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Fish and wildlife in the Swedish mountain region: resources, use and management

Fish and wildlife in the Swedish mountain region: resources, use and management International Journal of Biodiversity Science and Management 2 (2006) 334–342 Fish and wildlife in the Swedish mountain region: resources, use and management 1 2 1,2 3 4 5 T. Eriksson , J. Andersson , P. Byström , M. Hörnell-Willebrand , T. Laitila , C. Sandström and T. Willebrand Department of Aquaculture, Swedish University of Agricultural Sciences, Umeå, Sweden Department of Ecology and Environmental Science, Umeå University, Umeå, Sweden Department of Animal Ecology, Swedish University of Agricultural Sciences, Umeå, Sweden Currently: International Institute for Applied System Analysis (IIASA), Laxenburg, Austria Department of Forest Economics, Swedish University of Agricultural Sciences, Umeå, Sweden Department of Political Science, Umeå University, Umeå, Sweden Key words: Fish, wildlife, Arctic char, willow grouse, ecology, management, property regimes SUMMARY Hunting and fishing are an important part of life for many Swedes. The importance of fish and wildlife is even greater in the northern part of Sweden and especially in the Swedish mountain area, which covers a large geographical area but is sparsely populated. The bio- logical systems in the mountain ecosystems can be characterised as arctic with low produc- tivity and biodiversity. This article addresses two important species typical for this region (willow grouse and Arctic char), provides an up-to-date characterisation of current use and management, and presents a range of management models that frame the mecha- nisms of population dynamics of fish and game. It is concluded that lack of knowledge of population responses of fish and game in the mountain region to different harvesting options and management strategies is a practical obstacle, as is knowledge and under- standing of the costs and benefits associated with different ecological management strategies. However, it is also crucial to integrate other disciplines with ecology for success- ful management. INTRODUCTION Fish and wildlife have significant consumptive and importance of wildlife and freshwater fish has non-consumptive values (Mattsson 1990; Rockel declined over much of Europe, game and fish are and Kealy 1991; Murray and Simcox 2003). Wild- still used for food in many households in the moun- life, especially fish, is an important source of food tain region of Sweden (Ericsson et al. 2005). and cash in many societies. Hunting and fishing are Many communities in the Swedish mountain often central to the perception of the landscape region have faced substantial declines in jobs for people in rural areas (Greider and Garkovich previously generated by forestry, hydro-power and 1994; Steadman 2003). While the consumptive agriculture (Lundmark 2006). The most important Correspondence: T. Eriksson, Department of Aquaculture, Swedish University of Agricultural Sciences, SE-901 83 Umeå, Sweden. Email: torleif.eriksson@vabr.slu.se 334 Fish and wildlife in the Swedish mountain region Eriksson et al. source of employment today is the public sector. 6 kg/ha/year in lakes and from 2 to 15 kg/ha/year The region has also experienced net emigration, in running waters (Hammar 1996). Moose resulting in an imbalanced demographic popula- (Alces alces) and grouse species (willow grouse, rock tion structure. However, increased welfare and ptarmigan (Lagopus mutus), capercaillie (Tetrao leisure time in modern societies are believed to be a urogallus)and blackgrouse (Tetrao tetrix)) are the factor attracting more visitors to the region. A most popular game in the region. Willow grouse are general increase in tourism is therefore seen as a widespread, inhabiting primary arctic tundra, open- possible new source of employment, but this indus- ings in boreal forest, forest edge habitats, and try has not yet grown as quickly as other industries sub-alpine vegetation. Autumn densities range from have declined (Lundmark 2006). 1to50birds per km (Hörnell-Willebrand 2005). Wilderness, fish and wildlife are typical Our two focal species, Arctic char and willow resources of the region, and have led to the explora- grouse, show major differences in their life history tion of new areas for recreational hunting and fish- characteristics as well as in their position in the food ing. To date, however, there is limited knowledge of web. Arctic char and many other target species for the ecology and harvesting of these resources, so fishing are at the top of the food web (Walters and that it is difficult to estimate the potential of devel- Kitchell 2001; Karlsson and Byström 2005). As top oping fishing and hunting tourism. Increased pres- predator, Arctic char have strong effects on lower sure on these resources would certainly also result trophic levels, but are therefore commonly in competition and conflicts over access to them. resource-limited; individual growth rates are Furthermore, hunting and fishing rights are con- strongly resource-dependent (Byström and tested and overlapping between landowners and Andersson 2005; Byström 2006; Finstadt et al. 2006). the Sami, the indigenous people in the mountain In contrast, willow grouse is a typical herbivore region. However, this article does not discuss the specialised in low-digestible browsing during land-use or property rights system (for an overview winter. It does not show such an obvious food limi- of this conflict see Ekenberg 2000; Hahn 2000). tation as Arctic char, although it can be argued that Instead, it focuses on two important systems which food quality influences adult body condition and are typical for this region: willow grouse (Lagopus the subsequent vitality and survival of offspring lagopus) and salmonids (mainly Arctic char (Salve- (Brittas 1984). The proportion of eggs laid in linus alpinus). We give an up-to-date characterisa- spring that survive, and subsequently become tion of current use and management, and present a young grouse in autumn, varies greatly. The com- range of management models that frame the mech- plex fluctuation pattern and population changes anisms of population dynamics of fish and game. can be explained by trophic interactions with We then summarise research results on ecological predators and possibly food (vegetation) quality management strategies and economic values, and (Brittas 1984; Steen et al. 1988; Smith and discuss how these findings can be merged in a more Willebrand 1999) (Figure 1). complete and policy-relevant range of manage- Consequently, harvesting char or grouse will ment options, irrespective of land-use rights. potentially have different effects on population ECOLOGICAL CHARACTERISTICS SETTING THE FRAMEWORK The Swedish mountain region covers about 165,000 km . The area is rich in lakes, with about 61,000 lakes larger than one hectare and a total lake area of 8240 km . There are about 20,000 km of running water (National Board of Fisheries 2004). The species mainly fished are Arctic char (Salvelinus alpinus), whitefish (Coregonus spp.) and brown trout (Salmo trutta) in lakes, and European grayling (Thymallus thymallus) and brown trout in streams. Figure 1 Characteristic population fluctuation of Crude production estimates range from 0.5 to willow grouse at Lövhögen, Jämtland, 1962–1994 International Journal of Biodiversity Science and Management 335 Fish and wildlife in the Swedish mountain region Eriksson et al. dynamics. Arctic char, like fish in general, grow larger lakes, household fisheries in lakes, and recre- continuously throughout life, with the rate of ational fisheries in lakes and streams. Sport fishing growth depending on resource availability and is allowed in 85–90% of the waters in the mountain competition. It therefore takes a substantial num- area. There are no limitations in the number of ber of years for Arctic char (and other fish top pred- licences given in these waters, but a number ators) to reach a size suitable for harvest. This of waters are open only for a restricted number of pattern is even more pronounced in recreational fishing days. Furthermore, private waters, waters fisheries which aim for the largest individuals (Post used by the Sami organised in Reindeer Herding et al. 2002). Target char individuals may range in Communities (RHCs) and some sensitive waters age from 5–6 years to 20 years depending on the sys- are not included. In the waters open for recre- tem and size of individuals (Svenning and ational fishing, there are some limitations, such as Borgström 1995; Finstadt et al. 2006; Byström 2006). a bag limit and a minimum size limit. The presence of size structure in Arctic char, and The RHCs have rights to fish in all waters on fish populations in general, also make the interac- crown land. For all others, a permit is needed for tions between char and other fish species and their commercial or household fisheries with gillnets. resources complex and in many cases unpredict- These permits are personal and give fishing access able when it comes to the effects of harvesting large to one or more lakes. Household fishing has a individuals (Post et al. 2002; Walters and Kitchell limitation on the number of nets allowed. The 2001). Furthermore, even if the char population number of licences has been reduced during the may respond by compensatory increase, due to past 10 years (in Norrbotten county by 60% to increased resource availability for each individual, ~1000 licences, in Västerbotten county by 30% to the fact that char is a long-lived species, with a long ~650 licences). time to reach maturity, means that the compensa- Commercial catches of fish using gillnets have tion will not appear from one year to the next, even continuously decreased over the past 40 years. In though a female char may lay from 100 to more Norrbotten, the landings from the larger lakes have than 1000 eggs (Klemetsen et al. 2003). decreased from 180 tons in the 1960s to about In contrast to char, individuals in the grouse 40 tons in 2000. In recent years, fewer licences for population are similar in their survival and fecun- fishing with gillnets have been sold in Norrbotten dity from their first year. Willow grouse females lay and Västerbotten, while the numbers of sport fish- between 7 and 12 eggs, and have the potential to ing licences sold have remained more or less con- re-nest if the eggs are stolen or eaten. Even first-year stant (Figure 2). Estimated catches have decreased breeders seem to perform as well as older grouse. by approximately 25% over the last 5 years, mainly There is no apparent selection by hunters, and no due to a reduction of gillnet fishing. trophy characters that would potentially change the structure of the population. Harvest adds a mortal- ity source to the many different predators that prey on willow grouse (Smith and Willebrand 1999). Jämtland Norrbotten The complex trophic interaction, which induces an apparent strong stochastic component in the annual increase of the population, makes the response to harvesting difficult to predict (Pedersen et al. 2004; Hörnell-Willebrand 2005). The risk is that the small effects of harvesting might go undetected until a 50000 threshold is reached when the system could collapse (Willebrand and Hörnell 2001). 1960 1965 1970 1975 1980 1985 1990 1995 2000 2005 RIGHTS, RULES, USAGE AND VALUES Year Fishing Figure 2 Number of sport fishing licences sold in The fishing resource is traditionally used by a Jämtland 1960–2003 and Norrbotten 1995–2003 (data number of user groups: commercial fisheries in from County Administrations) 336 International Journal of Biodiversity Science and Management # Licenses sold Fish and wildlife in the Swedish mountain region Eriksson et al. whole: 43% and 13%, respectively. The utilisation Hunting of game meat also seems to play an important social The willow grouse hunting season runs from 25 role. The proportion of households consuming August to the last day of February in the southern game meat on a regular basis is far larger than the part of the mountain range and until 15 March in proportion of households with hunter members. the northern part. Willow grouse are hunted with For instance, in the mountain area, about three out shotguns, principally using pointing dogs; two- of four people live in households where game meat thirds of all hunting takes place during the first is used regularly. 10 days. In 1993, a decision in the parliament per- The fish and wildlife resource in the mountain mitted access to state-managed land (60,000 km ), area adds considerably to the welfare of anglers and and the interest in willow grouse hunting has since hunters in Sweden. For instance, with 50,000 hunt- increased; however, a significant proportion (about ing days for willow grouse and a consumer surplus 30%) of the hunting permits are bought by hunters of 200 SEK per hunting day (cf. Willebrand and from outside Sweden, especially from Norway and Paulrud 2004), there is a total consumer surplus of Italy. There is a general bag limit, of six to eight 10 million SEK per year. Similarly, with 1.2 million willow grouse, on state-managed land and access is tourist angling days and a consumer surplus of 200 regulated when the cumulative hunting effort in an SEK per day, angling in the mountain area gives a area reaches more than 3–5 days per km depend- total consumer surplus of over 200 million SEK per ing on county, i.e. approximately 30–50% of the year. To this number may be added the value of autumn population. Since 1997, the total number fishing by non-tourists. of hunter days for willow grouse in the mountain The costs for hunters of willow grouse are esti- region has been estimated to be between 43,000 mated at around 1000 SEK per hunting day and 51,000 days. (Willebrand and Paulrud 2004). This corresponds to a total cost of 50 million SEK per year, giving a total value of 60 million SEK per year for the hunt- THE ECONOMIC AND SOCIAL ing of willow grouse. Similarly, the cost per day for IMPORTANCE OF HUNTING AND tourist fishers is estimated at 400 SEK (Kriström FISHING IN THE MOUNTAIN et al. 2004; Laitila et al. 2006). With 1.2 million tour- REGION IN SWEDEN ist fishing days, the total cost is close to 500 million Hunting and fishing is an important part of life for SEK per year. Adding the consumer surplus yields a many Swedes. According to the National Board of total valuation of 700 million SEK per year. Thus, in Fisheries (2005), around 3 million Swedes stated total the values of hunting and tourist fishing that they were interested in recreational fishing, adds up to around about three-quarters of a billion with an estimated 29 million fishing days. Their SEK per year. This is a crude estimate of the value, total costs for recreational fishing during 2005 are but shows the level of importance of hunting and estimated at nearly 3 billion SEK. With regard to fishing in the mountain area. When judging the hunting, 286,000 Swedes bought a hunting licence importance of these values, one should bear in for the hunting season 2006/2007. The number of mind that the population of the communities in the hunters is one-tenth that of recreational fishers, but mountain area is only around 150,000 (Statistics their economic importance is relatively larger, with Sweden 2006). the market value of all game meat estimated at 700 million SEK per year (Kriström and Skånberg 2001; Boman 2003). MANAGEMENT STRATEGIES, NEW For the local population, the importance of fish IDEAS and wildlife is much larger in the northern part of Harvesting of Arctic char Sweden, especially in mountain communities. Ericsson et al. (2005) showed that around 75% and The effects of different harvesting methods and 50% of the population of the mountain communi- levels on char populations are currently low and ties live in households with family members who mainly extend to include the effects of gill-netting are anglers and hunters, respectively. This can be on densities of large char (Langeland 1995; compared with the proportions for Sweden as a Finstadt et al. 2001; Filipsson 2003). Depending on International Journal of Biodiversity Science and Management 337 Fish and wildlife in the Swedish mountain region Eriksson et al. whether target individuals of char mainly gain their Moreover, in such systems, strong disturbances energy from cannibalism (large-sized cannibals) or (mortality), if not directed towards the largest indi- gain most of their energy from feeding on viduals, increase individual growth rates, densities macroinvertebrates, the population responses to of larger char, and maximum size of char, thus pro- harvest probably differ. Previously it was thought viding managers with an option to improve quality that stunted Arctic char populations are a stable in terms of individual size (e.g. Langeland 1986; state, caused by over-exploitation of large cannibals Klemetsen et al. 2002). that regulated the densities of smaller individuals. An approach based on both theoretical analysis This ‘thinning’ by cannibals in turn increased the of different management options and experi- growth rate of intermediate-sized char and allowed mental studies in natural systems is needed. In this new individuals to enter the cannibalistic niche. context, recently, a new model technique (Physio- Recent studies, however, suggest that this is not the logical Structured Populations Models) has been case, and that the renewal and densities of large developed, incorporating the individual life history cannibals appear not to depend on cannibal densi- parameters, food-dependent growth and the pres- ties per se, but on the breakthrough of strong ence of size structure in fish populations (Persson recruitment pulses of char or other occasionally et al. 1998; Claessen et al. 2002; Persson et al. 2003). high densities of suitable resources (Griffith 1994; Within the mountain Mistra programme, a physio- Claessen et al. 2002; Byström 2006; Finstadt et al. logical structured population model has been 2006). Thus, in such systems with large cannibals, developed and parameterised for Arctic char. there is no space for compensatory responses and Analysis of this model reveals that it realistically the removal of a large cannibal does not open up captures and describes the dynamics and indivi- the cannibalistic niche for the new individual to dual-level characteristics of natural undisturbed enter; instead, if large cannibals are over-fished, char populations (Byström et al. in preparation). others may emerge again over time as a conse- We expect that the model will provide a powerful quence, for instance, of a breakthrough of a strong tool with which to analyse the consequences of recruitment pulse of char (Byström 2006). Densi- different harvesting options and the development ties of large cannibals are also relatively low: one of management strategies for char populations. or two individuals per hectare (Finstadt et al. 2001; Byström 2006; Byström unpublished data). Com- Harvesting of willow grouse populations bined with their high activity and the use of large spatial areas for cannibalism (search for victims), Aanes et al. (2002) used the population abundance densities of large cannibals are highly sensitive and harvest data from the southern mountain to gill-netting, for example (Finstadt et al. 2003; range in Sweden to estimate that the maximum Byström 2006). Similarly, angling from a boat in a sustainable yield of willow grouse is about 44%. cannibalistic system in a 17 ha lake resulted in the They also suggested that the best strategy to maxi- capture of 30% of the larger char (>30 cm) within a mise yield and minimise risk of extinction was few hours (Byström unpublished data), suggesting threshold harvesting or threshold combined with the same holds for vulnerability to angling. Thus, proportional harvesting. However, small-game hunt- high capture efficiency and lack of compensatory ing is rarely focused on maximising the bag but, response can make cannibalistic char systems very rather, to avoid reaching harvest levels that could sensitive to over-fishing in terms of the densities result in irreversible changes in the population and of large-sized cannibals. its food web. In a sense, it could be described as assur- In systems where target char individuals mainly ing that there will always be a safe distance to the derive their energy from macroinvertebrates, there estimated maximum sustainable yield without is much more space for compensatory responses, as having to resort to expensive monitoring. these individuals show a strong density dependence Catch per unit effort (CPUE) models have been in growth and stability in population dynamics widely used for fisheries, though with varying suc- (Langeland 1986; Johnsson 1994). In such systems, cess (Hilborn and Walter 1993). In contrast, the sustainable fishing mortality obviously depends Hörnell-Willebrand (2005) have shown that on lake productivity; knowledge of sustainable neither CPUE nor total bag of willow grouse hunt- harvesting levels is lacking but much needed. ing in the Swedish mountain is a reliable index of 338 International Journal of Biodiversity Science and Management Fish and wildlife in the Swedish mountain region Eriksson et al. population abundance. Hunter effort provided the demanding control system. The results show that most useful information in understanding harvest about 75% of a typical management area could be dynamics, and is a promising candidate as a base- open to excessive hunting that would lead to extinc- line variable to monitor the harvest of willow tion if it occurred in the whole area. The manage- grouse. We have shown that the catchability of ment system would only need to ensure that the willow grouse increased as the populations de- refuges are not utilised, once the required size and creased in several areas (Hornell-Willebrand distribution of buffers are established. This harvest 2005). Conversely, density-dependent mortality, in strategy would not work where ownership is combination with high catch rates, could reduce a divided, so that each landowner has a single grid to population below a critical level of of density which manage. Thus, a population in one area could inevitably leads to extinction. A fundamental ques- decline even with no change in hunting, if the har- tion is whether catchability continuously increases vest in the surrounding areas changed. Despite the as a population of grouse declines, or if a threshold advantages of buffer areas, they have not been is reached where hunter success begins to decline. widely adopted when developing harvest strategies More grouse are encountered at higher densities, for wildlife management. In fisheries, reserves or but a hunter’s interest in catching additional refuges have been proposed to reduce chances of grouse will decrease, leading to lower catches. Atti- over-harvesting, and this has also received some tudes and values of willow grouse hunters show that support, under certain conditions, from analyses of they do not maximise the number of grouse bagged general harvest models (see Jonzén et al. 2001). The (Willebrand and Paulrud 2004). Instead, the high- use of buffer areas when harvesting game has a est ranking factor for quality of hunting is associ- close relation to the source–sink concept (Pulliam ated with the performance of their pointing dog. 1988). A local harvest creates an artificial sink that Willow grouse hunting could well be characterised makes the buffer area develop into a source. as a dog sport; more than 90% of hunters use We suggest that the annual accumulated effort pointers for hunting. Second in quality ranking is the single best available tool for managing the came a high number of willow grouse encounters; harvest of small game. Setting limits to the total the bag was not important. This pattern is also allowable effort within an area probably has more described by Devers (2005), for dedicated ruffed potential for controlling harvests of small game grouse hunters in the Appalachian region, USA. than daily bag limits and adjusting the length of the Their satisfaction tended to be based on time in the open season. Additional control over the system field, the probability of flushing grouse, and work- can be obtained if areas are set aside as buffer and a ing with their dogs, not necessarily the number of limit is placed on the harvest effort in grids that are harvested grouse. However, there is an obvious gap open for hunting. in our knowledge about hunter expectations and the values forming hunter behaviour. Combining predator–prey theory from natural science with CONCLUSIONS human behaviour from sociology will provide a framework for understanding the relationships Fishing and the hunting of small game have between hunter behaviour (effort and efficiency) changed from being an important part of sub- and population density. sistence in rural areas to being mainly recreational. To further increase the safety margins of sport At present, the value of commercial catches are hunting, a spatial stochastic model of a fluctuating much lower than the estimated welfare values population of willow grouse was used to investigate created by recreational fishing and hunting in the the effects of buffer zones in managing harvest mountain area. The management of fish and small (Willebrand and Hörnell 2001). The analysis game was formerly characterised by a large-scale suggested that the use of buffer zones may provide a approach with a relatively limited active manage- simple strategy for harvest management. This ment: mostly the regulation of seasonal length and strategy is particularly advantageous when, rather restriction on access. Access to the large areas of than harvesting a maximum sustainable yield, the crown land under the direct management of the goal is to provide as many hunting opportunities state in the mountain region was limited, and pro- as possible without the need for a resource- vided little room for private commercial initiatives. International Journal of Biodiversity Science and Management 339 Fish and wildlife in the Swedish mountain region Eriksson et al. Differences in management between small game to set up a management strategy or harvesting and fish have mainly been that the disturbance regime that would be independent of harvest and effect of small-game hunting on other activities, management in surrounding areas. such as reindeer herding and tourism, has been of It is now possible to evaluate different harvesting high priority whereas, for fish, the regulation scenarios in different lakes using the recent model of fishing effort in running waters, identified as approach (Persson et al. 2003). This will likely be sensitive (i.e. tributaries and smaller streams), has a powerful management tool to evaluate the out- been seen as important. Little attention was paid to comes of different management scenarios. Predic- how the ecological characteristics of the different tions can be evaluated at both population and systems call for different management strategies. individual levels for any management system, as Questions regarding the management options long as careful monitoring of the key variables is in and use of fish and small game are presently in place. A similar approach is possible with willow focus due to unclear tenure and property rights. A grouse, using the spatial harvest model and CPUE. major ecological difference between char and However, if these models are to provide meaningful grouse populations is the spatial scale that has to be evaluation, management goals are also needed. considered when setting up management strate- Recent advances in the ecological understanding of gies. The char is associated with lake systems of fish and willow grouse in the mountain region have different size; populations of char in neighbouring greatly increased possibilities for experimental or lakes can show remarkably different structures. adaptive management. It would provide a number Hence, even lakes that are close to one another may of keys to the population ecology of these species need different management strategies, suggesting if such programmes could be launched in the that the management of mountain fish populations Swedish mountain region. Nevertheless, it is crucial is likely to be most effective on a small local scale. that other disciplines, such as economics, political Willow grouse, on the other hand, show large-scale science and sociology, find ways to integrate with movement, which has been suggested to be a major ecology. 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Slutbetänkande av jakt- Willebrand T and Paulrud A. Småviltjakt i Jämtland och fiskerättsutredningen. Statens Offentliga 2003 – så tycker jägarna. FjällMistra Rapport Utredningar; 2005:116 2004 no. 9, Fjällmistra, SLU, Umeå, Sweden [in Steadman RC. Is it really just a social construction? Swedish] The contribution of the physical environment to 342 International Journal of Biodiversity Science and Management http://www.deepdyve.com/assets/images/DeepDyve-Logo-lg.png International Journal of Biodiversity Science & Management Taylor & Francis

Fish and wildlife in the Swedish mountain region: resources, use and management

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International Journal of Biodiversity Science and Management 2 (2006) 334–342 Fish and wildlife in the Swedish mountain region: resources, use and management 1 2 1,2 3 4 5 T. Eriksson , J. Andersson , P. Byström , M. Hörnell-Willebrand , T. Laitila , C. Sandström and T. Willebrand Department of Aquaculture, Swedish University of Agricultural Sciences, Umeå, Sweden Department of Ecology and Environmental Science, Umeå University, Umeå, Sweden Department of Animal Ecology, Swedish University of Agricultural Sciences, Umeå, Sweden Currently: International Institute for Applied System Analysis (IIASA), Laxenburg, Austria Department of Forest Economics, Swedish University of Agricultural Sciences, Umeå, Sweden Department of Political Science, Umeå University, Umeå, Sweden Key words: Fish, wildlife, Arctic char, willow grouse, ecology, management, property regimes SUMMARY Hunting and fishing are an important part of life for many Swedes. The importance of fish and wildlife is even greater in the northern part of Sweden and especially in the Swedish mountain area, which covers a large geographical area but is sparsely populated. The bio- logical systems in the mountain ecosystems can be characterised as arctic with low produc- tivity and biodiversity. This article addresses two important species typical for this region (willow grouse and Arctic char), provides an up-to-date characterisation of current use and management, and presents a range of management models that frame the mecha- nisms of population dynamics of fish and game. It is concluded that lack of knowledge of population responses of fish and game in the mountain region to different harvesting options and management strategies is a practical obstacle, as is knowledge and under- standing of the costs and benefits associated with different ecological management strategies. However, it is also crucial to integrate other disciplines with ecology for success- ful management. INTRODUCTION Fish and wildlife have significant consumptive and importance of wildlife and freshwater fish has non-consumptive values (Mattsson 1990; Rockel declined over much of Europe, game and fish are and Kealy 1991; Murray and Simcox 2003). Wild- still used for food in many households in the moun- life, especially fish, is an important source of food tain region of Sweden (Ericsson et al. 2005). and cash in many societies. Hunting and fishing are Many communities in the Swedish mountain often central to the perception of the landscape region have faced substantial declines in jobs for people in rural areas (Greider and Garkovich previously generated by forestry, hydro-power and 1994; Steadman 2003). While the consumptive agriculture (Lundmark 2006). The most important Correspondence: T. Eriksson, Department of Aquaculture, Swedish University of Agricultural Sciences, SE-901 83 Umeå, Sweden. Email: torleif.eriksson@vabr.slu.se 334 Fish and wildlife in the Swedish mountain region Eriksson et al. source of employment today is the public sector. 6 kg/ha/year in lakes and from 2 to 15 kg/ha/year The region has also experienced net emigration, in running waters (Hammar 1996). Moose resulting in an imbalanced demographic popula- (Alces alces) and grouse species (willow grouse, rock tion structure. However, increased welfare and ptarmigan (Lagopus mutus), capercaillie (Tetrao leisure time in modern societies are believed to be a urogallus)and blackgrouse (Tetrao tetrix)) are the factor attracting more visitors to the region. A most popular game in the region. Willow grouse are general increase in tourism is therefore seen as a widespread, inhabiting primary arctic tundra, open- possible new source of employment, but this indus- ings in boreal forest, forest edge habitats, and try has not yet grown as quickly as other industries sub-alpine vegetation. Autumn densities range from have declined (Lundmark 2006). 1to50birds per km (Hörnell-Willebrand 2005). Wilderness, fish and wildlife are typical Our two focal species, Arctic char and willow resources of the region, and have led to the explora- grouse, show major differences in their life history tion of new areas for recreational hunting and fish- characteristics as well as in their position in the food ing. To date, however, there is limited knowledge of web. Arctic char and many other target species for the ecology and harvesting of these resources, so fishing are at the top of the food web (Walters and that it is difficult to estimate the potential of devel- Kitchell 2001; Karlsson and Byström 2005). As top oping fishing and hunting tourism. Increased pres- predator, Arctic char have strong effects on lower sure on these resources would certainly also result trophic levels, but are therefore commonly in competition and conflicts over access to them. resource-limited; individual growth rates are Furthermore, hunting and fishing rights are con- strongly resource-dependent (Byström and tested and overlapping between landowners and Andersson 2005; Byström 2006; Finstadt et al. 2006). the Sami, the indigenous people in the mountain In contrast, willow grouse is a typical herbivore region. However, this article does not discuss the specialised in low-digestible browsing during land-use or property rights system (for an overview winter. It does not show such an obvious food limi- of this conflict see Ekenberg 2000; Hahn 2000). tation as Arctic char, although it can be argued that Instead, it focuses on two important systems which food quality influences adult body condition and are typical for this region: willow grouse (Lagopus the subsequent vitality and survival of offspring lagopus) and salmonids (mainly Arctic char (Salve- (Brittas 1984). The proportion of eggs laid in linus alpinus). We give an up-to-date characterisa- spring that survive, and subsequently become tion of current use and management, and present a young grouse in autumn, varies greatly. The com- range of management models that frame the mech- plex fluctuation pattern and population changes anisms of population dynamics of fish and game. can be explained by trophic interactions with We then summarise research results on ecological predators and possibly food (vegetation) quality management strategies and economic values, and (Brittas 1984; Steen et al. 1988; Smith and discuss how these findings can be merged in a more Willebrand 1999) (Figure 1). complete and policy-relevant range of manage- Consequently, harvesting char or grouse will ment options, irrespective of land-use rights. potentially have different effects on population ECOLOGICAL CHARACTERISTICS SETTING THE FRAMEWORK The Swedish mountain region covers about 165,000 km . The area is rich in lakes, with about 61,000 lakes larger than one hectare and a total lake area of 8240 km . There are about 20,000 km of running water (National Board of Fisheries 2004). The species mainly fished are Arctic char (Salvelinus alpinus), whitefish (Coregonus spp.) and brown trout (Salmo trutta) in lakes, and European grayling (Thymallus thymallus) and brown trout in streams. Figure 1 Characteristic population fluctuation of Crude production estimates range from 0.5 to willow grouse at Lövhögen, Jämtland, 1962–1994 International Journal of Biodiversity Science and Management 335 Fish and wildlife in the Swedish mountain region Eriksson et al. dynamics. Arctic char, like fish in general, grow larger lakes, household fisheries in lakes, and recre- continuously throughout life, with the rate of ational fisheries in lakes and streams. Sport fishing growth depending on resource availability and is allowed in 85–90% of the waters in the mountain competition. It therefore takes a substantial num- area. There are no limitations in the number of ber of years for Arctic char (and other fish top pred- licences given in these waters, but a number ators) to reach a size suitable for harvest. This of waters are open only for a restricted number of pattern is even more pronounced in recreational fishing days. Furthermore, private waters, waters fisheries which aim for the largest individuals (Post used by the Sami organised in Reindeer Herding et al. 2002). Target char individuals may range in Communities (RHCs) and some sensitive waters age from 5–6 years to 20 years depending on the sys- are not included. In the waters open for recre- tem and size of individuals (Svenning and ational fishing, there are some limitations, such as Borgström 1995; Finstadt et al. 2006; Byström 2006). a bag limit and a minimum size limit. The presence of size structure in Arctic char, and The RHCs have rights to fish in all waters on fish populations in general, also make the interac- crown land. For all others, a permit is needed for tions between char and other fish species and their commercial or household fisheries with gillnets. resources complex and in many cases unpredict- These permits are personal and give fishing access able when it comes to the effects of harvesting large to one or more lakes. Household fishing has a individuals (Post et al. 2002; Walters and Kitchell limitation on the number of nets allowed. The 2001). Furthermore, even if the char population number of licences has been reduced during the may respond by compensatory increase, due to past 10 years (in Norrbotten county by 60% to increased resource availability for each individual, ~1000 licences, in Västerbotten county by 30% to the fact that char is a long-lived species, with a long ~650 licences). time to reach maturity, means that the compensa- Commercial catches of fish using gillnets have tion will not appear from one year to the next, even continuously decreased over the past 40 years. In though a female char may lay from 100 to more Norrbotten, the landings from the larger lakes have than 1000 eggs (Klemetsen et al. 2003). decreased from 180 tons in the 1960s to about In contrast to char, individuals in the grouse 40 tons in 2000. In recent years, fewer licences for population are similar in their survival and fecun- fishing with gillnets have been sold in Norrbotten dity from their first year. Willow grouse females lay and Västerbotten, while the numbers of sport fish- between 7 and 12 eggs, and have the potential to ing licences sold have remained more or less con- re-nest if the eggs are stolen or eaten. Even first-year stant (Figure 2). Estimated catches have decreased breeders seem to perform as well as older grouse. by approximately 25% over the last 5 years, mainly There is no apparent selection by hunters, and no due to a reduction of gillnet fishing. trophy characters that would potentially change the structure of the population. Harvest adds a mortal- ity source to the many different predators that prey on willow grouse (Smith and Willebrand 1999). Jämtland Norrbotten The complex trophic interaction, which induces an apparent strong stochastic component in the annual increase of the population, makes the response to harvesting difficult to predict (Pedersen et al. 2004; Hörnell-Willebrand 2005). The risk is that the small effects of harvesting might go undetected until a 50000 threshold is reached when the system could collapse (Willebrand and Hörnell 2001). 1960 1965 1970 1975 1980 1985 1990 1995 2000 2005 RIGHTS, RULES, USAGE AND VALUES Year Fishing Figure 2 Number of sport fishing licences sold in The fishing resource is traditionally used by a Jämtland 1960–2003 and Norrbotten 1995–2003 (data number of user groups: commercial fisheries in from County Administrations) 336 International Journal of Biodiversity Science and Management # Licenses sold Fish and wildlife in the Swedish mountain region Eriksson et al. whole: 43% and 13%, respectively. The utilisation Hunting of game meat also seems to play an important social The willow grouse hunting season runs from 25 role. The proportion of households consuming August to the last day of February in the southern game meat on a regular basis is far larger than the part of the mountain range and until 15 March in proportion of households with hunter members. the northern part. Willow grouse are hunted with For instance, in the mountain area, about three out shotguns, principally using pointing dogs; two- of four people live in households where game meat thirds of all hunting takes place during the first is used regularly. 10 days. In 1993, a decision in the parliament per- The fish and wildlife resource in the mountain mitted access to state-managed land (60,000 km ), area adds considerably to the welfare of anglers and and the interest in willow grouse hunting has since hunters in Sweden. For instance, with 50,000 hunt- increased; however, a significant proportion (about ing days for willow grouse and a consumer surplus 30%) of the hunting permits are bought by hunters of 200 SEK per hunting day (cf. Willebrand and from outside Sweden, especially from Norway and Paulrud 2004), there is a total consumer surplus of Italy. There is a general bag limit, of six to eight 10 million SEK per year. Similarly, with 1.2 million willow grouse, on state-managed land and access is tourist angling days and a consumer surplus of 200 regulated when the cumulative hunting effort in an SEK per day, angling in the mountain area gives a area reaches more than 3–5 days per km depend- total consumer surplus of over 200 million SEK per ing on county, i.e. approximately 30–50% of the year. To this number may be added the value of autumn population. Since 1997, the total number fishing by non-tourists. of hunter days for willow grouse in the mountain The costs for hunters of willow grouse are esti- region has been estimated to be between 43,000 mated at around 1000 SEK per hunting day and 51,000 days. (Willebrand and Paulrud 2004). This corresponds to a total cost of 50 million SEK per year, giving a total value of 60 million SEK per year for the hunt- THE ECONOMIC AND SOCIAL ing of willow grouse. Similarly, the cost per day for IMPORTANCE OF HUNTING AND tourist fishers is estimated at 400 SEK (Kriström FISHING IN THE MOUNTAIN et al. 2004; Laitila et al. 2006). With 1.2 million tour- REGION IN SWEDEN ist fishing days, the total cost is close to 500 million Hunting and fishing is an important part of life for SEK per year. Adding the consumer surplus yields a many Swedes. According to the National Board of total valuation of 700 million SEK per year. Thus, in Fisheries (2005), around 3 million Swedes stated total the values of hunting and tourist fishing that they were interested in recreational fishing, adds up to around about three-quarters of a billion with an estimated 29 million fishing days. Their SEK per year. This is a crude estimate of the value, total costs for recreational fishing during 2005 are but shows the level of importance of hunting and estimated at nearly 3 billion SEK. With regard to fishing in the mountain area. When judging the hunting, 286,000 Swedes bought a hunting licence importance of these values, one should bear in for the hunting season 2006/2007. The number of mind that the population of the communities in the hunters is one-tenth that of recreational fishers, but mountain area is only around 150,000 (Statistics their economic importance is relatively larger, with Sweden 2006). the market value of all game meat estimated at 700 million SEK per year (Kriström and Skånberg 2001; Boman 2003). MANAGEMENT STRATEGIES, NEW For the local population, the importance of fish IDEAS and wildlife is much larger in the northern part of Harvesting of Arctic char Sweden, especially in mountain communities. Ericsson et al. (2005) showed that around 75% and The effects of different harvesting methods and 50% of the population of the mountain communi- levels on char populations are currently low and ties live in households with family members who mainly extend to include the effects of gill-netting are anglers and hunters, respectively. This can be on densities of large char (Langeland 1995; compared with the proportions for Sweden as a Finstadt et al. 2001; Filipsson 2003). Depending on International Journal of Biodiversity Science and Management 337 Fish and wildlife in the Swedish mountain region Eriksson et al. whether target individuals of char mainly gain their Moreover, in such systems, strong disturbances energy from cannibalism (large-sized cannibals) or (mortality), if not directed towards the largest indi- gain most of their energy from feeding on viduals, increase individual growth rates, densities macroinvertebrates, the population responses to of larger char, and maximum size of char, thus pro- harvest probably differ. Previously it was thought viding managers with an option to improve quality that stunted Arctic char populations are a stable in terms of individual size (e.g. Langeland 1986; state, caused by over-exploitation of large cannibals Klemetsen et al. 2002). that regulated the densities of smaller individuals. An approach based on both theoretical analysis This ‘thinning’ by cannibals in turn increased the of different management options and experi- growth rate of intermediate-sized char and allowed mental studies in natural systems is needed. In this new individuals to enter the cannibalistic niche. context, recently, a new model technique (Physio- Recent studies, however, suggest that this is not the logical Structured Populations Models) has been case, and that the renewal and densities of large developed, incorporating the individual life history cannibals appear not to depend on cannibal densi- parameters, food-dependent growth and the pres- ties per se, but on the breakthrough of strong ence of size structure in fish populations (Persson recruitment pulses of char or other occasionally et al. 1998; Claessen et al. 2002; Persson et al. 2003). high densities of suitable resources (Griffith 1994; Within the mountain Mistra programme, a physio- Claessen et al. 2002; Byström 2006; Finstadt et al. logical structured population model has been 2006). Thus, in such systems with large cannibals, developed and parameterised for Arctic char. there is no space for compensatory responses and Analysis of this model reveals that it realistically the removal of a large cannibal does not open up captures and describes the dynamics and indivi- the cannibalistic niche for the new individual to dual-level characteristics of natural undisturbed enter; instead, if large cannibals are over-fished, char populations (Byström et al. in preparation). others may emerge again over time as a conse- We expect that the model will provide a powerful quence, for instance, of a breakthrough of a strong tool with which to analyse the consequences of recruitment pulse of char (Byström 2006). Densi- different harvesting options and the development ties of large cannibals are also relatively low: one of management strategies for char populations. or two individuals per hectare (Finstadt et al. 2001; Byström 2006; Byström unpublished data). Com- Harvesting of willow grouse populations bined with their high activity and the use of large spatial areas for cannibalism (search for victims), Aanes et al. (2002) used the population abundance densities of large cannibals are highly sensitive and harvest data from the southern mountain to gill-netting, for example (Finstadt et al. 2003; range in Sweden to estimate that the maximum Byström 2006). Similarly, angling from a boat in a sustainable yield of willow grouse is about 44%. cannibalistic system in a 17 ha lake resulted in the They also suggested that the best strategy to maxi- capture of 30% of the larger char (>30 cm) within a mise yield and minimise risk of extinction was few hours (Byström unpublished data), suggesting threshold harvesting or threshold combined with the same holds for vulnerability to angling. Thus, proportional harvesting. However, small-game hunt- high capture efficiency and lack of compensatory ing is rarely focused on maximising the bag but, response can make cannibalistic char systems very rather, to avoid reaching harvest levels that could sensitive to over-fishing in terms of the densities result in irreversible changes in the population and of large-sized cannibals. its food web. In a sense, it could be described as assur- In systems where target char individuals mainly ing that there will always be a safe distance to the derive their energy from macroinvertebrates, there estimated maximum sustainable yield without is much more space for compensatory responses, as having to resort to expensive monitoring. these individuals show a strong density dependence Catch per unit effort (CPUE) models have been in growth and stability in population dynamics widely used for fisheries, though with varying suc- (Langeland 1986; Johnsson 1994). In such systems, cess (Hilborn and Walter 1993). In contrast, the sustainable fishing mortality obviously depends Hörnell-Willebrand (2005) have shown that on lake productivity; knowledge of sustainable neither CPUE nor total bag of willow grouse hunt- harvesting levels is lacking but much needed. ing in the Swedish mountain is a reliable index of 338 International Journal of Biodiversity Science and Management Fish and wildlife in the Swedish mountain region Eriksson et al. population abundance. Hunter effort provided the demanding control system. The results show that most useful information in understanding harvest about 75% of a typical management area could be dynamics, and is a promising candidate as a base- open to excessive hunting that would lead to extinc- line variable to monitor the harvest of willow tion if it occurred in the whole area. The manage- grouse. We have shown that the catchability of ment system would only need to ensure that the willow grouse increased as the populations de- refuges are not utilised, once the required size and creased in several areas (Hornell-Willebrand distribution of buffers are established. This harvest 2005). Conversely, density-dependent mortality, in strategy would not work where ownership is combination with high catch rates, could reduce a divided, so that each landowner has a single grid to population below a critical level of of density which manage. Thus, a population in one area could inevitably leads to extinction. A fundamental ques- decline even with no change in hunting, if the har- tion is whether catchability continuously increases vest in the surrounding areas changed. Despite the as a population of grouse declines, or if a threshold advantages of buffer areas, they have not been is reached where hunter success begins to decline. widely adopted when developing harvest strategies More grouse are encountered at higher densities, for wildlife management. In fisheries, reserves or but a hunter’s interest in catching additional refuges have been proposed to reduce chances of grouse will decrease, leading to lower catches. Atti- over-harvesting, and this has also received some tudes and values of willow grouse hunters show that support, under certain conditions, from analyses of they do not maximise the number of grouse bagged general harvest models (see Jonzén et al. 2001). The (Willebrand and Paulrud 2004). Instead, the high- use of buffer areas when harvesting game has a est ranking factor for quality of hunting is associ- close relation to the source–sink concept (Pulliam ated with the performance of their pointing dog. 1988). A local harvest creates an artificial sink that Willow grouse hunting could well be characterised makes the buffer area develop into a source. as a dog sport; more than 90% of hunters use We suggest that the annual accumulated effort pointers for hunting. Second in quality ranking is the single best available tool for managing the came a high number of willow grouse encounters; harvest of small game. Setting limits to the total the bag was not important. This pattern is also allowable effort within an area probably has more described by Devers (2005), for dedicated ruffed potential for controlling harvests of small game grouse hunters in the Appalachian region, USA. than daily bag limits and adjusting the length of the Their satisfaction tended to be based on time in the open season. Additional control over the system field, the probability of flushing grouse, and work- can be obtained if areas are set aside as buffer and a ing with their dogs, not necessarily the number of limit is placed on the harvest effort in grids that are harvested grouse. However, there is an obvious gap open for hunting. in our knowledge about hunter expectations and the values forming hunter behaviour. Combining predator–prey theory from natural science with CONCLUSIONS human behaviour from sociology will provide a framework for understanding the relationships Fishing and the hunting of small game have between hunter behaviour (effort and efficiency) changed from being an important part of sub- and population density. sistence in rural areas to being mainly recreational. To further increase the safety margins of sport At present, the value of commercial catches are hunting, a spatial stochastic model of a fluctuating much lower than the estimated welfare values population of willow grouse was used to investigate created by recreational fishing and hunting in the the effects of buffer zones in managing harvest mountain area. The management of fish and small (Willebrand and Hörnell 2001). The analysis game was formerly characterised by a large-scale suggested that the use of buffer zones may provide a approach with a relatively limited active manage- simple strategy for harvest management. This ment: mostly the regulation of seasonal length and strategy is particularly advantageous when, rather restriction on access. Access to the large areas of than harvesting a maximum sustainable yield, the crown land under the direct management of the goal is to provide as many hunting opportunities state in the mountain region was limited, and pro- as possible without the need for a resource- vided little room for private commercial initiatives. International Journal of Biodiversity Science and Management 339 Fish and wildlife in the Swedish mountain region Eriksson et al. Differences in management between small game to set up a management strategy or harvesting and fish have mainly been that the disturbance regime that would be independent of harvest and effect of small-game hunting on other activities, management in surrounding areas. such as reindeer herding and tourism, has been of It is now possible to evaluate different harvesting high priority whereas, for fish, the regulation scenarios in different lakes using the recent model of fishing effort in running waters, identified as approach (Persson et al. 2003). This will likely be sensitive (i.e. tributaries and smaller streams), has a powerful management tool to evaluate the out- been seen as important. Little attention was paid to comes of different management scenarios. Predic- how the ecological characteristics of the different tions can be evaluated at both population and systems call for different management strategies. individual levels for any management system, as Questions regarding the management options long as careful monitoring of the key variables is in and use of fish and small game are presently in place. A similar approach is possible with willow focus due to unclear tenure and property rights. A grouse, using the spatial harvest model and CPUE. major ecological difference between char and However, if these models are to provide meaningful grouse populations is the spatial scale that has to be evaluation, management goals are also needed. considered when setting up management strate- Recent advances in the ecological understanding of gies. The char is associated with lake systems of fish and willow grouse in the mountain region have different size; populations of char in neighbouring greatly increased possibilities for experimental or lakes can show remarkably different structures. adaptive management. It would provide a number Hence, even lakes that are close to one another may of keys to the population ecology of these species need different management strategies, suggesting if such programmes could be launched in the that the management of mountain fish populations Swedish mountain region. Nevertheless, it is crucial is likely to be most effective on a small local scale. that other disciplines, such as economics, political Willow grouse, on the other hand, show large-scale science and sociology, find ways to integrate with movement, which has been suggested to be a major ecology. Prioritising only for increased ecological process for compensating locally high harvest rates understanding to reveal the necessary components (Willebrand and Hörnell 2001). Although the pre- will not be sufficient for successful management; it cise area depends on landscape features, it is not demands new skills among both researchers and unlikely that about 100–200 km would be needed stakeholders. REFERENCES Aanes S, Engen S, Saether BE, Willebrand T and Claessen D, Van Oss C, DeRoos AM and Persson L. Markström V. Sustainable harvesting strategies of The impact of size-dependent predation on willow ptarmigan in a fluctuating environment. population dynamics and individual life history. Ecological Applications 2002;12:281–90 Ecology 2002;83:1660–75 Boman M. Vilt och välfärdsekonomie. K Skogs- o Lantbr Devers PK. Population ecology and the effects of akad Tidskr 2003;142(2):35–9 hunting ruffed grouse (Bonasa umbellus) in the Brittas R. 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Journal

International Journal of Biodiversity Science & Management – Taylor & Francis

Published: Dec 1, 2006

Keywords: FISH; WILDLIFE; ARCTIC CHAR; WILLOW GROUSE; ECOLOGY; MANAGEMENT; PROPERTY REGIMES

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