths. Within this category, 8 deer died from legal harvest. 3 from wounding loss, and 1 deer was harvested illegally. Legal harvest was the prlmar!- source of rnorralin7 within males, accounting for 23 of 31 deaths. One male died from wounding loss. Natural mortalit!. events accounted lor S
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male deaths; 4 males died from covote predation. 2 from unidentified predators, 1 from starvation, and 1 from bobcat (Felis rzlfus) predation. We observed no vehicular trauma mortality Natural (0.07-0.17) and human-induced (0.05-0.07) mortality rates within females did not differ among years (all P>O.j5), suggesting minimal weather o r harvest-regulation effects and validating our pooling of data. Survival rates peaked during summer and were minimal during autumn for all
Table 2. Compar~son i wh~te-tailed o deer annual cause-speciiic rnortalirv rates tor sex and age classes o n the MeadWestvaco Wildlite ,ind Ecosvstem Research Forest West \ / i r g ~ n ~ Iron1 a 1999-2002. Standard deviations are reported in parentheses. Class ~Llales Females
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sex and age classes ( T ~ b l e >-exling females had 1). a 6i'% greater probabilih of annual survival than yearling males (Table 1). Similarly,adult females had a 66'!<, greater probability of annual survival than adult males (Table 1). Annual human-induced mortality was greater for yearling and adult males than females (Table 2). Annual natural mortality was greater in adult females than adult males but did not differ w i t h yearlings (Table 2). During winter mortality rates were similar between males and females within both fawns (Z= 0.51. P=O.61) and yearlings ( Z = l . i 8 , P=0.13). However. adult females had greater (Z=-?.O(>. P= 0.003) mortality rates in winter than adult males. Morrality rates during autumn were greater for yearling i Z = i . 9 7 . P*:: ... .+ .-.,
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inciuced (%=O.58.IJ=0.56)and natural (%= -1.88.1' = 0 . 0 6 )mortalic-. We observed no differences in con~parisons between summer mortality rates of adult and yearlil~g female> (Z= -0.7(3,I-'= 0 . 4 5 ) .sunlnler mortalinrates of adult and yearling males (Z=0.00.P= 1.00), autumn nlortality rates of a d u l ~ and !-earling . females (%= -0.53. ~ = 0 . 6 0 )and autumn mortalin rates of adult and yearling males (Z=0.30,P=0.70). Within females. no differences in mortality rates occurred during winter between adults and yearlings (%=().-2, P=O.-i7j. between adults and Pawns (%= -1.77,P=0.08).o r between yearlings and fawns (Z=-2.01, P=O.OLij. Within males. we observed no differences in mortality rates during winter between adults and yearlings ( Z = - 1.66,P=0.10) or between yearlings and fawns (Z= 0.37, P= 0.71). However, male fawns had greater ( Z = 2.50, P= 0.0 1) mortality rates than adult males during winter.
Discussion
Ricca et al. (3002) compared annual survival rates among white-railed deer populations throughout North America. Within hunted populations, adult e l vear old) male survival rates were greatest in southern Texas (0.65-0.74, DeYoung 1989) and irast ia northern Michigali (G.22-0.35,'v'an Deelci; et al. 1997). Our observed adult male annual survival rates (0.32-0.23) represent the lowest range among those reported. Adult 01 year old) female survival rates within hunted populations were greatest in northeastern Minnesota (0.79-0.80. Nelson and Mech 1986) and northern Michigan ( 0 . 7 - 0 . 8 9 , Van Deelen e t al. 199-1 and least in Montana (0.43-0.83. Dusek e t al. 1989). Similarly, our observed adult female annual survival rates (0.86-0.88) represent the highest range among those reported and were similar to or exceeded unhunted populations m c c a e t al. 1002). Estimates of neonate and young fawn survival are scarce. particularly within the eastern United States. Recent15 however, Vreeland (3002) reported that fawn sun7iral from birth to recruitment age cposthunting season) in Pennsylvania ranged from 0.28-0.59. depending on the study area and year. l Our estimate o f fan711 s u n ~ i r afrom birth to capture 10.5') was consistent with Vreeland (3002)and. similarly we suspect that coyotes. black bears. and bohcats preved upon neonate and young fan7nsduring the summer months because these predators 21lso killed older deer during the months that tbllowed.
Thc primar!- hitman-induced mortalin frlctoiinfluencing white-tailed deer population ecolog!- in north-central West Virginia is thr clifferential harvesi betxeen sexes (Table 1).This fincling is not unique to this region (Van Deelen ei a!. 199'). Langclon (2001) reported pre-hunting season adult ( L l year old) sex ratios (ma1e:female) for spotlight and camer:) surveys on tht. RI\XW/ERf as -: 100 and 26:100, respectively Others have attributed skewed sex ratios directly to male-biased hunting (Nelson and Mech 1981. Dusek et al. 1989, Nixon et al. 1991 ). Though infrequentlj- observecl. natural mortalin. was primarily from predation. Coyotes accounted for 6190 (14 of 23) of all known natural mortality predation events and 78% (14 of 18) of all k ~ l o w n events, including 7 adults. As in New Brunswick (Whitlaw et al. 1998). coyotes were the primary predator of adult deer. Traditionally, coyotes have been considered opportunistic predators (Bekoff 1977, Van Vuren and Thompson 1982). However. recent work in southern Alberta suggests that coyotes regularly participate in group hunts, specifically for deer when alternate prey is not available (Lingle 3000,. Our observations from mortaiiy sites suggest that multiple coyotes participated in deer kills. Black bear predation occurred in summer or early autumn and was not limited to fawns. Summer n ; ~ r t f , ai'k as notably low f ~ each -mL w~ r and sex class but similar to summer mortality for adult deer in New Brunswick (Wl~itlaw al. 1998). et In agricultural regions of Illinois. Nixon et al. (1 991) found the highest summer mortality rates within vearling females (0.15);however. all summer mortalities within their study were human-induced. Van Deelen et al. (1997) reported the highest summer mortality rates within yearling males (0.16), with half of the mortalities from stanration and half from unknown causes. W observed onl~7 summer mor2 talities. both from natural causes and within females (Table 1). Low natural mortality rates within vearlings and adults during summer suggest adequate nutrition for maintenance. mhanced hiding cover. and the possibility that predators were using alternate prey (Lingle 2000). Our estimate of fawn survival from June-December suggests these alternate prey likely included neonate ;ind young fawns. L4utumn mortality ot both ?-earlings and adults reflect selection by hunters for male ciecr. The greatest seasonal mortality rates for 31 age and sex 1 classes were observed for adult males during autumn. which is a finding not unique to our stud!area (Nixon et al. 1991). Winter ~nortalitvwithin
r ) ~ * p
km-ns was less than observed in northern hlichiyan p a n I>cele~i er a1. 199') and nort11e:tstern Mnnesot:~(Nelson and Mech 1986). but grealer than e:~s~-central Illinoi5 (Nixon er al. 1991). \-an Ikelen ct 31. (199') obsenFeclwinter mortalit!. rate? of 0.20 and 0.00 for adult and ).earling males. respecti\.el!.. This relationship was reversed in our stud! Hoa-rver. onl\- 1 adult male survived into the xvintrr. thus limiting meaningful comparisons. Our observed winter mortality ratc for yearling males ( 0 . 2 7 was less than reported from east-central Illinois (Nixon et al. 1991). However. 7 of 3 reported winter mortalities within yearling males from Nixon et al. (1991) were human-induced, whereas all of ours were from natural causes. High natural winter mortality within yearling males may suggest nutritional deficits and corresponding increase in vulnerability to predators (Nelson and Mech 1986). Our data illustrate the limitations of in-utero reproductive measures (counts of corpora lutea early in sgestation o r fetuses later in gestation) as indicators of population health. ~McCullough (19'9) demonstrated that n-hile both reprodrictiv~ and recruitment rates are density dependent, the)follow differing curves. Therefore, it ma!- be inappropriate to manage deer populations for maxiliiulli reprodactive ou:put because t k associated increases in fawn mortality may result in decreases in net recruitment rates. Because winter is the most nutritionally stressful season for deer in the central Appalachians and reproductive rates :Ire indicators of health prior to the breeding season p e n t w o r t h et al. 1990, Gee et al. 1994, Kroll and Jacobson 1995). we do not recommend in-utero reproductive measurements as the sole indicator of population health in this region. Without estimates of recruitment (a function of reproductive and survival rates) and other population measures, reproductive rates do not provide reliable management information. Many wildlife managers use visual observations collected incidentally or methodically (e.g.. from apotlight count data) to generate estimates o f recruitment or the "fawn crop" (Gee ct al. 1994. Qoll and Jacobson 1995) without considering biases .~ssociatedwith the temporal detectability or behavior of deer t McCullough ct al. I99it). Using spotlight counts. .McC~~llough (1982) found biases in the fan7n:adult female ratio in every month cxcept April. with the ratio being underestimated rroni June-March. Sirnllarly Langdon ( 200 1 ) louncl .I marketll!- smaller ratio of fawns::ldult females 1 1
October than in 1:inuarj: despitc the lom- incidence of harvested adult females. Consequentl!; the tradttionnl tirninl;: of spotlight count5 ii.e.. pxhunting se:ison) ma!. not generatc reliable estimates of recruitment. Our d;lt:~m d Langdon (2001) suggest that in the centrrtl Appa1achi:tnr of' \Vest Virgini:~ recruitment rates ma!. be mosr reliahl!- estim;~tecl from observational o r spotlight count dat:t collected from Februar!.-April. the months nlheii our capture ratios w e r r similar to the other estimates or extrapolations. Population status and health should be indexed through direct comparisons of reproductive and recruitment rates. Comparisons of reproductive data (as a measure of prior nutritional condition and an estimate of fawn production) and recruitment data (collected during winter months with minimal bias) could b e expressed as a crude fawn survival rate (?;I fawns:100 adult females born that reached recruitment age). Changes in crude fawn survival rates can guide mamgement recommendations independent of density estimates because both reproductive and recruitment rates arc functions of density relative to carrying capacity (McCullough 1979).
Management implications
Our observed survival. mortality. capture, and reproductive data are characteristic of a population shaped by moderate fawn recruitment and excessive harvest of yearling male deer. We recommend a reduction in female deer to promote forest regeneration in harvested stands and to protect and conserve mature forest community components, such as vernal woodland, herbaceous flora, or understory vegetative cover necessary for ground-nesting neotropical migratory songbirds. Failure t o respond to white-tailed deer overabundance on the MWWERF and throughout the central Appalachians of West Virginia may produce alternative steadystate forest conditions of low economic utility and reduced wildlife and biodiversity values (deCalesta and Stout 199'. Stromayer and Warren 199-.Wller and Alverson 1 9 T . Pedersen and Wallis 2004). Van Deelen et al. (199') noted that in northern Michigan. differential mortalin. rates hetween sexes were the product of male-biased hunting regulations and recommended increasing antlerless permits ro relieve hunting pressure [In males. Likmise. we recommend [hat t a t r wildlife agencies wlthin this region continue ro iibcr;llize :mrler-
Deer:populat~on-characteristics Carnpbellaet al.
2 9 1 1
ciet-r- 1ourn:il of\Vilcliite hlamgemrnr (10' i 7 i - 3 8 0 It-s.\ deer season Hecmsc nnny hunters in the central Appal~~chians nor willing to h a r v r s ~ L I I L .. 19-0. \X lldl~fcrn;~na$nien~ant1 the tl\.nnn~lc\01 are ungularr p)pul:iuona. .4ppl1etl I31olog!- I : 18;-1iO antlerless deer. being comfortnblc with tlie trxli- Lil.\l~l, . 5 . I.. I<. A . (r~lllL\l1\51EH \\I> 5 . 1; . I\ 11 tionai deer management paradigm of' buck-onl! O O O . ?'hc pcogrsph!. o f inlper~inlent: r:irp3iog c o ~ i I:L&~.I, sen-:lrlon t o n x d \ cr~tical biodlrrrs~r! area\ Page.; Ii')-200 Ilunting. n ce 7recoinmend that educational programs ill 13. A. Stein. L. S. hutner. and 1 . 5. Ad:tnl>. editon I'rrcioui (via start. wildlifc agencies, extension service.s. and l ~ e r i r ~ g ctlie status of hlodivrrsir! in thr Ln~tecl St:itei .: private orpnizations) continue to bc directed at LISA OsIord 1Jnlverhlh Press. Ke~\-Yurk. Xe\~>i)rl<, hunters ~ l arc reluctant to harvest a~itlerless ~ o deer (LI\-I.R. R I K. 195-1. A portablr deer trap and catch-nc~ (e.g..see Alt ZOO?). These programs shoulcl clearl!. Californl:~Fish and C,ame 40: j(~--Z'j I\ outline the ecological and biological need and ben- COI.I> . 1LJ. 200-t. l i e effects of prescri1,etl fires. c:tnopi, g q ~ s and deer lterhtvor! on tree spccics cotnposiwn 1mp11c:tefits of harvesting female deer (c.g..see Hamilton e t tions for succession t l i r o ~ I)~ssrrk~tion. linivrrsit!. 0 1 we al. 1995). F~~rtherniore. reconlmend that corpoI'itt~lx~rgli. P~ttshurgh. I'ennsy1vanl:i. I!Sh rate landowners pursue cooperative agreements : \ 1 1 ~ . 1 . . . \ S H L I ~. . I S 2001 with hunters, implementing incentive (e.g.. earn-:iEuropem gyps) moth (L~,?ilniztrrr disprrr L.) outbreaks: ;I buck) programs to encourage hunters to harvest rcvlr\v of thc literilturc. United Sc:ltrs Department of Agriculture Forest Service, Northtrastern Rese:irch Station. female deer.
,
Acknowkdginents. Financial support was provided b ~ tlie MeadWiestvaco Corporation and the United States Department of Agriculture's, National Research Initiative Program (grant 00-35101-9284). Editorial comments to earlier drafts of this nianuscript were gmciously provided by R.J.Warren, J. I? Carroll. and R. L. tiendrick. W are indebted tu the 30 field technicians who assisted in data collection and J. L. Crum. P D. Keyser, and D. A. Osborn for their logistical assistance. Deer were captured and liand i d under the auspices o Scicniiiic Coiicction T Permits 43-1999. 16-1000, 3001.008. and 2007.009 from the West Virginia Division of Natural Resources.
General Technical Report NE-278. Newtown Squarr. I'mnsylvania. USA. : 1).4vi1)so~.W l<.,xNll1 F ~'ETTLF? 1997. Field manual o f wildlife diseases in the southeastern United States Second editton Soutl~rasternCoopentlve Wildlife Disease Stud!. I Jniversity of (;eorgia. Athens. USA. 5. I)ECAI.EST.I.D. 1994 Deer and diversity in .Ulegheny hardwood forrsrs. managing an unlikely challenge. Landscape and Urtxo Planning 28. 4--53. IIFCALLST.&. 1). S..ANI) L. STOI:T. 199-. Relative deer densln and S. susta~nabilin.:a conceptual framework for integrating deer management with ecosystem management. Wildlife Socieq Bulletin 75: 252-258 De\oltsc.. C. A. 198'). Morrality of adult white-tailc~ldeer 111 south Texas. Journ;il of Wildlife Management 53: i .i-5 I 8 1 (i. J. B. B. 1989 Dl SEX. L.. R. J. NL\'L\cKIE.D. HERRK;ES.AND COMPTON. Population ecology of white-ratled deer along the lower k~ellowstoneRiver. Wildlife ,Monographs 104. ERERHARI)T. 1985. Assessing the dynamics of wild popul;~L. L. t~ons. .loumal of Wildlife Management 49: 99-- 1012. R. FIS(:HER.,I. 1996. White-tailed deer herd health check on the Westvaco Wildlife and Ecosystem Research Forest Southe:lstern Cooperative Wildlife Disease Study. Final Report to the West Virginia Division of Natural Resources. Charleston. USA. FORD.W .M.. 5. McC.n-, M. A. MENZEL. D. WEI)STW, . H. T. V(: C C;I(EENI%EK~~,J. E PAGEU.ANI) M E R I ( I 2 0 0.5 . Influence of eleJ. F ~~ vation ;uld forest n p e o n shrew communin. assemblage and s p e c ~ e distr~burlon the central and southern Appalach~an s in .Mountains. In I . E Xlerritt and S. B Churchfield. rdirors i c c o n d lnternat~onal C o n q e s s of the Biolo~!, o! the 5orlc1ciae. Carnegle Museum d Natural Histon 5peci;il I'ubl~c:~tion. press In \V .\vv 1001 Soricid .ibunckmce in parI.?)RI), M.. I . I.. ROI)RIGI'E. n:tI owrstory removal harvests :ind nparun :ireas ~n :in indus[nal torest landscape oi the c e ~ i r n l \ppalacluans. Forest Ecolog!, and Management 1 ;7. 159- 168 S. 1W8 I:I~EI)EIUCK~I:~.T. Impacts of lo,qginr: and developnienr on ~ r n r r ; l l .ippalachlan forests. Vatunl .\re;is ,loumal 18.
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1 W 1 tl;~rvcst stratc1 I O . .1 . I ) L ( I I eclitor> gies Pages .i:-i-111 fi \: ,\Iiller am! K. L hl:~rchinton. ()ualir!- whirctails: t h e n-li! ;mcl hou CIS qu:~lit?deer Inanaprnlrnr. Stackpole Hooks. Mechanicsburg. Penns!'lvani;l. 1!Sh. 1 1 : ~ 19(1S H I R. E , L I . , \ I R C L II ~ , G 7 . I Lannon-netting tlcer ,lournal of l'ildlitc hla~ugement-52. 19-195 HLISEI.1 > I . , A N D T K. Fl I 1.~1: 1985 Evnluati~jnof surviv:iI and 1. : cause-specific mortality rates uaing telcmetr! d a t z lournai o f W~ldlite Ivlanapemrnr 4% 66%-67-1. H o w ~ r lS.H . . 5. L. S m n . r \ ~ iD. 5. DECALESIA . ) 200.5 Whttr-tailed on d e r r ~ n ~ p ; i c r the vepetatlon dynamics of a northcrn hartlwood forest. Ecological Applicat~ons1 i:98-118 l0O5 I H...\ J A C ~ L ( \ I O N Starting out. Page5 1%46 KIWI.J. C..ANI) i l l K. \' iMiller :1nd R. L. Marchinton. edirors Quality whiretniis t h e why and how of qua11t)- deer man:cgemrnt ht~kpole Hooks. Mechan~csburg, Pennsylvania. IJSA. of Ls~c,t)os. C.A. 200 I . A con~parison white-railed d r e r population esrlmation methods 1 1 West Virginia. Thesis. West 1 Virgmia University Morgantown, USA. hK L L. 5. 2000. Seasonal variation in coyorr feeding hehnviour . .. . and mortality of w-Me-tailed deer and mule deer. Canadian Journal of Zoology 78:85-99. MCCLLI.OI:C;~~, 15179. The George Reserve deer herd: popuD R. larion ecology of a K-selected species. University of M~chiganPress, Ann Arbor. I!SA. MCCCLLOIIGH. 1981. Evaluation of nighc spotlighting as a D. R. deer study technique. Journal of Wildlik Management 46: 96.3-9-3. D. P. A McCuu.o~:c,~. R., E W. WWCERLY,1. GARCIA. N D R. R. E v m 1 9 9 4 Source:. of inaccuracy in black-tailed deer herd composition counts. Journal of Wildlife Management 58. ?Ic)-329. M I ~ A E E. D. 1%-. Bark stripping by white-railed deer in \Vest I., Virginia. Northern lournal of Applied Forestns t : 96-9'. MICHAEI.. D. 1991. Impact of deer browsing o n regeneration E. of balsam fir in Canaan\JalleyWestVirgln~a.Northern Iournal of Applied Foresrr). 9: 89-90. L. NELIOP.. E.,ANI) D. MECH.1981. Deer social organization and .M. wolf predation In northeastern Minnesota. Wildlife Monograph T . N E L ~ N . E.,AND D.MEcI-I.1986. Mortalin of white-tailrd deer M. L. in northeastern Minnesota. Journal of Wildlife Management 50:691-698. L. J. E CHEUWG.1091. NIXON. M., I? HANSEN. P A . BREW'ER.ANI~ C. Ecology of white-railed deer in an ~ntensivelyfarmed regloll of Illinois Wildlife Monograph 118. G. 1960. Sratist~calcalc~~lations for PAIILIK. I . . .\ND D 5. ROB~ON. change-in-rdtio estimators of popularlon panmeters. Journal ofwildlife Managemem 53: 1-37. I-'EDERSEN. B. S..ANI~A. M W ~ ~ ~ Effect5 o f white-tailed deer A 2004. . herbivon- on forest gap dynamcs In a wildlife preserve, I Pennsvlvan~a. JSh. Natural .Lreah Journal 2-i:SL-94. Rm .\. .M.h., K. G. AYTHO~T, H. J.WKSON.I ) 5.A. WOI.FE.200L D AN 5urvlval ot Coiumbran white-talled deer In mestern Oregon. ,loumal ofX'ildlife Management 66. l 2 i i - 1 3 6 . I ? M I ~ ' I I 1) W' I ? 9 i he southern .\ppalachian hardwoi~d region.
P;~pc. I-?-l.?i I I I I.\\' I$:trrcrt.ctlirni~.Kcgloni silviculture. n: \X Xeu-lorl, tlir I n ~ r e d htate5 ~ o l i n iJr?.m d >on.. !\e\v'~orl~. I is:\ Soh.\l. ll 1 ...\ Y I ) TI. KOHII 1905 Uionwrr! Th~rti : echrion \\: E l Frecrn:ui and Conipa~i!~. hex\-l'orl,. hcn-Yorl,. I:S.\ SOI-10, R . 4 \ 1 1 K. 5 . WI!.I.IP 2 0 0 0 chwvic\\. of lieniiocl. 11 K s and l~e:il111 1';1pr+-b-XO I I I I< 2 . M c ~ l a n ~ ~ S..Sh~elds. 11. 1; Souto. editors. I'roceecllng\: symposnum ot sust:unable rn:m agrrnent of Iirmlock ccos!xernh in r a s c r n k l r t l i Anieric:~ Unitccl State\ Drpartrneni ol' Agr~culturr Forest Srrvicc. hortlieasrern 1~rse:irch bration. <;enrr;~l1kchnic;tl Rrpor: NE-26-. lic\vtown Squ:~rc. I'rnnsylvanis. USA SPEKLI:. A. 199-, Adding v;~lut. p i e c t h- piccc. Alount:lln ! Prornisr '. I - 5 . STFIWEV\OIL. 19Oji1. AN introductwfi t ~ tllt ~~plan(I S I. ) fores~ repon P ~ g e s 1-0 1 1 1 S. I.. Stephrnson. editor. I 'pland torests of West Virginia McCl;iin Prtnting. I'arsons West Virginia. USA. SIWHENWN. 19936. Upland forest vegetation. I'ages I1 -3-1 S L 1 1 1 S. L. Stephenson. cdttor Upland forests of W s t \irgmi:l. hlcChin Printing. Parsons.U;est Virginia, USA. R. STIPE~,J 1999. Trojan horse in Appalachian forests. Pages 73-79 i71 R I? Eckerlin. editor. Proceedings of the Appalachian Biogeography Symposium. \'irgini.r Museum of h:ttural Histoy Special Publication N o . -, M;minsville. USA. STRO~~\\YEI:. AND R. J.%RI:E~ 10% Are overabundant derr K.A., herds in the eastern United Scares creating alternative srable states in foresr plant communities? Wildlife Sociery Bulletin 15:32--234. TILGHMAN, N. G. 1989 Impacts of white-railed deer on forest regetirr~tion In northwestern Pennsylvania. Journal of\Vildlife Management 53: 521-532. IJYITED STATESDEI'ARTMENT 0 . AGRICITTURE 1 FORES~. SER\'T(X ?()()-J Northcutern forest Inventory and analysis: West Virginla results. U n l ~ r d States Ikpartmenr of Agr~culturr Forest Servlcc. Northeastern Research htarion. Newtown Square. Pmnsylvanra. USA. Available online ar: hrrp://mr.fs.fed.us/ ne/fia/srares/wv/indrx.htd (accessed 1 May 1004). T. . A~ . 1 THOMI'SOIL. V A D E E L E ~K.. H. CAVI'A. 111, 1. B.H h l i l : ~ ~ .N1 1'. D. ~ 1 9 T Mortalin patterns of white-railed deer in Mich~gan's iipper Peninsula. Journal of Wildlife Management 61: 903-910 L. D..ANI) L. SHERIFE S. 1090. Survwal estimation when VANGIL~ER. fates of some animals are unknown. Tmns:icrions of the Missour1 Academy of Sc~ence 5--68. Zt: VAN VLREN. . , . ~ NS.) E . T H ~ M I W V . J 1981. Opportunlsric feedD I R. ing b. coyotcs. Northwest Sciencr 56: 131- 135. ! L. . \ J r ~ < w ~ .I.. \ N I ) D. E. ULLKCY 198't. Phvsiol~gy:ind ntltrltlon. Pagcs ?I -1 It; 117 L. K. Halls. ediror. Whitr-tailed deer eco1og)- and management Stackpole Books. Harr~sburg, I'ennsvlvania. IiSA. \JREELA.W.J 3001. F a w ~ sun.lva1 In Pennsylvania Prnnsvlvania ( h m e Yews -3(0). 12- 10. W,ux D.A.. \Y[) I E.B O W N ~1984. I ' r o c ~ d ~ i r e s r\.aIuat~nl: . for preclat~on livestock and wildlife. Texas Agricultural Estens~on on Senxcc. (:c)lle~c >cation. I!S.i. X A I I R . I .1 I Iv . L ~ 1O 90-. The whire-t:cilcd deer: ;L !;evstone herbivire. Wildlife Socien- I k ~ l l e t ~ n 25.21--226. (.. 'Xrr:~\#l.\\~). .i 3000 Effects of diameter-lim~t.ind rwo-~#ed t ~ m h e rliarvestin!: on wnghird populations o n :in ~nduscrnl \Vest \ir$mxa lorest 111 ccntml W s t \irzinra. L)~ssertat~on. 1 inivrraln. .\loryntown i 'S.4. \VFYI?YOI