Total red deer harvests had increased by 400–700% in most countries over the last 30 years. Exponential increases occurred in eight out of the 11 countries examined. The exceptions were Austria and Germany, which had stable harvests, and Poland, where a strong increase had been followed by a recent decline (Fig. 2). Harvest growth rates varied from 0·009 in Austria, where the annual harvest fluctuated between 31 000 and 46 000, to 0·075 in Sweden, which had the smallest harvest, rising from around 90 red deer shot in 1965 to 1420 in 2002. However, harvest growth rates were only significantly greater than 0 in France and Norway after making a conservative Bonferroni correction for multiple t-tests (Table 1). During the study period, we found no evidence of a change in harvest growth over time in any country (F1,365 = 1·653, P= 0·199; see Appendix S2 in the supplementary material), suggesting that deer populations were growing at a faster rate than the harvests.
Figure 2. Trends in size of the (ln) total red deer harvests in 11 European countries since 1965. For management systems, indicated by numbered labels, refer to Table 1.
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Differences in national harvest growth rates were not significant (F10,356 = 1·55, P= 0·12) because of high variability over time within countries. However, harvest growth rates differed significantly between management systems, being higher in countries where the harvest was controlled by licence and land or hunting ground ownership (management system 2; Fig. 1 and Table 1) than in countries with other management systems (F1,365 = 9·72, P= 0·002). Denmark, France, Norway and Sweden were also the countries with the highest percentage of inhabitants who were hunters in 2004 (average 3·5% of the population compared with 0·9%). Indeed, the percentage of hunters in the population explained nearly half of the variation in harvest growth rate between countries (slope = 0·009, F1,10 = 7·86, P= 0·021; Fig. 3a).
Figure 3. Relationship between (a) harvest growth rate and the percentage of inhabitants that were registered hunters in 2004 in 11 European countries, (b) harvest size and hunter numbers in the focal countries and (c) harvest size and deer range area in the focal countries (note the log-scaled x-axis for b and c).
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In support of prediction P2, harvest growth rate was significantly negatively influenced by the proportion of females in the adult harvest at the national level (Table 2a). Once the proportion of females in the adult harvest was fitted, the management system explained no additional variation (F3,220 = 0·367, P= 0·777), suggesting that this term accounted for much of the difference in harvest growth rate between management systems.
Table 2. The effects of the proportion of females (PropF) in the adult harvest on the annual rate of increase in harvest in (a) the national harvest in seven countries (Austria, France, Hungary, Norway, Scotland, Slovenia and Switzerland) modelled by GLS, and (b) the regional harvest in each focal country together with the effect of hunting tradition and their interaction, modelled with a linear mixed model fitting region as a random effect on the intercept
| || || ||Estimate (SE)||F||d.f.||P||Random effect (95% CI)|
|(a)||National||PropF||−0·284 (0·096)|| 8·779|| 1|| 0·003|| |
|(b)||France||PropF||−0·244 (0·095)|| 6·527||1,1235|| 0·0107||0·521 (0·377–0·719)|
|Tradition*||−0·032 (0·017)|| 3·408|| 1,78|| 0·069|| |
|Hungary||PropF||−1·243 (0·225)||12·01|| 1,544||< 0·001||0·422 (0·204–0·874)|
|Tradition|| 0·013 (0·243)||40·88|| 1,17||< 0·001|| |
|PropF–tradition|| 0·904 (0·432)|| 4·383|| 1,544|| 0·037|| |
|Norway||PropF||−0·032 (0·044)|| 2·467||1,4504|| 0·116||0·297 (0·227–0·388)|
|Tradition|| 0·122 (0·033)||23·21|| 1,212||< 0·001|| |
|PropF–tradition||−0·166 (0·081)|| 4·167||1,4504|| 0·041|| |
|Scotland||PropF||−0·112 (0·139)||16·95||1,1632||< 0·001||0·403 (0·242–0·671)|
|Tradition|| 0·177 (0·100)|| 5·014|| 1,55|| 0·029|| |
|PropF–tradition||−0·436 (0·187)|| 5·414||1,1632|| 0·020|| |
Within the focal countries there was, as predicted (P3), a positive relationship between ln(harvest size) and red deer range in all countries (Fig. 3c; r2 = 0·82–0·97; see Appendix S3 in the supplementary material) and with hunter numbers in Hungary and Norway (Fig. 3b; r2 = 0·83–0·95). However, harvest size in France increased despite a decline in hunter numbers (Fig. 3b; r2 = −0·97). Neither hunter numbers (F = 0·008, d.f. = 1, P= 0·931) nor range size (F = 0·244, d.f. = 1, P= 0·622) had any effect on harvest growth rates.
There was considerable spatial variation in harvest growth rates within focal countries (Table 3 and Fig. 4). The highest Hr values tended to occur in areas of recent colonization at the periphery of the red deer range (Fig. 4). The high variance between areas in France and Scotland was partly associated with a pronounced difference in regional harvest growth rates between traditional deer hunting areas and areas into which deer and deer hunting have expanded, being significantly faster in these non-traditional areas (Table 3). In contrast, there was no difference in the harvest growth rate between traditional and non-traditional areas in Hungary and Norway (Table 3). Thus prediction P1 was only partially supported. The variability in harvest growth rate was greater in non-traditional than traditional areas in all countries (Table 3). In traditional areas, the harvest growth rate reflected an increase in harvest density (i.e. harvest per unit area) within regions, while in non-traditional areas it reflected a combination of both an increase in harvest density and an increase in the number of regions in which harvesting was carried out (Table 3). However, while the number of municipalities harvesting red deer for at least 5 years in Norway increased during the study period by 83%, the contribution of these new areas to the total harvest in 2002 was only 13% and, in Hungary, the non-traditional areas contributed only 3·6% to the 2002 harvest. In comparison, in Scotland the number of management blocks increased by 65% and non-traditional areas accounted for 21% of the 2002 harvest, while in France the number of departments in non-traditional areas increased by 25% and their contribution to the total harvest rose from 26% in 1973 to 50% in 2002. The increase in total size of the Hungarian and Norwegian harvests was therefore most influenced by an increase in harvest density in traditional areas, while in France and Scotland range expansion was an important contributory factor.
Table 3. (a) Linear mixed model showing the fixed effects of country and hunting tradition on the change in harvest between years, with region fitted as a random effect. (b) Mean harvest growth rates (Hr) in traditional and non-traditional red deer hunting areas, 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 were included
|(a)|| ||d.f.||F||P|| |
|Intercept|| || 1||347·6||< 0·001|| |
|Country|| || 3|| 4·693|| 0·003|| |
|Tradition|| || 1|| 5·833|| 0·016|| |
|Country–tradition|| || 3|| 3·594|| 0·014|| |
|Residual|| ||10254|| || || |
|(b)|| ||ni||nmax||Hr (SD)||Hr range|
|France||Traditional|| 26|| 29|| 0·062 (0·248)||−0·006–0·111|
|Non-traditional|| 34|| 52|| 0·119 (0·417)***||−0·058–0·418|
|Hungary||Traditional|| 13|| 13|| 0·075 (0·172)|| 0·040–0·110|
|Non-traditional|| 0|| 6|| 0·075 (0·763)||−0·030–0·155|
|Norway||Traditional|| 92|| 92|| 0·061 (0·312)||−0·004–0·141|
|Non-traditional|| 0||122|| 0·064 (0·543)||−0·154–0·260|
|Scotland||Traditional|| 34|| 34|| 0·037 (0·285)**||−0·008–0·155|
|Non-traditional|| 0|| 22|| 0·096 (0·512)||−0·004–0·500|
Figure 4. The spatial variation in the rate of growth in harvest size in (a) France at the department scale, (b) Norway at the municipality scale and (c) Scotland at the scale of management blocks. Traditional hunting areas are outlined in red. Areas with no harvest include those areas where the total number of deer harvested over the study period was less than 50 or where harvesting took place in fewer than 5 years.
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As at the national level, the proportion of females in the adult harvest also had a negative effect on regional Hr in all focal countries, adding further support to prediction P2. However, the relationships differed between traditional and non-traditional areas in all cases except France (Table 2b), being stronger in non-traditional than traditional areas in Hungary but only significant in traditional areas in Norway and Scotland (Table 2b).