Tetrao urogallus Linnaeus, 1758
Text, map and photo from
Storch I. (2000) : Grouse Action Plan 2000-2004,
reproduced here with the Editor's agreement
Common
names:
Song ji
Chinese
Veliki
tetrijeb
Croatian
Tetrev
hlu_ec
Czech
Tiur
Danish
Capercaillie
English
Metso (male),
koppelo (female)
Finnish
Grand
tétras
French
Auerhuhn
German
Agriokourkos
Greek
Gallo
cedrone
Italian
Storfugl, Tiur
(male), røy (female)
Norwegian
Gluszec
Polish
Glukhar
Russian
Divji
Petelin
Slovenian
Urogallo
Spanish
Tjäder
Swedish
Conservation Status
1996 IUCN: Lower risk
(least concern)
CITES 1998: not listed in Appendices.
Bern Convention: T. u. cantabricus (Appendix II)
EU Birds Directive: Annex I, Annex II/2, Annex III/2
National red data books: listed in western, central, and
southeastern European countries.
The subspecies T. u. cantabricus, the Cantabrian capercaillie (Castroviejo 1967, 1975, Castroviejo et al. 1974), qualifies to be listed as endangered according to the IUCN Red List Categories under criteria EN; C2a (see Appendix 2). The subspecies inhabits a 6,000km_ range in the Cantabrian mountains of northern Spain (Castroviejo, pers. comm.). The range is severely fragmented and separated from its nearest neighbouring population in the Pyrenees (T. u. aquitanus) by a distance of 300km (J. Castroviejo pers. comm.). Numbers have been declining by an estimated 25-50% over the past 10-15 years. The population is presently estimated as <400 adult males, equivalent to a total population size of <1000 birds (Cantabrian Capercaillie Group, unpubl. data).
Taxonomy
There are 12 subspecies recognised (Potapov and Flint 1989, del Hoyo et al. 1994); two are geographically isolated, T. u. cantabricus in the Cantabrian mountains of Spain, that mostly lives on holly Ilex aquifolium instead of conifers as winter food (Castroviejo 1967, 1975, Castroviejo et al. 1974), and T. u. aquitanus in the Pyrenees of France and Spain.
Distribution
Eurasia. Contiguous distribution in the boreal forest from Scandinavia to eastern Siberia (approx. 125° east); the south-western part of the range in western and central Europe is fragmented primarily due to the patchy distribution of montane conifer forests and secondarily due to habitat loss (see Klaus et al. 1989).
Population size and trend
The capercaillie still occupies most of its original range, although serious declines in western and central Europe have resulted in local extinctions. In central Europe, many populations have disappeared. Most of the remaining ones are small (<200 birds) and probably isolated. In the boreal forests of Fennoscandia and western Russia, capercaillie numbers dropped following the beginning of intensive clearcutting. In general, the species is listed as threatened in western, central, and south-eastern Europe, but still occurs in considerable numbers throughout most of its boreal range from Scandinavia to eastern Siberia. However, in some parts of the boreal forest, e.g. Finland, the species regionally is considered vulnerable.(See Klaus et al. 1989, Office national de la Chasse 1994 a,b).
Habitat and ecology
The capercaillie is adapted to boreal climax forests. Its primary habitat is a landscape dominated by old-growth forest intermixed with bogs and patches of younger successional stages following natural disturbance such as wind-blow, snow-break, and fire. Capercaillie habitats are characterised by coniferous trees, open structure with moderate canopy cover, and rich ground vegetation dominated by bilberry Vaccinium myrtillus and other ericaceous shrubs. The birds feed almost exclusively on conifer needles in winter but on leaves, buds, flowers, fruits of various herbs and shrubs in summer (Jacob 1987). The Cantabrian subspecies is known to feed on holly in winter, but apparently some populations winter on beech buds and herbs (J. Ramón Obeso pers. comm.). Young capercaillie chicks rely on invertebrates, especially caterpillars on Vaccinium. In the temperate zone, e.g. in central Europe, Capercaillie habitats are restricted to montane regions. Capercaillie depend on particular habitat structures, but are rather flexible with regard to conifer species and forest age. In most areas, old, natural or semi-natural forests are the capercaillie´s stronghold. However, if the structure of the vegetation is suitable, the species may use young and commercial forests as well. The capercaillie is often referred to as an indicator species of healthy old forest communities in montane and boreal ecosystems (See Klaus et al. 1989, Potapov and Flint 1989, Rolstad 1989, Gjerde 1991, Ménoni 1991, Borchtchevski 1993, Storch 1995, 1997a, Graf 1998, Fischer 1999).
Hunting and cultural importance
The capercaillie has a long history as a game bird (Klaus et al. 1989, Potapov and Flint 1989) since the middle ages and even before. In central Europe, the capercaillie has received particular attention as a highly-valued hunting trophy. At times, e.g. since the 18th century, the capercaillie hunt was reserved for aristocrats. In its central European strongholds, the capercaillie has been a traditional element of local folklore until the present day. Despite its great cultural importance, however, capercaillie hunting generally has played a minor economic role in central Europe.
Throughout central Europe, capercaillie have been hunted mostly in spring at the lek, and hunters prefer to shoot supposedly high-ranking cocks. An often-discussed problem related to the spring hunt is that it may disturb mating and result in reduced reproductive success. In Scotland, the Pyrenees, Fennoscandia, and Russia, capercaillie of both sexes are mostly hunted in autumn. The autumn hunt is often considered to be less critical in terms of population dynamics, because leks are not disturbed, and because hunting losses are assumed to be at least partly compensated by reduced winter mortality among the survivors. Thorough tests of this latter assumption however are still lacking.
Whereas trophy hunting and taxidermy are the major motivation for hunting capercaillie in western and central Europe, the species is hunted for food and sport in northern Europe and mostly for food in eastern Europe and Russia. In recent times, however, trophy hunting by westerners is gaining economic importance in eastern Europe. In the boreal forest, grouse hunting has long played a major economic role and is still culturally important. In Russia, the hunting bag of capercaillie during the early 1990s was estimated at 700,000 birds annually (Grabuzov 1995, Flint 1995).
Since the 1970s, capercaillie hunting has been restricted or banned in all western and central European countries. Hunting bans did not reverse the negative population trends, however. In most areas, hunting has not been a primary cause of population declines (Klaus et al. 1989). There are several countries where capercaillie are still hunted although the species is listed as endangered in the national Red Lists (e.g. Austria, Bulgaria). There is growing opposition to this practice from conservationists and the anti-hunting movement. Moderate, strictly controlled hunting, however, may also have a positive overall effect on capercaillie conservation. For example, in parts of Austria, where the hunting rights belong to small, private land owners, the chances to sell the right to shoot a capercaillie cock to guest hunters every other year appears to be a significant incentive for active habitat preservation.
Principal threats
Habitat degradation. Loss and deterioration of habitats are assumed to be the major causes of declining capercaillie numbers. Habitat changes occur at various levels of spatial scale: As a habitat specialist, the Capercaillie is sensitive to changes in habitat structure, i.e. features at forest stand level. Due to its large spatial requirements the Capercaillie is also susceptible to changes at the landscape scale, such as forest fragmentation (e.g. Rolstad and Wegge 1989, Rolstad 1991, Ménoni et al 1997, Storch 1997a,b) and forestry practices have major influence on capercaillie populations. In the boreal forest, capercaillie numbers declined parallel to the onset of intensive clearcutting (Rolstad and Wegge 1989, Rolstad 1991). In Russia and some other eastern European countries, political and socio-economic developments will influence future forestry practices and also the intensity of poaching, and thus have the potential to create significant threats for the capercaillie and other old-forest species. In central Europe, capercaillie abundance was highest at times when human land-use practices, e.g. collection of forest litter and cattle grazing, favoured open forest structures. During the past decades, increasing standing timber volumes throughout central Europe were paralleled by declining capercaillie numbers. Forestry practices probably are the one major factor limiting capercaillie numbers (Klaus et al. 1989, Rolstad and Wegge 1989, Rolstad 1989, 1991, Storch 1997). In the UK, however, the major threat possibly leading to the extinction of capercaillie is mortality from striking fences (see below).
Small population size. In western and central Europe, deterioration and fragmentation of habitats has resulted in isolated populations, many of which are now threatened by small size. Small populations of e.g. <100 birds are generally vulnerable and show a high risk of extinction due to chance events, or loss of genetic variation (Klaus 1994, Loneux and Ruwet 1997, Westemeier 1998; see 2.6.2). Measures to preserve habitats need to be initiated well before a population is at the edge of extinction. The size and spatial distribution of suitable habitats at a landscape scale needs much more attention than it has received in the past (Rolstad 1991, Ménoni et al. 1997, Storch 1997a, b).
Pollution. In parts of central Europe, large scale pollution through wind and rain resulted in soil eutrophication and thus vegetational changes disadvantageous for the ericaceous shrubs preferred by capercaillie (see Klaus and Bergmann 1994).
Predation. Predation pressure on capercaillie seems to have significantly increased during the past three decades. Changes in land-use, e.g. forest fragmentation and agricultural fertilisation, and declining persuation of predators have resulted in increasing densities of small and medium sized mammalian and avian predators (Reynolds 1991). In addition, large-scale vaccination of foxes against rabies in central Europe is contributing to constantly high fox populations (Vos 1995). Regionally, increasing populations of wild boar have become a major cause of nest losses.
Human disturbance. Disturbance by tourism and leisure activities such as hiking, skiing, mountain-biking, etc. are viewed as a serious threat to local capercaillie populations. In some regions of Europe, programmes to limit the effects of human disturbance on capercaillie and other grouse have been initiated (e.g. Austria, France, Germany, Switzerland) (e.g. Ménoni and Magnani 1998, Zeitler and Glänzer 1998).
Collisions. Regionally, e.g. in Fennoscandia, collisions with high-tension power lines may kill significant numbers of capercaillie; >20,000 mortalities annually have been estimated in Norway (Beveranger 1995). Locally, collisions with deer fences may cause significant mortality among capercaillie. Fences that are erected in relation with woodland management schemes to exclude deer have become a serious threat for capercaillie in Scotland and account for an estimated 50% of the annual mortality (J. Oswald, pers. comm.; see Baines and Summers 1997).
Exploitation. Some poaching may occur throughout the range. Especially in eastern Europe, birds may be shot in excess of the legal hunting bags or outside the season; this is partly related to poor law enforcement and the tight economic situation in some regions. Poaching for food was reported greatly to affect populations in Ukraine; poaching and sport and trophy hunting in Spain was reported to have serious effects on the population dynamics of the capercaillie in Spain, and moderate effects in Greece; hunting is illegal in these countries and better law enforcement is desirable. Legal and illegal hunting was also felt greatly to influence the declining populations in Andorra, Bulgaria and Romania, calling for improved legal protection and law enforcement. The lek mating system generally makes the species susceptible to over-exploitation, because diplaying males are an easy target and known leks may be destroyed with little effort, as reported from Russia.
Research needs
Biology, food habits, habitat requirements, spatial requirements, behaviour and mating system are generally well studied. The book by Klaus et al. (1989) provides an extended review. Several case studies have explored the effects of landscape-scale habitat features such as habitat fragmentation on capercaillie populations (e.g. Rolstad 1989, Gjerde 1991, Ménoni 1991, Storch 1997a,b). Information is almost completely lacking on juvenile dispersal rates and dispersal distances and their roles in population genetics, dynamics and persistence. Larger-scale habitat relationships and population dynamics, and minimum requirements in population size (Storch 1995) and in habitat area and connections can be considered research topics with the greatest relevance to conservation in those regions where the species is endangered.
Current conservation measures
Legal protection. The degree of legal protection for the capercaillie varies among the range countries. In general, the species is fully and effectively protected in western and central European countries with small and declining populations. In other countries, e.g. Austria, Italy, and France, hunting is strictly regulated and only allowed in certain areas and during limited hunting seasons. In some eastern (e.g. Bulgaria, Romania) and southern (Spain, Greece) European countries, illegal hunting may have serious effects on capercaillie populations.
Protected areas. Only a minor proportion of the species´ range is covered by protected areas, and most are too small for self-sustaining, viable populations. The role of protected areas for capercaillie conservation is generally considered to be limited. In Greece, Spain, and Andorra, where habitat degradation and poaching appear to threaten the remaining populations, protected areas are believed to be critical for the survival of the species. In the Cantabrian mountains of Spain, protected areas for capercaillie conservation are under development. Protected areas may effectively maintain a capercaillie population if the area is large enough (>>10.000 ha; see Storch 1995), if the habitat is and remains suitable, and if utilisation and disturbances are strictly regulated and controlled.
Surveys and monitoring. In Europe, regular surveys or monitoring are common as a tool in planning harvests (e.g. Austria, Fennoscandia, France, UK), and in regions with small remnant populations (e.g. parts of Germany, Switzerland). Whether lek or autumn counts are conducted is largely related to regional hunting traditions. The reliability of lek counts is questionable because larger leks are difficult do overlook, the spatial organisation and attendance of a lek may change over the season, and small leks and individually displaying cocks may be missed. Lek counts are usually done by hunters or game keepers who might have their own agenda; the method has a great potential for misuse and involves a high risk of disturbance. However, if properly organised, lek counts may provide a spring population index for males. Well organised autumn counts, e.g. the Finnish wildlife triangle scheme, (Lindén et al. 1996) generally give reliable estimates of population structure and trends. Transect counts of indirect capercaillie sign such as feathers and droppings may provide a population index suitable for monitoring and regional comparisons in areas with at least moderate population densities (Storch unpubl.).
Habitat preservation. Habitat management is considered the most important conservation measure. In most regions, the major challenge is to integrate forestry practices with capercaillie habitat requirements. In this context, the argument that the capercaillie is an indicator of a healthy old forest ecosystem is helpful. In parts of Europe, recent policy changes, at least by the state forest agencies towards increasing "naturalness" of the forest, may be favourable to capercaillie habitat conservation.
Predator control. In most regions of Europe with small and highly endangered populations, predation is experienced as the major proximate threat to the capercaillie, and to many conservation practitioners, predator control seems to be desirable (e.g. Kaphegyi 1998). A significant reduction of predators will probably result in improved capercaillie survival (Reynolds 1991). Maintaining low predator densities on a large scale, however, is technically difficult and morally questionable, and not easily accepted by the public (Kaphegyi 1998). Therefore, in most areas predator control cannot be a sustainable approach to capercaillie conservation.
Captive breeding and release. Captive breeding of capercaillies has made significant progress over the past two decades, and release of capercaillies reared in captivity has become a common, however unsuccessful, conservation tool in central Europe. Newly released birds suffer great mortality from predation, which is at least partly related to rearing and releasing techniques. As a result, there is still not a single example of a self-sustaining capercaillie population established from birds reared in captivity, although over the past 20 years more than 3000 birds have been released in nine different projects in Germany alone. The prospects for translocations, i.e. release of birds caught in the wild elsewhere, might be better. Experience with capercaillie release projects have been summarised by Klaus (1997).
Education.
Capercaillie conservation measures are frequently accompanied by
information and education of land owners and foresters regarding
habitat needs, and of tourist and sport organisations such as
skiing and alpine clubs regarding the avoidance of disturbance in
grouse habitats (e.g. Zeitler and Glänzer 1998).
Priority conservation measures
On a global scale, the Cantabrian capercaillie, the only subspecies that is globally endangered, has highest priority for capercaillie conservation. Recommendations concerning the Cantabrian capercaillie are described in Chapter 4. Below, general conservation needs for the capercaillie are listed; their order of priority however may vary between different parts of the range.
Habitat preservation. Integrating forestry and capercaillie conservation is most important where industrial forestry is introduced, and where populations are declining. The major goal is maintaining on a large scale a relatively open forest structure with a well developed ground vegetation.
Maintaining spatial connections. Measures are needed to prevent further declines of the still existing larger populations of western and central Europe. Maintaining or restoring spatial connection among these populations, e.g. in the Alps, seems to be vital. The recent history of capercaillie in central Europe teaches us that once populations have become small and isolated, the chances for population restoration are poor. Preventive measures are needed for the remaining larger populations and metapopulation systems. Landscape scale aspects of capercaillie habitats need to be considered where capercaillie populations are spatially structured, e.g. in the Alps and Pyrenees (e.g. Ménoni et al. 1997, Storch 1997b).
Reduction of human disturbance. In areas with small, declining, or threatened populations, human disturbance due to sport and leisure activities should be minimised, particularly where wintering, display, and brood habitats are limited.
Monitoring. Where the species is endangered, sound monitoring programmes of populations and habitats should be established to assess population trends and the success of conservation measures.
Law enforcement. Locally, where the species appears to be threatened by poaching, better law enforcement is needed (see above).
Correspondents
David Baines, Massimo Bocca, Ayurzanyn Bold, Jordi Canut Bartra, Miran Cas, Javier Castroviejo, Roman Dziedzic, Volodymyr Domashlinets, Michael Fasel, Marijan Grubesic, David Jenkins, Lazlo Kalaber, Wolfgang Kantner, Siegfried Klaus, Harto Lindén, Antonio Lucio, Christian Marti, Ann Matschke, Emmanuel Ménoni, Alexander Mikityuk, Pierre Mollet, Jimmy Oswald, Tatjana Pavlushchick, Roald Potapov, José Ramón Obeso, Athanassios Sfougaris, Peter Shurulinkov, Torstein Storaas, Ilse Storch, Sun Yue-Hua, Albin Zeitler.
Key publications
Borchtchevski 1993. Population biology of the capercaillie. Principles of the structural organisation. Izdatelstvo Tsnil ochotnitshjego chosjaistva i zapovednikov, Moscow, 268 pp. (ISBN 5-87560-011-X)
Gjerde, I. 1991. Winter ecology of a dimorphic herbivore: temporal ans spatial relationships and habitat selection of male and female capercaillie. Ph.D. thesis, University of Bergen, Norway.
Klaus, S., Andreev, A.V., Bergmann, H.-H., Müller, F. , Porkert, J. and Wiesner, J. 1989. Die Auerhühner. Die Neue Brehm-Bücherei. Band 86. Westarp Wissenschaften, Magdeburg, Germany.
Ménoni, E. 1991. Écologie et dynamique des population du Grand Tétras dans les Pyrénées, avec des references speciales a la biologie de la réproduction chez les poules - quelques application a sa conservation. Ph.D. thesis, Univ. Toulouse, France.
Office National de la Chasse 1994. Plan de restauration pour quelque galliformes européens galliformes: gelinotte, grand tétras, tétras-lyre et perdix bartavelle &endash; tome 1. Gibier Faune Sauvage 11(1), 348 p.
Office National de la Chasse 1994. Restoration plans for some European galliformes: hazel grouse, capercaillie and black grouse; part 2. Gibier Faune Sauvage 11(2), 222 p.
Rolstad, J. 1989. Habitat and range use of capercaillie in southcentral scandinavian boreal forests. Ph.D. thesis, Agric. Univ., As, Norway.
Storch, I. 1995. Habitat requirements of capercaillie. Proc. Int. Symp. Grouse 6:151-154.
Storch, I. 1997a. The importance of scale in habitat conservation for an endangered species: the capercaillie in central Europe. Pages 310-330 in: J. A. Bissonette, editor. Wildlife and Landscape Ecology: effects of pattern and scale. Springer Verlag, New York.