Temporal and spatial development of red deer harvesting in Europe: biological and cultural factors

Journal of Applied Ecology

2006

43

721ndash734

copy 2006 The Authors Journal compilation copy 2006 British Ecological Society

Blackwell Publishing Ltd

Temporal and spatial development of red deer harvesting in Europe biological and cultural factors

JOS M MILNERdagger CHRISTOPHE BONENFANTdaggerDagger ATLE MYSTERUDdagger JEAN-MICHEL GAILLARDDagger SAacuteNDOR CSAacuteNYIsect and NILS CHR STENSETHdagger

Hedmark University College Department Forestry and Wildlife Management Evenstad N-2480 Koppang Norway

dagger

Centre for Ecological and Evolutionary Synthesis Department Biology University of Oslo PO Box 1066 Blindern N-0316 Oslo Norway

Dagger

Biomeacutetrie et Biologie Evolutive Universiteacute Claude Bernard Lyon 1 43 Bvd du 11 Novembre 1918 69622 Villeurbanne Cedex France and

sect

St Stephen University Department Wildlife Biology and Management H-2103 Goumldoumlllotilde Hungary

Summary

1

Deer numbers have increased dramatically throughout Europe and North Americaover the last century but empirical analyses of variation in harvesting and the influenceof biological and cultural factors are lacking

2

We examined trends in size and composition of red deer

Cervus elaphus

harvests overthe last three to four decades in 11 European countries with contrasting deer productivitymanagement strategies and hunting traditions

3

The harvest increased exponentially in all countries except Austria and Germanywhere it was stable and Poland where it has declined in recent years Harvest growthrates ranged from 0middot009 in Austria to 0middot075 in Sweden and depended on the managementsystem and harvest composition being negatively related to the proportion of femalesin the adult harvest

4

Within four focal countries (France Hungary Norway and Scotland) there was con-siderable spatial variation in harvest growth rates These tended to be higher in recentlycolonized areas than in traditional hunting areas and were often higher than the maximumpossible population growth rate Range expansion was an important component of theincrease in total harvest in France and Scotland but not in Hungary or Norway

5

Harvest composition was available for seven countries all of which showed a strongincrease in the proportion of calves in the harvest The sex ratio of the adult harvestwas relatively stable being strongly male-biased in Norway and marginally female-biased elsewhere The proportion of males in the harvest was unrelated to trophyhunting objectives

6

Synthesis and applications

Our study emphasizes that cultural aspects of managementneed to be accounted for as well as biological factors when interpreting the patterns ofharvest growth and composition across Europe Widespread sustained harvest growthhas occurred suggesting continued growth of deer populations with consequent socialand economic impacts Population control is therefore a major challenge for the futurecurrently hampered by inadequate population data and a decreasing number of huntersin some countries Increasing the motivation of hunters to harvest female deer is onepossible solution although this may conflict with hunting traditions and economicconsiderations in some areas

Key-words

deer management exploitation large herbivores population dynamics selectivehunting sustainable management ungulates wildlife cropping


(2006)

43

721ndash734doi 101111j1365-2664200601183x

Correspondence Jos Milner Hedmark University College Department of Forestry and Wildlife Management EvenstadN-2480 Koppang Norway (fax 0047 62430851 e-mail josmilnerhihmno)

722

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et al



43

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Introduction

The numbers of many deer species have increaseddramatically throughout Europe and North Americaover the last century (Gill 1990 Clutton-Brock amp Albon1992 McShea Underwood amp Rappole 1997 Cocircteacute

et al

2004 Gordon Hester amp Festa-Bianchet 2004) as a resultof both increases in density and range expansion Thereare a number of causes both direct and indirect with nosingle explanation applicable to all countries or species(Fuller amp Gill 2001) Factors unrelated to cervid manage-ment include increased availability of forage resultingfrom changes in land use (Ahleacuten 1975) agricultural andsilvicultural practises (Mysterud

et al

2002) and theclimate (Mysterud

et al

2003) However the directmanagement of cervids has also played a part Forexample protection from overhunting and poachingthe use of selective harvesting regimes (Langvatn amp Loison1999 Solberg

et al

1999) supplementary feeding inwinter (Peek

et al

2002) and reintroduction programmes(McShea Underwood amp Rappole 1997 Mattioli

et al

2001 Leduc amp Klein 2004) have all contributed to risingnumbers As a result in many areas management objec-tives are beginning to change from species protectionto population control (Buckland

et al

1996 Brown

et al

2000 Cocircteacute

et al

2004 Gordon Hester amp Festa-Bianchet 2004)

Red deer

Cervus elaphus

L are widely distributed acrossEurope (Fig 1 Koubek amp Zima 1999) and have increasedin both abundance and geographical range in recentdecades (Gill 1990 Csaacutenyi 1992 Mattioli

et al

2001Clutton-Brock Coulson amp Milner 2004) althoughreliable quantitative data on population sizes aregenerally lacking Red deer are primarily found inforest or woodland-edge habitats often associated withland of low productivity including open moorland inScotland The widespread distribution and large bodysize of red deer makes it an important game species(Koubek amp Zima 1999) Hunting rights in Europe aregenerally held by the landowner (Gill 1990) in contrastto North America where hunting is accessible to a widesector of the general public (OrsquoGara 2002) Consequentlyhunting in Europe tends to be limited to landowners(state or private) andor their associates and payingleaseholders or trophy-hunting clients The size of reddeer harvests has increased considerably in manyEuropean countries over recent decades (Gill 1990)

Harvesting represents a major source of mortality inmany North American elk

Cervus elaphus

L populationswhere predators are absent (Ballard

et al

2000 Bieder-beck Boulay amp Jackson 2001 McCorquodale Wisemanamp Marcum 2003) While few empirical studies havedocumented this in Europe it is also likely to be true formany red deer populations (Langvatn amp Loison 1999)although predation may play a more significant rolein some protected areas (Jedrzejewski

et al

2002) Con-sequently selective harvesting is likely to influencestrongly population composition dynamics (Langvatnamp Loison 1999 for moose Alces alces see Solberg

et al

1999) and genetics (Coltman

et al

2003) At the Euro-pean scale the basis for red deer management variesdepending on cultural traditions and the environmentalconstraints of habitat and latitude (Bonenfant

et al

2004Loe

et al

2005) We might therefore expect harvest ratesselectivity and composition to differ between countrieswith consequences for population dynamics Howeverit is currently not possible to address empirically howregional variation in selective harvesting regimes hasaffected population dynamics as the required data simplydo not exist for most red deer populations In the absenceof such data we aimed to describe the main patterns inthe size of the red deer harvest over the last three to fourdecades in 11 European countries with contrasting hunt-ing objectives management strategies and environmentalconditions In addition we investigated trends in the com-position of the harvest in seven of the countries and madea more in-depth analysis of spatial variation in fourfocal countries France Hungary Norway and Scotland

Harvest development is likely to follow general trendsin population development (Forchhammer

et al

1998and references therein) although often with a 1ndash2-yeartime-lag (Fryxell

et al

1991 Solberg

et al

1999) Harvestsize is also influenced by policy and cultural factorsincluding harvesting strategies (Giles amp Findlay 2004)and quotas which explain some of the time lags(Solberg

et al

1999) We therefore also investigatedthe following predictions (i) P1 harvest growth rateswill be highest where deer populations are growingfastest Consequently we expect harvest growth ratesto be lower in traditional deer hunting areas than inareas of recent colonization where lower densities andthus higher recruitment rates are likely (unless habitatdiffers markedly) (ii) P2 harvest growth rates will belower with a higher proportion of adult females in theharvest because of the sensitivity of population growthto adult female survival (Gaillard Festa-Bianchet ampYoccoz 1998) assuming a close link between popu-lation growth and harvest growth (iii) P3 there will bepositive covariation between harvest size hunternumbers and size of red deer range (iv) P4 a higherproportion of adult males relative to females will occurin the harvest in trophy-hunting cultures (Ginsberg ampMilner-Gulland 1994)

Materials and methods

Annual statistics for the size of the total red deer harvestwere obtained for Austria Denmark France GermanyHungary Norway Poland Scotland Slovenia Swedenand Switzerland from national bodies in the respectivecountries (Table 1) The year of harvest was taken fromthe start of the hunting season Harvest compositionin terms of the number of calves adult (includingyearling) males and females shot were available forseven countries (Table 1) Data describing populationsize and structure were unfortunately unavailable formost of the countries studied However as our investiga-tion was of variation at a continental scale we assumed

723

Red deer harvesting in Europe



43

721ndash734

the spatiotemporal variations in age and sex structureat smaller spatial scales would be dampened

The number of hunters per capita in 2004 (FACE 2004)was used as an indicator of the strength of the huntingculture in each country while the deer managementsystem was classified according to the groupings identifiedby Gill (1990 Table 1) Overall hunting objectivesrepresentative of a given country were based on personalcommunications with experts In recent years objectiveshave been shifting towards population control in severalcountries (eg SNH 1994)

A more detailed analysis of the spatial variation inharvest growth and composition was carried out at aregional level in France Hungary Norway and Scotlandwhich are representative of the main deer managementsystems in Europe In France and Norway huntingis regulated by a quota of licences issued by wildlifeauthorities to landowners or hunting-right holders(management system 2 Fig 1) In Hungary hunting-rightholders (hunting clubs state farms and forests) are

Fig 1 Map of the current distribution of red deer across Europe by 50-km squares (Societas Europaea Mammalogica Koubekamp Zima 1999) showing management systems in each country based on Gill (1990) The few small symbols represent presumeddistribution from data collected before 1970

724

J M M

ilner

et al

copy 2006 T

he Authors

Journal compilation

copy 2006 B

ritish E

cological Society

Journal of Applied

Ecology

43

721

ndash734

Table 1

Description of hunting systems harvest data and harvest growth rates from each study country

Country RangedaggerManagementsystemDagger

Hunting objective

No hunters ( of population)sect

Hunting season (M males F females)

Harvest data period (harvest composition)

Harvest growth rate

H

r

(SD)Probability

H

r

gt 0

Proportion males in adult harvest (2002)

Proportion calves in harvest (2002) Data source

Austria 34 1 Trophy minimise forest damage 1middot4

M 0108ndash3112F 0107ndash3101para

1965ndash2002 (1965ndash2002)

0middot009 (0middot066) 0middot210 0middot42 0middot34 Zentralstelle Oesterreichischer Jagdverbaende

Denmark 15 2 Recreation meat 3middot2 M 0109ndash3101 1965ndash2002 0middot042 (0middot131) 0middot030 Asferg amp Olesen (2004)F 0110ndash3101

France 155 2 Trophy meat 2middot3 1511ndash3101 1973ndash2002 (1983ndash2002)

0middot056 (0middot077) lt 0middot001 0middot48 0middot30 Office National de la Chasse et de la Faune Sauvage

Germany 113 1 Trophy minimise forest damage

0middot4 0108ndash3101daggerdagger 1982ndash2002 0middot012 (0middot069) 0middot230 Deutscher Jagdschutz-Verband

Hungary 34 1 Trophy population control

0middot5 0109ndash3101 1965ndash2002 (1969ndash2002)

0middot050 (0middot125) 0middot010 0middot36 0middot39 Hungarian Game Management Database

Norway 66 2 Meat recreation 4middot8 1009ndash1511 1965ndash2002 (1977ndash2002)

0middot062 (0middot091) lt 0middot001 0middot56 0middot23 Statistics Norway

Poland 116 1 Trophy 0middot3 M 2108ndash2802F 0109ndash1501

1965ndash2002DaggerDagger 0middot029 (0middot139) 0middot148 Central Statistical Office Warsawparapara

Scotland 41 4 Trophy population control

1middot3 M 107ndash2010F 2110ndash1502

1965ndash2002 (1965ndash2002)

0middot031 (0middot091) 0middot024 0middot48 0middot14 Deer Commission for Scotland

Slovenia 7 1 Trophy population control


1969ndash2002 (1969ndash2002)

0middot041 (0middot141) 0middot052 0middot46 0middot44 Lovska Zveza Slovenije

Sweden 17 2 Recreation meat 3middot6 1608ndash3101 1965ndash2002 0middot075 (0middot216) 0middot021 Svenska JaumlgarefoumlrbundetSwitzerland 16 13 Recreation

trophy meat0middot4 0109ndash3101 1965ndash2002

(1965ndash2002)0middot035 (0middot148) 0middot082 0middot57 0middot19 Bundesamt fuumlr Umwelt Wald

und Landschaft

daggerRelative distributional range given as no 50-km

2

occupied (Societas Europaea Mammalogica Koubek amp Zima 1999)DaggerManagement system as per Gill (1990) fig 3middot1 where 1 systems requiring detailed harvest plan 2 systems where harvest is controlled by licence and land or hunting ground ownership 3 system where harvest is controlled by licence only 4 system where harvest is controlled by landowner only (Fig 1)sectFACE (2004) Note data for Scotland not available so value for whole of UK is presentedparaSeason for yearlings begins 0106 all dates vary

plusmn

1 month between provincesdaggerdaggerSeason for yearling females 0106ndash3101 yearling males 0106ndash2802DaggerDaggerMissing data in 1977 1986ndash89 1991ndash93paraparaStatistical Yearbook of the Republic of Poland Warsaw 1965 1968 1970 1975 1981 1985 1997 2002

H

r

significantly greater than zero assuming probability threshold of

P

= 0middot0045 after Bonferroni correction

725




43

721ndash734

required to provide annual counts detailed harvest plansand reports for an age- and sex-structured cull (manage-ment system 1 Fig 1) In Scotland individual landownersor groups of neighbouring landowners decide how manydeer to hunt on their land (management system 4 Fig 1)Further background information about the focal countriesis given in Appendix S1 in the supplementary material

French harvest data came from a total of 84 depart-ments ranging from 60 departments in 1973 to a peakof 81 in the late 1990s Hungarian data came from the19 counties of which 13 had a red deer harvest in 1965and all have harvested deer since 1979 In Norwaythe number of municipalities harvesting deer increasedfrom 93 in 1965 to 222 in 2002 The number of calvesvs adults in the harvest was available from 1966 witha full breakdown by age and sex from 1977 Scottishdata came from the statutory cull returns for individuallandholdings grouped into blocks corresponding toDeer Commission for Scotland (DCS) counting areasThe number of blocks increased from 34 in 1965 to 74in 2002 However the total harvest was an underestimatebecause of a lag in reporting of shot animals on newlyhunted landholdings (M Daniels personal communica-tion) The total harvest within traditional red deer hunt-ing areas was less subject to this bias Red deer shot incommercial forestry plantations were included Thetotal number of hunting license holders per year (large andsmall game) in France (partial time series Directiondes Etudes et de la Recherche de lrsquoONCFS) Hungary(Hungarian Game Management Database) and Norway(from 1971 Statistics Norway) was used as an indexof hunting effort while the sum of areas of each regionwhere harvesting occurred was used as an index of thered deer range in all focal countries

Harvest growth rate

The harvest growth rate (

H

r

) was calculated for eachcountry as the mean of the annual change in harvestsize log

n

(

h

t+

1

)

minus

log

n

(

h

t

) where

h

t

is the size of the totalharvest in year

t

It is analogous to population growthrate (Caughley 1977) with which it appears to bepositively correlated (S Csaacutenyi amp JM Milner unpub-lished data) and as such is necessarily a temporaryphenomenon At a smaller spatial scale harvest growthwithin focal countries was determined by calculating

H

r

for each region with at least 5 years harvest dataHowever some regions had a zero harvest in years whenno deer were culled so 1 was added to enable naturallogarithms to be taken (Sokal amp Rohlf 1981) Regional

H

r

was therefore calculated as the average of log

n

(

h

t

+1

+ 1)

minus

log

n

(

h

t

+ 1) for each region

Comparison between countries

For each country

t

-tests were used to assess whether

H

r

was significantly different from zero making a Bonferroni

correction for 11 tests (threshold

P

-value = 0middot0045 Sokalamp Rohlf 1981) Harvest growth rates were then comparedbetween countries and management systems over timefitting linear models using generalized least squares (GLSsee Appendix S2 in the supplementary material) Thesemodelled both the mean and variance of the responsevariable (Pinheiro amp Bates 2000) and allowed us toaccount for the observed unequal variances betweencountries (Bartlettrsquos test

T

= 85middot98 df = 10

P

lt 0middot001)when comparing

H

r

H

r

was the response variable year(covariable) either country (11-level factor) or manage-ment system (four-level factor) and the respective first-order interactions with year were entered as explanatoryvariables The variances were allowed to differ inde-pendently according to the levels of the country factorFinally we checked the model adequacy residualnormality and homoscedasticity against our data setgraphically (Venables amp Ripley 1999)

A similar procedure was used to investigate the influenceof hunter numbers and range area (explanatory variables)on harvest size and harvest growth rate in the focalcountries (excluding Scotland for analysis of hunternumbers) Here ln(national harvest size) and

H

r

atthe national scale were considered as the responsevariables respectively and country was fitted as anadditional explanatory variable Hunter numbers andrange area were used to explain the observed time trendin ln(national harvest size) rather than fitting thecovariable year with which they were strongly correlatedAgain the variance was allowed to differ independentlyaccording to the levels of the country factor Similarlythe influence of the proportion of females in the adultharvest on the annual rate of increase in harvests wasexplored using GLS models Country year and theirinteraction were fitted as covariables but none wassignificant

Comparisons within focal countries

Within each focal country regions were classified byhunting tradition as either traditional or non-traditionalred deer hunting areas (two-level factor) In HungaryNorway and Scotland regions were classified as tradi-tional if deer were harvested there from the beginningof the study period In France department-level dataprovided a poorer spatial resolution so traditionalareas were taken as those in which red deer occurred in1900 (Leduc amp Klein 2004) The time series for each ofthe non-traditional areas began in the first year ofharvesting

Differences in

H

r

between traditional and non-traditional areas across countries were investigated withina mixed-model framework that accounted for the correla-tion structure arising at the country level because datawere collected at the region scale A linear mixed modelwas used fitting country as a random effect on both theintercept and the regression slope and accounting forunequal variances between countries (Pinheiro amp Bates2000) Subsequently comparisons within countries were

726

J M Milner

et al



43

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made with linear models having established that includ-ing region as a random effect did not significantly improvemodel fit (France likelihood ratio 0middot007

P

= 0middot932Hungary L ratio 0middot009

P

= 0middot923 Scotland L ratio =0middot003

P

= 1middot0) except in Norway (L ratio = 8middot547

P

=0middot036) Similarly a mixed-model approach was takento investigate the effect of proportion of females in theadult harvest tradition and their interaction on regional

H

r

in the focal countries Region was fitted as a randomeffect on both the intercept and the regression slopeIn all these analyses year was dropped as a fixed effectbecause it was not significant in any model

Harvest composition

Changes in harvest composition (proportion of calvesmales and females) were investigated in the seven countriesfor which composition data were available and withintraditional and non-traditional hunting areas in thefocal countries The dependent variables were binomialtreating number of calves as 1 and non-calves as 0and similarly for the number of males (1) vs females(0) in the total adult harvests Binomial responses wereanalysed with generalized estimation equation (GEE)models (Diggle

et al

2002) We used a logit link functionwith a binomial error distribution and an lsquoexchangeablersquomatrix for the error correlation structure (Liang amp Zeger1986) GEE allowed us to handle the high degree of

overdispersion originating from the spatial and temporalcorrelation structure of the data for example countryor region and year For national models linear andquadratic year terms either country or managementsystem and their interactions with year were fittedas fixed effects and country as the grouping factor Asthere were marked differences in harvest compositionbetween countries the regional harvest of each focalcountry was modelled separately with tradition yearand their interaction fitted as fixed effects and region asthe grouping factor The effects of the explanatoryvariables were tested using the Wald statistic basedon naive

z

statistics and SE values which follows a chi-square distribution with

k

degrees of freedom where

k

is the number of parameters to be tested All analyseswere carried out using R 2middot0middot1 (R Development CoreTeam 2004)

Results

Total red deer harvests had increased by 400ndash700 in mostcountries over the last 30 years Exponential increasesoccurred in eight out of the 11 countries examinedThe exceptions were Austria and Germany which hadstable harvests and Poland where a strong increasehad been followed by a recent decline (Fig 2) Harvest

Fig 2 Trends in size of the (ln) total red deer harvests in 11 European countries since 1965 For management systems indicated by numbered labels referto Table 1

727




43

721ndash734

growth rates varied from 0middot009 in Austria where theannual harvest fluctuated between 31 000 and 46 000to 0middot075 in Sweden which had the smallest harvestrising from around 90 red deer shot in 1965 to 1420 in2002 However harvest growth rates were only signifi-cantly greater than 0 in France and Norway aftermaking a conservative Bonferroni correction formultiple

t

-tests (Table 1) During the study period wefound no evidence of a change in harvest growthover time in any country (

F

1365

= 1middot653

P

= 0middot199 seeAppendix S2 in the supplementary material) suggestingthat deer populations were growing at a faster rate thanthe harvests

Differences in national harvest growth rates werenot significant (

F

10356

= 1middot55

P

= 0middot12) because of highvariability over time within countries However harvestgrowth rates differed significantly between managementsystems being higher in countries where the harvestwas controlled by licence and land or hunting groundownership (management system 2 Fig 1 and Table 1)than in countries with other management systems (

F

1365

=9middot72

P

= 0middot002) Denmark France Norway and Swedenwere also the countries with the highest percentageof inhabitants who were hunters in 2004 (average 3middot5of the population compared with 0middot9) Indeed thepercentage of hunters in the population explained nearlyhalf of the variation in harvest growth rate betweencountries (slope = 0middot009

F

110

= 7middot86

P

= 0middot021 Fig 3a)In support of prediction P2 harvest growth rate was

significantly negatively influenced by the proportionof females in the adult harvest at the national level(Table 2a) Once the proportion of females in the adultharvest was fitted the management system explainedno additional variation (

F

3220

= 0middot367

P

= 0middot777) suggest-ing that this term accounted for much of the differencein harvest growth rate between management systems

Within the focal countries there was as predicted (P3)a positive relationship between ln(harvest size) andred deer range in all countries (Fig 3c r2 = 0middot82ndash0middot97see Appendix S3 in the supplementary material) andwith hunter numbers in Hungary and Norway (Fig 3br 2 = 0middot83ndash0middot95) However harvest size in France increaseddespite a decline in hunter numbers (Fig 3b r2 = minus0middot97)Neither hunter numbers (F = 0middot008 df = 1 P = 0middot931)nor range size (F = 0middot244 df = 1 P = 0middot622) had anyeffect on harvest growth rates

There was considerable spatial variation in harvestgrowth rates within focal countries (Table 3 and Fig 4)The highest Hr values tended to occur in areas of recentcolonization at the periphery of the red deer range(Fig 4) The high variance between areas in Franceand Scotland was partly associated with a pronounceddifference in regional harvest growth rates betweentraditional deer hunting areas and areas into which deerand deer hunting have expanded being significantlyfaster in these non-traditional areas (Table 3) Incontrast there was no difference in the harvest growthrate between traditional and non-traditional areasin Hungary and Norway (Table 3) Thus prediction P1

was only partially supported The variability in harvestgrowth rate was greater in non-traditional than tradi-tional areas in all countries (Table 3) In traditionalareas the harvest growth rate reflected an increase inharvest density (ie harvest per unit area) within regionswhile in non-traditional areas it reflected a combinationof both an increase in harvest density and an increase inthe number of regions in which harvesting was carried out(Table 3) However while the number of municipalitiesharvesting red deer for at least 5 years in Norwayincreased during the study period by 83 the contribu-tion of these new areas to the total harvest in 2002 wasonly 13 and in Hungary the non-traditional areascontributed only 3middot6 to the 2002 harvest In com-parison in Scotland the number of management blocks

Fig 3 Relationship between (a) harvest growth rate and thepercentage of inhabitants that were registered hunters in 2004in 11 European countries (b) harvest size and hunter numbersin the focal countries and (c) harvest size and deer range area inthe focal countries (note the log-scaled x-axis for b and c)

728J M Milner et al

copy 2006 The Authors Journal compilation copy 2006 British Ecological Society Journal of Applied Ecology 43 721ndash734

increased by 65 and non-traditional areas accountedfor 21 of the 2002 harvest while in France the numberof departments in non-traditional areas increased by25 and their contribution to the total harvest rosefrom 26 in 1973 to 50 in 2002 The increase intotal size of the Hungarian and Norwegian harvestswas therefore most influenced by an increase inharvest density in traditional areas while in Franceand Scotland range expansion was an important con-tributory factor

As at the national level the proportion of females inthe adult harvest also had a negative effect on regionalHr in all focal countries adding further support to predic-tion P2 However the relationships differed betweentraditional and non-traditional areas in all cases exceptFrance (Table 2b) being stronger in non-traditionalthan traditional areas in Hungary but only significantin traditional areas in Norway and Scotland (Table 2b)

Calf harvest

The composition of the harvest has changed dramati-cally over recent decades in the seven countries for whichdata were available (Fig 5) In particular there was astrong increase in calf harvesting in all countries manyof which prohibited or rarely practised this before theearly 1970s Averaged across countries the proportionof calves in the harvest increased more than fivefoldfrom 5 in 1965 to 29 in 2002 although in recent yearsit had reached a plateau in several countries (Fig 5a)By 2002 calves accounted for between 14 (Scotland)and 44 (Slovenia) of the harvest (Table 1) The changein the proportion of calves in the harvest over timediffered significantly between countries (countryndashyearinteraction χ2 = 108middot11 df = 6 P lt 0middot001 countryndash

Table 2 The effects of the proportion of females (PropF) in the adult harvest on the annual rate of increase in harvest in (a) thenational harvest in seven countries (Austria France Hungary Norway Scotland Slovenia and Switzerland) modelled by GLSand (b) the regional harvest in each focal country together with the effect of hunting tradition and their interaction modelled witha linear mixed model fitting region as a random effect on the intercept

Estimate (SE) F df P Random effect (95 CI)

(a) National PropF minus0middot284 (0middot096) 8middot779 1 0middot003(b) France PropF minus0middot244 (0middot095) 6middot527 11235 0middot0107 0middot521 (0middot377ndash0middot719)

Tradition minus0middot032 (0middot017) 3middot408 178 0middot069Hungary PropF minus1middot243 (0middot225) 12middot01 1544 lt 0middot001 0middot422 (0middot204ndash0middot874)

Tradition 0middot013 (0middot243) 40middot88 117 lt 0middot001PropFndashtradition 0middot904 (0middot432) 4middot383 1544 0middot037

Norway PropF minus0middot032 (0middot044) 2middot467 14504 0middot116 0middot297 (0middot227ndash0middot388)Tradition 0middot122 (0middot033) 23middot21 1212 lt 0middot001PropFndashtradition minus0middot166 (0middot081) 4middot167 14504 0middot041

Scotland PropF minus0middot112 (0middot139) 16middot95 11632 lt 0middot001 0middot403 (0middot242ndash0middot671)Tradition 0middot177 (0middot100) 5middot014 155 0middot029PropFndashtradition minus0middot436 (0middot187) 5middot414 11632 0middot020

Non-significant term not included in the minimal model F-value calculated as the change from the minimal model

Table 3 (a) Linear mixed model showing the fixed effects of country and hunting tradition on the change in harvest betweenyears with region fitted as a random effect (b) Mean harvest growth rates (Hr) in traditional and non-traditional red deer huntingareas where ni is the number of land units with red deer hunting in the initial year (1965 for Norway and Scotland 1973 for France)and nmax is the maximum number of land units during the study period Only land units with harvesting in at least 5 years wereincluded

(a) df F PIntercept 1 347middot6 lt 0middot001Country 3 4middot693 0middot003Tradition 1 5middot833 0middot016Countryndashtradition 3 3middot594 0middot014Residual 10254(b) ni nmax Hr (SD) Hr rangeFrance Traditional 26 29 0middot062 (0middot248) minus0middot006ndash0middot111

Non-traditional 34 52 0middot119 (0middot417) minus0middot058ndash0middot418Hungary Traditional 13 13 0middot075 (0middot172) 0middot040ndash0middot110

Non-traditional 0 6 0middot075 (0middot763) minus0middot030ndash0middot155Norway Traditional 92 92 0middot061 (0middot312) minus0middot004ndash0middot141

Non-traditional 0 122 0middot064 (0middot543) minus0middot154ndash0middot260Scotland Traditional 34 34 0middot037 (0middot285) minus0middot008ndash0middot155

Non-traditional 0 22 0middot096 (0middot512) minus0middot004ndash0middot500

P lt 0middot01 P lt 0middot001 significant difference in harvest growth rate between traditional and non-traditional areas within a country

729Red deer harvesting in Europe


year2 χ2 = 107middot02 df = 6 P lt 0middot001) managementsystems (management systemndashyear interaction χ2 =38middot32 df = 2 P lt 0middot001 management systemndashyear2χ2 = 37middot9 df = 2 P lt 0middot001) and within focal countriesbetween traditional and non-traditional areas (seeAppendix S4 in the supplementary material) Differ-ences between countries are likely to reflect in partbiological differences such as calving rates being lowerin Scotland (Clutton-Brock amp Albon 1989 MilnerAlexander amp Griffin 2002) than in countries withhigh female productivity (Langvatn et al 1996 2004Bonenfant et al 2002) In contrast differences betweenmanagement systems are more likely to reflect culturalfactors related to harvesting strategies For examplecentral European countries (management system 1)harvested a significantly higher proportion of calves thancountries where harvesting was controlled by licenceand ownership (management system 2) which showedthe fastest increase in calf harvesting and in turn harvesteda greater proportion of calves than other countries(Table 1 and Fig 5a main effect of managementsystem χ2 = 732middot92 df = 2 P lt 0middot001) Calves generallyaccounted for a higher proportion of the harvest in

traditional than non-traditional areas of focal countrieswith lower year-to-year variability (see Appendix S4 inthe supplementary material)

Adult harvest

As a consequence of an increased calf harvest adultsaccounted for a smaller proportion of the total nationalharvests in later years in all countries However withinthe adult harvest the proportion of males shot remainedrelatively stable at just below 50 in most countriesalthough there were considerable periodic fluctuationsin many cases (Fig 5b) Norway was the only countrywith a strongly male-biased harvest The trends at eitherextreme were a significant decline in the proportionof males shot in Slovenia from 0middot53 in 1969 to 0middot46 in2002 (χ2 = 16middot77 df = 1 P lt 0middot001) and a small butsignificant increase in France from 0middot46 in 1983 to 0middot48in 2002 (χ2 = 4middot27 df = 1 P lt 0middot001) The proportion ofmales in the adult harvest differed significantly betweencountries and over time (countryndashyear interactionχ2 = 14middot11 df = 6 P = 0middot028 year2 χ2 = 9middot37 df = 1P = 0middot003) Contrary to P4 there was no evidence of a

Fig 4 The spatial variation in the rate of growth in harvest size in (a) France at the department scale (b) Norway at themunicipality scale and (c) Scotland at the scale of management blocks Traditional hunting areas are outlined in red Areas withno harvest include those areas where the total number of deer harvested over the study period was less than 50 or where harvestingtook place in fewer than 5 years

730J M Milner et al


relationship between the proportion of males in theadult harvest and trophy hunting culture (Table 1)

Differences in the proportion of males in the adultharvest between management systems were lesspronounced than differences in calf harvesting Gener-ally countries with management systems 1 and 3 had alower proportion of males in the harvest than countriesin other systems However differences in the proportionof males in the harvest in Norway and France both undermanagement system 2 (Fig 5b) were too extreme tomake a comparison between management systems mean-ingful Within the focal countries males accounted fora larger part of the adult harvest in non-traditional thantraditional deer hunting areas in all countries withharvest composition tending to be more variable fromyear-to-year in non-traditional areas (see Appendix S4in the supplementary material)

Discussion

Harvesting has a fundamental influence on the popu-lation dynamics of ungulates often accounting for themajority of mortality (Langvatn amp Loison 1999) andincreasing adult female mortality the vital rate withthe highest elasticity (ie the factor in which a givenvariation affects dynamics the most Gaillard Festa-Bianchet amp Yoccoz 1998) However in contrast with

the huge body of theoretical and empirical literatureof other key processes such as density dependenceand climate (Fowler 1987 Sinclair 1989 Lande Engenamp Saeligther 2003 Turchin 2003) there are extremely fewempirical studies addressing how different harvestingstrategies have affected the development and com-position of ungulate populations (Gordon Hester ampFesta-Bianchet 2004 but see Fryxell et al 1991 Lang-vatn amp Loison 1999 Solberg et al 1999 McCullough2001) As a preliminary step towards addressing thiswe have shown the extent to which red deer harvests haveincreased across Europe as a result of both biologicaland cultural factors including increasing deer rangeand hunter numbers in at least some countries (P3)Harvest growth rates and harvest composition differedbetween management systems across Europe Althoughvariation in the harvest of adult females explainedpart of this (P2) differences were not related to huntingobjective in terms of meat vs trophies (P4) and wereonly partially related to differences in harvest growthrates between traditional and non-traditional areas (P1)While harvest growth rates do not reflect populationgrowth rates perfectly the continued growth and apparentsustainability of the harvests over time is indicativeof steadily growing red deer populations in manyEuropean countries despite differences in the culturalaspects of management Such increases are indicative

Fig 5 Changes in harvest composition in terms of (a) the proportion of calves in the total harvest and (b) the proportion of adult males in the total adultharvest Yearlings are included with adults in all cases For management systems indicated by numbered labels refer to Table 1



of an underestimation of population sizes (Csaacutenyiamp Toacuteth 2000 Gaillard Loison amp Toiumlgo 2003) andorunderharvesting particularly of females (Clutton-Brock Coulson amp Milner 2004 Giles amp Findlay 2004)but appear to be independent of the red deer manage-ment system

Across Europe harvest growth rates of red deer werehighest in management systems where the harvest wascontrolled by licence and land or hunting groundownership (management system 2 Fig 1) Differencesbetween systems are more likely to reflect differencesbetween groups of countries with similar huntingcultures and general approaches to management thanto be a direct result of the system for controlling harvest-ing The intensity of harvesting ie harvesting rate islikely to strongly influence harvest growth and maydiffer between management systems but in the absenceof population data it is not possible to identify the extentto which this occurred Based on available data harvestcomposition was partly responsible for the higher harvestgrowth of management system 2 countries (althoughNorway harvested more males than Denmark Franceand Sweden) but these countries also showed a stronghunting culture indicated by a high percentage ofinhabitants who were hunters Hence harvesting ratesmay be higher and management more focused towardssatisfying huntersrsquo demands For example managementpractises such as introducing new deer populations inFrance in the 1950ndash60s (Leduc amp Klein 2004) and takinga male-biased harvest in Norway (Langvatn amp Loison1999) were taken to increase population size and hencehunting opportunities in low-density areas

Harvest composition is strongly influenced by culturalfactors including management strategies huntingtraditions and economic considerations as well as bybiological factors Changes in calf harvesting strategyduring the study period were at least partly the result oflegislative and management policy changes In terms ofthe adult harvest this was female-biased in all countriesexcept Norway Contrary to prediction P4 the propor-tion of males in the harvest was unrelated to trophyhunting objectives unlike the harvesting of manyAfrican ungulates (Ginsberg amp Milner-Gulland 1994)As is typical for Scandinavia meat rather than trophieswas the major hunting objective in Norway yet theharvest was more male-biased than in many primarilytrophy-hunting countries While large males may betargeted by meat hunters for their higher yield thisdoes not account for the female bias in other countriesHowever trophy size is strongly influenced by age (Kruuket al 2002) so trophy hunters tend to be reluctant toshoot young males which make up most of the malepopulation Consequently relatively few animals are shotas trophies even in areas where trophy hunting is themost economically important management objective

For example in Scotland the sporting cull accounts foronly 25 of the total harvest (Reynolds amp Staines 1997)In addition managers may harvest males suboptimallyto avoid overharvesting (Milner-Gulland Coulson ampClutton-Brock 2004) Good-quality trophies are gen-erally achieved at lower population densities requiringfemale numbers to be controlled As a result secondarymanagement objectives such as meat yield or popula-tion control often account for the greater proportionof the total harvest Furthermore many populations ofdimorphic ungulates are female-biased (Clutton-Brockamp Lonergan 1994) so despite near parity in the harvesta greater proportion of the males in the population maynone the less be harvested Indeed in Scotland wherethe total red deer harvest is female-biased approximately16 of males compared with 14 of females wereharvested during the 1990s (Deer Commission forScotland unpublished data)

Deer and hunter populations are both dynamic entitiesthat are constantly interacting (Brown et al 2000)Feedback between cultural factors hunting traditionsand biological processes are likely to be involved in thedevelopment of game populations For example hunt-ing culture (including a mixture of facts and beliefs)will influence size and composition of yield which willaffect population growth rates and consequently man-agement decisions (Fryxell et al 1991 Cocircteacute et al 2004Gordon Hester amp Festa-Bianchet 2004) These may inturn feed back into hunting culture In addition thehunting methods allowed will influence the ability toharvest selectively (Martinez et al 2005) For examplein Hungary driven-shooting of male cervids is prohib-ited by law in order to ensure that the goals of selectivemale harvesting will not be jeopardized

In North America with declining numbers ofhunters and growing deer populations it is now ques-tionable whether sport hunting can reliably controldeer numbers (McShea Underwood amp Rappole 1997Brown et al 2000 Cocircteacute et al 2004) A similar situationis likely in parts of Europe such as France although inNorway and Hungary hunter numbers have risen andit may be that increased availability of game has stim-ulated the recruitment of new hunters

Deer harvests have risen both as a result of increasedharvest densities (reflecting population densities) withinexisting ranges and as a result of considerable expan-sion of the range of red deer throughout Europe (Gill1990) Consequently national harvests hide muchinteresting spatial variation Within the focal countriesthere has been extensive range expansion around manydiscrete introduced populations in France (Leduc ampKlein 2004) and in Alpine areas while in Norway there

732J M Milner et al


has been a primarily southerly and easterly expansionsince 1920 (Langvatn amp Albon 1986 Mysterud et al2002) The establishment of commercial forestry plan-tations in Hungary UK and western parts of Norwayhas also allowed red deer to move into new areas(Clutton-Brock amp Albon 1989 Csaacutenyi 1999 Fuller amp Gill2001) However the influence of deer range expansionon total harvest size was much greater in France andScotland than in Hungary and Norway Environmen-tal constraints such as snow depth or the duration ofsnow cover in winter may limit the rate of colonizationof inland ranges in Norway In addition hunting innew areas of Norway often starts as soon as red deerestablish because of fears of agricultural or forest dam-age which may severely restrict population growth Incontrast in France there is frequently a considerabletime lag between population establishment and thestart of hunting (Bonenfant amp Klein 2004) Much ofthe spatial variation in France and Scotland wasassociated with differences in harvest growth betweentraditional and non-traditional areas (P1) Particularlylarge increases in harvest size occurred in non-traditionalareas at the edges of the deer distribution (Fig 4) Inthese areas harvest growth rates were often higher thanthe maximum possible population growth rate for reddeer assuming a finite rate of population increase(λ) for red deer of 1middot25ndash1middot35 (Gaillard et al 2000) It isunlikely that dispersal could account for all of this(Clutton-Brock et al 2002) suggesting that culturalfactors associated with the expansion of hunting inter-est or improvements in hunting competence must havebeen involved as well as the predicted biologicalfactors related to rapid growth during colonization Inthe traditional hunting areas of France and Scotland itis unclear whether the lower rate of increase in harvestsize reflected a slower increase in deer numbers as aresult of either density-dependent factors or differ-ences in harvest off-take However in Norway wherepopulation density is generally well below carryingcapacity (Mysterud et al 2001) and in Hungary wherepopulation growth rate is almost at the maximumpossible (Csaacutenyi 1992) harvest growth rates weresimilar in traditional and non-traditional areas Alter-natively in traditional areas in France and Scotlandthere may be a greater lag in keeping abreast of popu-lation growth For example in Scotland the size of thecull is often set on the basis of what it has always been(Buckland et al 1996) and a tendency to target thefemale cull towards old rather than the most productivehinds (Trenkel et al 1998) could exacerbate problemsof population control

Red deer numbers have increased considerably through-out Europe over the last 30 years regardless of manage-ment system and cultural hunting traditions Our findingssuggest that deer population growth has been fasterthan harvest growth Changes in harvest size and

composition coincided with changes in managementpolicies hunter numbers and red deer range Furthershifts can be expected in the future as the social impactsof high deer numbers become increasingly felt (Fulleramp Gill 2001 Cocircteacute et al 2004 Gordon Hester amp Festa-Bianchet 2004) and as an awareness of the need forpopulation control develops (Gordon Hester amp Festa-Bianchet 2004) The increase in calf harvesting is anindicator that control efforts are being made in manycountries but changing the adult harvest is moreproblematic because hunting traditions and economicconsiderations often conflict with population controlobjectives

An important challenge for the future managementof red deer in Europe is to improve the availability andquality of population data on which managementdecisions are based Despite being one of Europersquoslargest and most economically important mammalswith a wide distribution and a probable population sizeof 1ndash2 million animals sound data on red deer popula-tions and their dynamics are patchy scarce or inconsistentAgainst this background harvesting must balancechanging hunting requirements and capabilities withthe increasing social and economic impacts of deer andother ungulates

Acknowledgements

We thank all the agencies who provided harvest statisticsdata and Kamil Barton Juan Carranza Petter KjellanderSandro Lovari Michael Petrak Karoline Schmidt andSilvano Toso for helping locate data In particular wethank ONCFS SSB and DCS for digital maps andfurther assistance Mike Daniels Franccedilois KleinKaroline Schmidt Erling Solberg and Vebjoslashrn Veibergprovided additional help information and discussionand Nigel Yoccoz provided statistical advice Financialsupport was from the Norwegian Research Council(NFR 156367530)

References

Ahleacuten I (1975) Winter habitats of moose and deer in relationto land use in Scandinavia Swedish Wildlife Research 945ndash192

Asferg T amp Olesen CR (2004) Danmarks Hjortevildt NaturogMuseum 43(4) Aringrhus Denmark

Ballard WB Whitlaw HA Wakeling BF Brown RL de VosJC amp Wallace MC (2000) Survival of female elk in northernArizona Journal of Wildlife Management 64 500ndash504

Biederbeck HH Boulay MC amp Jackson DH (2001)Effects of hunting regulations on bull elk survival and agestructure Wildlife Society Bulletin 29 1271ndash1277

Bonenfant C amp Klein F (2004) Evolution de la population deCerf (Cervus elaphus L) du Parc National des CeacutevennesReport SGME2004 ONCFS Bar-le-Duc France

Bonenfant C Gaillard J-M Klein F amp Loison A (2002)Sex- and age-dependent effects of population density on lifehistory traits of red deer Cervus elaphus in a temperateforest Ecography 25 446ndash458

Bonenfant C Loe LE Mysterud A Langvatn RStenseth NC Gaillard J-M amp Klein F (2004) Multiple



causes of sexual segregation in European red deer enlighten-ment from varying breeding phenology at high and lowlatitude Proceedings of the Royal Society of London B 271883ndash892

Brown TL Dekker DJ Riley SJ Enck JW Lauber TBCurtis PD amp Mattfield GF (2000) The future of huntingas a mechanism to control white-tailed deer populationsWildlife Society Bulletin 28 797ndash807

Buckland ST Ahmadi S Staines BW Gordon IJ ampYoungson RW (1996) Estimating the minimum popula-tion size that allows a given annual number of mature reddeer stags to be culled sustainably Journal of AppliedEcology 33 118ndash130

Caughley G (1977) Analysis of Vertebrate PopulationsWiley Chichester UK

Clutton-Brock TH amp Albon SD (1989) Red Deer in theHighlands BSP Professional Books Oxford UK

Clutton-Brock TH amp Albon SD (1992) Trial and error inthe Highlands Nature 358 11ndash12

Clutton-Brock TH amp Lonergan ME (1994) Cullingregimes and sex ratio biases in highland red deer Journal ofApplied Ecology 31 521ndash527

Clutton-Brock TH Coulson T amp Milner JM (2004)Red deer stocks in the Highlands of Scotland Nature 429261ndash262

Clutton-Brock TH Coulson TN Milner-Gulland EJThomson D amp Armstrong HM (2002) Sex differences inemigration and mortality affects optimal management ofdeer populations Nature 415 633ndash637

Coltman DW OrsquoDonoghue P Jorgensen JT Hogg JTStrobeck C amp Festa-Bianchet M (2003) Undesirableevolutionary consequences of trophy hunting Nature 426655ndash658

Cocircteacute SD Rooney TP Trembley J-P Dussault C ampWaller DM (2004) Ecological impacts of deer overabundanceAnnual Review of Ecology and Systematics 35 113ndash147

Csaacutenyi S (1992) Red deer population dynamics in Hungarymanagement statistics versus modeling The Biology of Deer(ed RD Brown) pp 37ndash42 Springer-Verlag New York NY

Csaacutenyi S (1999) A giacutemszarvasaacutellomaacuteny terjeszkedeacutese azalfoumlldoumln [Expansion of red deer in the Hungarian GreatPlain] Vadbioloacutegia 6 43ndash48 [in Hungarian with Englishsummary]

Csaacutenyi S amp Toacuteth P (2000) Populaacutecioacute-rekonstrukcioacutealkalmazaacutesa a hazai giacutemszarvas aacutellomaacuteny leacutetszaacutemaacutenakmeghataacuterozaacutesaacutera [Population reconstruction as a tools toestimate the past population size of red deer in Hungary]Vadbioloacutegia 7 27ndash37 [in Hungarian with English summary]

Diggle P Heagerty P Liang K-Y amp Zeger S (2002)Analysis of Longitudinal Data Oxford University PressNew York NY

FACE (2004) Census of the Number of Hunters in EuropeFederation of Associations for Hunting and Conservationof the EU Brussels Belgium httpwwwface-europeorgfs-huntinghtm (15042004)

Forchhammer M Stenseth NC Post E amp Langvatn R(1998) Population dynamics of Norwegian red deerdensity-dependence and climatic variation Proceedings ofthe Royal Society of London B 265 341ndash350

Fowler CW (1987) A review of density dependence in popula-tions of large mammals Current Mammalogy 1 401ndash441

Fryxell JM Hussell DJT Lambert AB amp Smith PC(1991) Time lags and population fluctuations in white-tailed deer Journal of Wildlife Management 55 377ndash385

Fuller RJ amp Gill RMA (2001) Ecological impacts ofincreasing numbers of deer in British woodland Forestry74 193ndash199

Gaillard J-M Festa-Bianchet M amp Yoccoz NG (1998)Population dynamics of large herbivores variable recruitmentwith constant adult survival Trends in Ecology and Evolution13 58ndash63

Gaillard J-M Festa-Bianchet M Yoccoz NG Loison Aamp Toigo C (2000) Temporal variation in fitness componentsand population dynamics of large herbivores Annual Reviewof Ecology and Systematics 31 367ndash393

Gaillard J-M Loison A amp Toiumlgo C (2003) Variation in lifehistory traits and realistic population models for wildlifemanagement the case of ungulates Animal Behavior andWildlife Conservation (eds M Festa-Bianchet amp M Apollonio)pp 115ndash132 Island Press Washington DC

Giles BG amp Findlay CS (2004) Effectiveness of a selectiveharvest system in regulating deer populations in OntarioJournal of Wildlife Management 68 266ndash277

Gill R (1990) Monitoring the Status of European and NorthAmerican Cervids The Global Environment MonitoringSystem Information Series 8 United Nations EnvironmentProgramme Nairobi Kenya

Ginsberg JR amp Milner-Gulland EJ (1994) Sex-biasedharvesting and population dynamics in ungulates implica-tions for conservation and sustainable use ConservationBiology 8 157ndash166

Gordon IJ Hester AJ amp Festa-Bianchet M (2004) Themanagement of wild large herbivores to meet economicconservation and environmental objectives Journal of AppliedEcology 41 1021ndash1031

Jedrzejewski W Schmidt K Theuerkauf J JedrzejewskaB Selva N Zub K amp Szymura L (2002) Kill rates andpredation by wolves on ungulate populations in Bialowiezaprimeval forest Ecology 83 1341ndash1356

Koubek P amp Zima J (1999) Cervus elaphus The Atlas ofEuropean Mammals (eds AJ Mitchell-Jones G Amori WBogdanowicz B Kryszligtufek PJH Reijnders F Spitzen-berger M Stubbe JBM Thissen V Vohralik amp J Zima)pp 388ndash389 Academic Press London UK

Kruuk LEB Slate J Pemberton JM Brotherstone SGuinness F amp Clutton-Brock T (2002) Antler size in reddeer heritability and selection but no evolution Evolution56 1683ndash1695

Lande R Engen S amp Saeligther B-E (2003) StochasticPopulation Dynamics in Ecology and Conservation OxfordUniversity Press Oxford UK

Langvatn R amp Albon SD (1986) Geographic clines in bodyweight of Norwegian red deer a novel explanation ofBergmannrsquos rule Holarctic Ecology 9 285ndash293

Langvatn R amp Loison A (1999) Consequences of harvestingon age structure sex ratio and population dynamics of reddeer Cervus elaphus in central Norway Wildlife Biology 5213ndash223

Langvatn R Albon SD Burkey T amp Clutton-Brock TH(1996) Climate plant phenology and variation in age of firstreproduction in a temperate herbivore Journal of AnimalEcology 65 653ndash670

Langvatn R Mysterud A Stenseth NC amp Yoccoz NG(2004) Timing and synchrony of ovulation in red deerconstrained by short northern summers American Naturalist163 763ndash772

Leduc D amp Klein F (2004) Lrsquoorigine du cerfs franccedilais de1900 agrave nos jours Faune Sauvage 264 27ndash29

Liang KY amp Zeger SL (1986) Longitudinal data-analysisusing generalized linear-models Biometrika 73 13ndash22

Loe LE Bonenfant C Mysterud A Gaillard J-MLangvatn R Stenseth NC Klein F Calenge CErgon T amp Pettorelli N (2005) Climate predictability andbreeding phenology in red deer timing and synchronyof rutting and calving in Norway and France Journal ofAnimal Ecology 74 579ndash588

McCorquodale SM Wiseman R amp Marcum CL (2003)Survival and harvest vulnerability of elk in the Cascade Rangeof Washington Journal of Wildlife Management 67 248ndash257

McCullough DR (2001) Population manipulations ofNorth American deer Odocoileus spp balancing high yieldwith sustainability Wildlife Biology 7 161ndash170

734J M Milner et al


McShea WJ Underwood HB amp Rappole JH (1997) TheScience of Overabundance Deer Ecology and PopulationManagement Smithsonian Institute Press WashingtonDC

Martinez M Vigal CR Jones OR Coulson T ampSan Miguel A (2005) Different hunting strategies selectfor different weights in red deer Biology Letters 1 353ndash356

Mattioli S Meneguzo PG Brugnoli A amp Nicoloso S(2001) Red deer in Italy recent changes in range and numbersHystrix Italian Journal of Mammalogy 12 27ndash35

Milner JM Alexander JS amp Griffin AM (2002) A HighlandDeer Herd and its Habitat Red Lion House London UK

Milner-Gulland EJ Coulson T amp Clutton-Brock TH(2004) Sex differences and data quality as determinants ofincome from hunting red deer Cervus elaphus WildlifeBiology 10 187ndash182

Mysterud A Langvatn R Yoccoz NG amp Stenseth NC(2002) Large-scale habitat variability delayed density effectsand red deer populations in Norway Journal of AnimalEcology 71 569ndash580

Mysterud A Stenseth NC Yoccoz NG Ottersen Gamp Langvatn R (2003) The response of the terrestrialecosystems to climate variability associated with theNorth Atlantic Oscillation The North Atlantic OscillationsClimatic Significance and Environmental Impact (edsJW Hurrell A Belgrano G Ottersen amp Y Kushnir)pp 235ndash262 American Geophysical Union WashingtonDC

Mysterud A Yoccoz NG Stenseth NC amp Langvatn R(2001) Effects of age sex and density on body weight ofNorwegian red deer evidence of density-dependentsenescence Proceedings of the Royal Society London B 268911ndash919

OrsquoGara BW (2002) Hunting red deer and elk old and newworlds North American Elk Ecology and Management (edsDE Toweill amp JW Thomas) pp 649ndash699 WildlifeManagement Institute Smithsonian Institution PressWashington DC

Peek JM Schmidt KT Dorrance MJ amp Smith BL(2002) Supplemental feeding and farming of elk NorthAmerican Elk Ecology and Management (eds DE Toweillamp JW Thomas) pp 617ndash647 Wildlife ManagementInstitute Smithsonian Institution Press Washington DC

Pinheiro JC amp Bates DM (2000) Mixed-Effects Models inS and S-PLUS Springer-Verlag New York NY

R Development Core Team (2004) R A Language andEnvironment for Statistical Computing R Foundation forStatistical Computing Vienna Austria

Reynolds P amp Staines B (1997) Deer management inScotland Conservation and the Use of Wildlife Resources(ed M Bolton) pp 171ndash198 Chapman amp Hall LondonUK

Sinclair ARE (1989) Population regulation in animalsEcological Concepts (ed JM Cherrett) pp 197ndash241Blackwell Science Oxford UK

SNH (1994) Red Deer and the Natural Heritage ScottishNatural Heritage Battleby UK

Sokal RR amp Rohlf FJ (1981) Biometry WH FreemanNew York NY

Solberg EJ Saeligther B-E Strand O amp Loison A (1999)Dynamics of a harvested moose population in a variableenvironment Journal of Animal Ecology 68 186ndash204

Trenkel VM Partridge LW Gordon IJ Buckland STElston DA amp McLean C (1998) The management of reddeer on Scottish open hills results of a survey conducted in1995 Scottish Geographical Magazine 114 57ndash62

Turchin P (2003) Complex Population Dynamics A TheoreticalEmpirical Synthesis Princeton University Press PrincetonNJ

Venables WN amp Ripley BD (1999) Modern Applied Statistieswith S-PLUS Springer New York

Received 13 January 2006 final copy received 14 March 2006Editor E J Milner-Gulland

Supplementary material

The following supplementary material is availableas part of the online article (full text) from httpwwwblackwell-synergycom

Appendix S1 Harvest management in the focal countries

Appendix S2 Changes in harvest growth rate over timein the 11 study countries

Appendix S3 GLS models of harvest size in relationto hunter numbers and red deer range area in the focalcountries

Appendix S4 Harvest composition within the focalcountries

Figure S2 Harvest growth rate over time in the 11study countries

Figure S4a The proportion of calves in the harvestsof traditional and non-traditional areas in the focalcountries

Figure S4b The proportion of males in the adultharvests of traditional and non-traditional areas in thefocal countries

Table S2 GLS model of harvest growth rate

722

J M Milner

et al



43

721ndash734

Introduction

The numbers of many deer species have increaseddramatically throughout Europe and North Americaover the last century (Gill 1990 Clutton-Brock amp Albon1992 McShea Underwood amp Rappole 1997 Cocircteacute

et al

2004 Gordon Hester amp Festa-Bianchet 2004) as a resultof both increases in density and range expansion Thereare a number of causes both direct and indirect with nosingle explanation applicable to all countries or species(Fuller amp Gill 2001) Factors unrelated to cervid manage-ment include increased availability of forage resultingfrom changes in land use (Ahleacuten 1975) agricultural andsilvicultural practises (Mysterud

et al

2002) and theclimate (Mysterud

et al

2003) However the directmanagement of cervids has also played a part Forexample protection from overhunting and poachingthe use of selective harvesting regimes (Langvatn amp Loison1999 Solberg

et al

1999) supplementary feeding inwinter (Peek

et al

2002) and reintroduction programmes(McShea Underwood amp Rappole 1997 Mattioli

et al

2001 Leduc amp Klein 2004) have all contributed to risingnumbers As a result in many areas management objec-tives are beginning to change from species protectionto population control (Buckland

et al

1996 Brown

et al

2000 Cocircteacute

et al

2004 Gordon Hester amp Festa-Bianchet 2004)

Red deer

Cervus elaphus

L are widely distributed acrossEurope (Fig 1 Koubek amp Zima 1999) and have increasedin both abundance and geographical range in recentdecades (Gill 1990 Csaacutenyi 1992 Mattioli

et al

2001Clutton-Brock Coulson amp Milner 2004) althoughreliable quantitative data on population sizes aregenerally lacking Red deer are primarily found inforest or woodland-edge habitats often associated withland of low productivity including open moorland inScotland The widespread distribution and large bodysize of red deer makes it an important game species(Koubek amp Zima 1999) Hunting rights in Europe aregenerally held by the landowner (Gill 1990) in contrastto North America where hunting is accessible to a widesector of the general public (OrsquoGara 2002) Consequentlyhunting in Europe tends to be limited to landowners(state or private) andor their associates and payingleaseholders or trophy-hunting clients The size of reddeer harvests has increased considerably in manyEuropean countries over recent decades (Gill 1990)

Harvesting represents a major source of mortality inmany North American elk

Cervus elaphus

L populationswhere predators are absent (Ballard

et al

2000 Bieder-beck Boulay amp Jackson 2001 McCorquodale Wisemanamp Marcum 2003) While few empirical studies havedocumented this in Europe it is also likely to be true formany red deer populations (Langvatn amp Loison 1999)although predation may play a more significant rolein some protected areas (Jedrzejewski

et al

2002) Con-sequently selective harvesting is likely to influencestrongly population composition dynamics (Langvatnamp Loison 1999 for moose Alces alces see Solberg

et al

1999) and genetics (Coltman

et al

2003) At the Euro-pean scale the basis for red deer management variesdepending on cultural traditions and the environmentalconstraints of habitat and latitude (Bonenfant

et al

2004Loe

et al

2005) We might therefore expect harvest ratesselectivity and composition to differ between countrieswith consequences for population dynamics Howeverit is currently not possible to address empirically howregional variation in selective harvesting regimes hasaffected population dynamics as the required data simplydo not exist for most red deer populations In the absenceof such data we aimed to describe the main patterns inthe size of the red deer harvest over the last three to fourdecades in 11 European countries with contrasting hunt-ing objectives management strategies and environmentalconditions In addition we investigated trends in the com-position of the harvest in seven of the countries and madea more in-depth analysis of spatial variation in fourfocal countries France Hungary Norway and Scotland

Harvest development is likely to follow general trendsin population development (Forchhammer

et al

1998and references therein) although often with a 1ndash2-yeartime-lag (Fryxell

et al

1991 Solberg

et al

1999) Harvestsize is also influenced by policy and cultural factorsincluding harvesting strategies (Giles amp Findlay 2004)and quotas which explain some of the time lags(Solberg

et al

1999) We therefore also investigatedthe following predictions (i) P1 harvest growth rateswill be highest where deer populations are growingfastest Consequently we expect harvest growth ratesto be lower in traditional deer hunting areas than inareas of recent colonization where lower densities andthus higher recruitment rates are likely (unless habitatdiffers markedly) (ii) P2 harvest growth rates will belower with a higher proportion of adult females in theharvest because of the sensitivity of population growthto adult female survival (Gaillard Festa-Bianchet ampYoccoz 1998) assuming a close link between popu-lation growth and harvest growth (iii) P3 there will bepositive covariation between harvest size hunternumbers and size of red deer range (iv) P4 a higherproportion of adult males relative to females will occurin the harvest in trophy-hunting cultures (Ginsberg ampMilner-Gulland 1994)

Materials and methods

Annual statistics for the size of the total red deer harvestwere obtained for Austria Denmark France GermanyHungary Norway Poland Scotland Slovenia Swedenand Switzerland from national bodies in the respectivecountries (Table 1) The year of harvest was taken fromthe start of the hunting season Harvest compositionin terms of the number of calves adult (includingyearling) males and females shot were available forseven countries (Table 1) Data describing populationsize and structure were unfortunately unavailable formost of the countries studied However as our investiga-tion was of variation at a continental scale we assumed

723




43

721ndash734





724

J M M

ilner

et al

copy 2006 T

he Authors

Journal compilation

copy 2006 B

ritish E

cological Society

Journal of Applied

Ecology

43

721

ndash734

Table 1



Hunting objective




Harvest growth rate

H

r

(SD)Probability

H

r

gt 0





1965ndash2002 (1965ndash2002)
















1965ndash2002 (1965ndash2002)




1969ndash2002 (1969ndash2002)





und Landschaft


2


plusmn


H

r


P


725




43

721ndash734



Harvest growth rate


H

r


n

(

h

t+

1

)

minus

log

n

(

h

t

) where

h

t


t


H

r


H

r


n

(

h

t

+1

+ 1)

minus

log

n

(

h

t



For each country

t


H

r



P


T


P


H

r

H

r



H

r




Differences in

H

r


726

J M Milner

et al



43

721ndash734


P


P


P


P


H

r


Harvest composition


et al



z


k


k


Results



727




43

721ndash734


t


F

1365

= 1middot653

P



F

10356

= 1middot55

P


F

1365

=9middot72

P


F

110

= 7middot86

P



F

3220

= 0middot367

P






728J M Milner et al




Calf harvest





















Adult harvest



730J M Milner et al




Discussion













732J M Milner et al






Acknowledgements


References


















































734J M Milner et al

































723




43

721ndash734





724

J M M

ilner

et al

copy 2006 T

he Authors

Journal compilation

copy 2006 B

ritish E

cological Society

Journal of Applied

Ecology

43

721

ndash734

Table 1



Hunting objective




Harvest growth rate

H

r

(SD)Probability

H

r

gt 0





1965ndash2002 (1965ndash2002)
















1965ndash2002 (1965ndash2002)




1969ndash2002 (1969ndash2002)





und Landschaft


2


plusmn


H

r


P


725




43

721ndash734



Harvest growth rate


H

r


n

(

h

t+

1

)

minus

log

n

(

h

t

) where

h

t


t


H

r


H

r


n

(

h

t

+1

+ 1)

minus

log

n

(

h

t



For each country

t


H

r



P


T


P


H

r

H

r



H

r




Differences in

H

r


726

J M Milner

et al



43

721ndash734


P


P


P


P


H

r


Harvest composition


et al



z


k


k


Results



727




43

721ndash734


t


F

1365

= 1middot653

P



F

10356

= 1middot55

P


F

1365

=9middot72

P


F

110

= 7middot86

P



F

3220

= 0middot367

P






728J M Milner et al




Calf harvest





















Adult harvest



730J M Milner et al




Discussion













732J M Milner et al






Acknowledgements


References


















































734J M Milner et al

































724

J M M

ilner

et al

copy 2006 T

he Authors

Journal compilation

copy 2006 B

ritish E

cological Society

Journal of Applied

Ecology

43

721

ndash734

Table 1



Hunting objective




Harvest growth rate

H

r

(SD)Probability

H

r

gt 0





1965ndash2002 (1965ndash2002)
















1965ndash2002 (1965ndash2002)




1969ndash2002 (1969ndash2002)





und Landschaft


2


plusmn


H

r


P


725




43

721ndash734



Harvest growth rate


H

r


n

(

h

t+

1

)

minus

log

n

(

h

t

) where

h

t


t


H

r


H

r


n

(

h

t

+1

+ 1)

minus

log

n

(

h

t



For each country

t


H

r



P


T


P


H

r

H

r



H

r




Differences in

H

r


726

J M Milner

et al



43

721ndash734


P


P


P


P


H

r


Harvest composition


et al



z


k


k


Results



727




43

721ndash734


t


F

1365

= 1middot653

P



F

10356

= 1middot55

P


F

1365

=9middot72

P


F

110

= 7middot86

P



F

3220

= 0middot367

P






728J M Milner et al




Calf harvest





















Adult harvest



730J M Milner et al




Discussion













732J M Milner et al






Acknowledgements


References


















































734J M Milner et al

































725




43

721ndash734



Harvest growth rate


H

r


n

(

h

t+

1

)

minus

log

n

(

h

t

) where

h

t


t


H

r


H

r


n

(

h

t

+1

+ 1)

minus

log

n

(

h

t



For each country

t


H

r



P


T


P


H

r

H

r



H

r




Differences in

H

r


726

J M Milner

et al



43

721ndash734


P


P


P


P


H

r


Harvest composition


et al



z


k


k


Results



727




43

721ndash734


t


F

1365

= 1middot653

P



F

10356

= 1middot55

P


F

1365

=9middot72

P


F

110

= 7middot86

P



F

3220

= 0middot367

P






728J M Milner et al




Calf harvest





















Adult harvest



730J M Milner et al




Discussion













732J M Milner et al






Acknowledgements


References


















































734J M Milner et al

































726

J M Milner

et al



43

721ndash734


P


P


P


P


H

r


Harvest composition


et al



z


k


k


Results



727




43

721ndash734


t


F

1365

= 1middot653

P



F

10356

= 1middot55

P


F

1365

=9middot72

P


F

110

= 7middot86

P



F

3220

= 0middot367

P






728J M Milner et al




Calf harvest





















Adult harvest



730J M Milner et al




Discussion













732J M Milner et al






Acknowledgements


References


















































734J M Milner et al

































727




43

721ndash734


t


F

1365

= 1middot653

P



F

10356

= 1middot55

P


F

1365

=9middot72

P


F

110

= 7middot86

P



F

3220

= 0middot367

P






728J M Milner et al




Calf harvest





















Adult harvest



730J M Milner et al




Discussion













732J M Milner et al






Acknowledgements


References


















































734J M Milner et al

































728J M Milner et al




Calf harvest





















Adult harvest



730J M Milner et al




Discussion













732J M Milner et al






Acknowledgements


References


















































734J M Milner et al





































Adult harvest



730J M Milner et al




Discussion













732J M Milner et al






Acknowledgements


References


















































734J M Milner et al

































730J M Milner et al




Discussion













732J M Milner et al






Acknowledgements


References


















































734J M Milner et al










































732J M Milner et al






Acknowledgements


References


















































734J M Milner et al











































































734J M Milner et al

































Temporal and spatial development of red deer harvesting in Europe: biological and cultural factors

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Transcript of Temporal and spatial development of red deer harvesting in Europe: biological and cultural factors