Cantabrian Capercaillie
Encyclopedia
The Cantabrian Capercaillie (Tetrao urogallus cantabricus) is a subspecies
Subspecies
Subspecies in biological classification, is either a taxonomic rank subordinate to species, ora taxonomic unit in that rank . A subspecies cannot be recognized in isolation: a species will either be recognized as having no subspecies at all or two or more, never just one...

 of the Western Capercaillie in the grouse
Grouse
Grouse are a group of birds from the order Galliformes. They are sometimes considered a family Tetraonidae, though the American Ornithologists' Union and many others include grouse as a subfamily Tetraoninae in the family Phasianidae...

 family Tetraonidae.

Description

The capercaillie is a large grouse, 80 to 115 cm (31 to 45 inches) in length, with the female much smaller than the male. It has dark grey plumage
Plumage
Plumage refers both to the layer of feathers that cover a bird and the pattern, colour, and arrangement of those feathers. The pattern and colours of plumage vary between species and subspecies and can also vary between different age classes, sexes, and season. Within species there can also be a...

 with fine blackish vermiculation (wavelike pattern) around the head and neck. The breast is glossy greenish-black. It has a long, rounded tail, an ivory-white bill
Beak
The beak, bill or rostrum is an external anatomical structure of birds which is used for eating and for grooming, manipulating objects, killing prey, fighting, probing for food, courtship and feeding young...

, and a scarlet crest (World Association of Zoos and Aquaria 2009, unpaginated).

Distribution and habitat

The subspecies once ranged the length of the Cantabrian Mountains
Cantabrian Mountains
The Cantabrian Mountains or Cantabrian Range are one of the main systems of mountain ranges in Spain.They extend for more than approximately 180 miles across northern Spain, from the western limit of the Pyrenees to the edges of the Galician Massif close to Galicia, along the coast of the...

 from northern Portugal
Portugal
Portugal , officially the Portuguese Republic is a country situated in southwestern Europe on the Iberian Peninsula. Portugal is the westernmost country of Europe, and is bordered by the Atlantic Ocean to the West and South and by Spain to the North and East. The Atlantic archipelagos of the...

, through Galicia, Asturias
Asturias
The Principality of Asturias is an autonomous community of the Kingdom of Spain, coextensive with the former Kingdom of Asturias in the Middle Ages...

 and León
León, Spain
León is the capital of the province of León in the autonomous community of Castile and León, situated in the northwest of Spain. Its city population of 136,985 makes it the largest municipality in the province, accounting for more than one quarter of the province's population...

, to Cantabria
Cantabria
Cantabria is a Spanish historical region and autonomous community with Santander as its capital city. It is bordered on the east by the Basque Autonomous Community , on the south by Castile and León , on the west by the Principality of Asturias, and on the north by the Cantabrian Sea.Cantabria...

 in northern Spain
Spain
Spain , officially the Kingdom of Spain languages]] under the European Charter for Regional or Minority Languages. In each of these, Spain's official name is as follows:;;;;;;), is a country and member state of the European Union located in southwestern Europe on the Iberian Peninsula...

 (IUCN Redbook 1979, p. 1). Its range has since contracted to the mountains in northwest Spain. It inhabits an area of 1,700 km2 (656 mi2), and is separated from the nearest neighbouring subspecies (T. u. aquitanicus) in the Pyrenees
Pyrenees
The Pyrenees is a range of mountains in southwest Europe that forms a natural border between France and Spain...

 by a distance of more than 300 km (186 mi) (Quevedo et al. 2006b, p. 268).

The Cantabrian subspecies is not as reliant as other capercaillies on pine forest, a type of habitat which is relatively scarce in the Cantabrian mountains. The bird is able to feed in deciduous woodland, and occurs in mature beech forest and mixed forests of beech and oaks (at elevations ranging from 800 to 1,800 m (2,600 to 5,900 ft). The capercaillie also uses other microhabitat types such as broom
Genista
Genista is a genus of legumes which includes many species of broom. Many of these brooms are notorious as noxious weeds.Species include:*Genista aetnensis - Mount Etna broom*Genista anglica - petty whin, needle furze*Genista canariensis...

, meadow and heath
Erica
Erica ,the heaths or heathers, is a genus of approximately 860 species of flowering plants in the family Ericaceae. The English common names "heath" and "heather" are shared by some closely related genera of similar appearance....

 selectively throughout the year (Quevedo et al. 2006b, p. 271). It feeds on beech buds, birch catkins, and holly leaves. It also feeds on bilberry
Vaccinium myrtillus
Vaccinium myrtillus is an almost Holarctic species of shrub with edible fruit, usually simply referred to as "bilberry" or "whortleberry". It is more precidely called Common Bilberry or Blue Whortleberry, to distinguish it from its Vaccinium relatives...

, a common component of its diet (Rodriguez and Obeso 2000 as reported in Pollo et al. 2005, p. 398).

Conservation and status

Storch et al. estimates the population to be 627 birds, of which approximately 500 are adults, according to the most recent population data collected from 2000 through 2003 (2006, p. 654). Population estimates for species of grouse are commonly assessed by counting males that gather during the leks (traditional places where males assemble during the mating season and engage in competitive displays that attract females). Pollo et al. (2005, p. 397) estimated a 60-to-70 percentdecline in the number of male leks since 1981. This is equivalent to an average decline of 3 percent per year, or 22 percent over 8 years. There is also evidence of a 30-percent decline in lek occupancy in the northern watershed of the species’ range
Range (biology)
In biology, the range or distribution of a species is the geographical area within which that species can be found. Within that range, dispersion is variation in local density.The term is often qualified:...

 between 2000 and 2005 (Bañuelos and Quevedo, unpublished data, as reported in Storch et al. 2006, p. 654).

Based on data collected between 2000 and 2003 by Pollo et al. (2005, p. 401), the distribution of Cantabrian capercaillie on the southern slope of the Cantabrian Mountains is fragmented into 13 small subpopulations: four in the western area and 9 in the eastern. Six subpopulations (5 in the eastern and 1 in the western) contained only one singing male, which indicates a very small subpopulation, since presence of singing males is a direct correlate to population numbers.

The area occupied by Cantabrian Capercaillie in 1981–1982 covered up to approximately 2,070 km2 (799 mi2) of the southern slope 972 km2 (375 mi2) in the west and 1,098 km2 (424 mi2) in the east). Between 2000 and 2003, the area of occupancy had declined to 693 km2 (268 mi2), specifically 413 km2 (159 mi2) in the west and 280 km2 (108 mi2) in the east. Thus, over a 22–year period, there was a 66-percent reduction in the areas occupied by this subspecies on the southern slope of the Cantabrian Mountains (Pollo et al. 2005, p. 401).

Based on this data, the subpopulation in the eastern portion of the range appears to be declining at a faster rate than the subpopulation in the western portion of the range.

Although Storch, et al. 2006 (p. 653) noted that the Cantabrian Capercaillie meets the criteria to be listed as ‘‘Endangered’’ on the IUCN Red List
IUCN Red List
The IUCN Red List of Threatened Species , founded in 1963, is the world's most comprehensive inventory of the global conservation status of biological species. The International Union for Conservation of Nature is the world's main authority on the conservation status of species...

 due to ‘‘rapid population declines, small population size, and severely fragmented range’’, it is not classified as such by the IUCN. The species is classified as ‘‘Endangered’’ in Spain under the National Catalog of Endangered Species. The species has not been formally considered for listing in the CITES Appendices (http://www.cites.org).

In 2011 United States Fish and Wildlife Service
United States Fish and Wildlife Service
The United States Fish and Wildlife Service is a federal government agency within the United States Department of the Interior dedicated to the management of fish, wildlife, and natural habitats...

 listed the Cantabrian capercaillie as an endangered species.

Habitat destruction

Numerous limiting factors influence the population dynamics of the capercaillie throughout its range, including habitat degradation, loss, and fragmentation (Storch 2000, p. 83; 2007, p. 96). Forest structure plays an important role in determining habitat suitability and occupancy. Quevedo et al. (2006b, p. 274) found that open forest structure with well-distributed bilberry shrubs were the preferred habitat type of Cantabrian capercaillie. Management of forest resources for timber production has caused and continues to cause significant changes in forest structure such as: species composition, density and height of tress, forest patch size, and understory vegetation (Pollo et al. 2005, p. 406).

The historic range occupied by this subspecies (3,500 km2 (1,350 mi2)) has declined by more than 50 percent (Quevedo et al. 2006b, p. 268). The current range is severely fragmented, with low forest habitat cover (22 percent of the landscape) and most of the suitable habitat remaining in small
patches less than 10 hectares (ha) (25 acres (ac)) in size (Garcia et al. 2005, p. 34). Patches of good-quality habitat are scarce and discontinuous, particularly in the central parts of the range
(Quevedo et al. 2006b, p. 269), and leks in the smaller forest patches have been abandoned during the last few decades. The leks that remain occupied are now located farther from forest edges than
those occupied in the 1980s (Quevedo et al. 2006b, p. 271).

Based on population surveys, forest
Forest
A forest, also referred to as a wood or the woods, is an area with a high density of trees. As with cities, depending where you are in the world, what is considered a forest may vary significantly in size and have various classification according to how and what of the forest is composed...

 fragments containing occupied leks in 2000 were significantly larger than fragments containing leks in the 1980s that have since been abandoned (Quevedo et al. 2006b, p. 271). The forest fragments from which the Cantabrian capercaillie has disappeared since the 1980s are small in size, and are the most isolated from other forest patches. In addition, the Cantabrian capercaillie have disappeared from forest patches located closest to the edge of the range in both the eastern and western subpopulations of the south slope of the Cantabrian Mountains,
suggesting that forest fragmentation is playing an important role in the population dynamics of this subspecies (Quevedo et al. 2006b, p. 271). Research conducted on other subspecies of
capercaillie indicate that the size of forest patches is correlated to the number of males that gather in leks to display, and that below a certain forest patch size, leks are abandoned
(Quevedo et al. 2006b, p. 273). In highly fragmented landscapes, forest patches are embedded in a matrix of other habitats, and forest dwellers like capercaillies frequently encounter open areas within their home range. Quevedo et al. (2006a, p. 197) developed a habitat suitability model for
the Cantarian capercaillie that assessed the relationship between forest patch size and occupancy. He determined that the subspecies still remains in habitat units that show habitat suitability
indices below the cut-off values of the two best predictive models (decline and general), which may indicate a high risk of local extinction. Other researchers suggested that, should further habitat
Habitat
* Habitat , a place where a species lives and grows*Human habitat, a place where humans live, work or play** Space habitat, a space station intended as a permanent settlement...

 or connectivity loss occur, the Cantabrian capercaillie population may become so disaggregated that the few isolated subpopulations will be too small to ensure their own long-term persistence (Grimm and Storch 2000, p. 224).

A demographic model based on Bavarian alpine populations of capercaillie suggest a minimum viable
population size of the order of 500 birds (Grimm and Storch 2000, p. 222). However, genetic data show clear signs of reduced variability in populations with numbers of individuals in the range of fewer than 1,000 birds, which indicates that a demographic minimum population of 500 birds may be too small to maintain high genetic variability (Segelbacher et al. 2003, p. 1779). Genetic consequences of habitat fragmentation exist for this species in the form of increased genetic
differentiation due to increased isolation of populations (Segelbacher et al. 2003, p. 1779). Therefore, anthropogenic habitat deterioration and fragmentation not only leads to range contractions and extinctions, but may also have significant genetic, and thus, evolutionary consequences for the surviving populations (Segelbacher et al. 2003, p. 1779).

Recent population surveys show this subspecies is continuing to decline throughout its current range, andsubpopulations may be isolated from one another due to range contractions in the eastern and western portions of its range, leaving the central portion of the subspecies range abandoned (Pollo et al. 2005, p. 401). Some remaining populations may already have a high risk of local extinction (Quevedo et al. 2006a, p. 197). Management of forest resources for timber production
continues to negatively affect forest structure, thereby affecting the quality,
quantity, and distribution of suitable habitat available for this subspecies. In addition, the structure of the matrix of habitats located between forest patches is likely affecting the ability of capercaillies to disperse between subpopulations. Therefore, we find that present or threatened destruction, modification, or curtailment of the
habitat or range is a threat to the continued existence of the Cantabrian capercaillie throughout its range.

Overhunting

Currently hunting of the Cantabrian capercaillie is illegal in Spain; however, illegal hunting still occurs (Storch 2000,
p. 83; 2007, p. 96). Because this species congregates in leks, individuals are
particularly easy targets, and poaching of protected grouse is considered common (Storch 2000, p. 15). It is
unknown what the incidence of poaching is or what impact it is having on this subspecies; however, given the
limited number of birds remaining and the reduced genetic variability already
evident at current population levels, the further loss of breeding adults could
have substantial impact on the subspecies.

Disease and predation

Diseases and parasites have been proposed as factors associated with the decline of populations of other species
within the same family of birds as the capercaillie (Tetraonidae) (Obeso et al. 2000, p. 191). In an attempt to determine if parasites were contributing to the decline of the Cantabrian
capercaillie, researchers collected and analyzed fecal samples in 1998 from various localities across the range of this
subspecies. The prevalence of common parasites (Eimeria sp. and Capillaria sp.) was present in 58 percent and 25
percent of the samples collected, respectively. However, both the intensity and average intensity of these parasites were very low compared to other populations of species of birds in the Tetraonidae family. Other parasites were found infrequently. The researchers concluded that it was unlikely that intestinal parasites were causing the decline of the Cantabrian capercaillie.

Inadequacy of existing protection

Although it meets the qualifications, the Cantabrian capercaillie is currently not classified as endangered by the
IUCN. Nor is the species listed under any Appendix of the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES).

This subspecies is currently classified as ‘‘vulnerable’’ in Spain under the National Catalog of Endangered Species,
which affords it special protection (e.g., additional regulation of activities in the forests of its range, regulation of trails
and roads in the area, elimination of poaching, and protection of areas important to young). Although it is
classified as vulnerable, as mentioned above (see Factor B), illegal hunting still occurs.

This subspecies is also afforded special protection under the Bern Convention (Convention on the Conservation of European Wildlife and Natural Habitats
Convention on the Conservation of European Wildlife and Natural Habitats
The Bern Convention on the Conservation of European Wildlife and Natural Habitats 1979, also known as the Bern Convention , came into force on June 1, 1982....

; European Treaty Series/104; Council of Europe 1979).
The Cantabrian capercaillie is listed as ‘‘strictly protected’’ under Appendix II, which requires member states to ensure
the conservation of the listed taxa and their habitats. Under this Convention, protections of Appendix-II species
include the prohibition of:
  • deliberate capture, keeping and killing of the species;
  • deliberate damage or destruction of breeding sites;
  • deliberate disturbance during the breeding season;
  • deliberate taking or destruction of eggs
    Bird egg
    Bird eggs are laid by females and incubated for a time that varies according to the species; a single young hatches from each egg. Average clutch sizes range from one to about 17...

    ;
  • the possession or trade of any individual of the species.

It is difficult to assess the effectiveness of this designation in preventing further loss of Cantabrian capercaillie or its habitat. However, the EU Habitats Directive is a response to the Bern Convention.
The Habitats Directive (92/43/EEC) addresses the protection of habitat and species listed as endangered at the European scale (European Union 2008). Several habitat types valuable to capercaillie have been included in this Directive, such as in Appendix I, Section 9, Forests.

The EU Birds Directive
Birds Directive
The Birds Directive is a European Union directive adopted in 2009. It replaces Council Directive 79/409/EEC of 2 April 1979 on the conservation of wild birds which was modified several times and had become very unclear...

 lists the capercaillie as an Annex 1 species, "subject to special habitat conservation measures in order to ensure their survival.’’
Under this Directive, a network of Special Protection Area
Special Protection Area
A Special Protection Area or SPA is a designation under the European Union Directive on the Conservation of Wild Birds.Under the Directive, Member States of the European Union have a duty to safeguard the habitats of migratory birds and certain particularly threatened birds.Together with Special...

s (SPAs) comprising suitable habitat for Annex I species is to
be designated. This network of SPAs and other protected sites are collectively referred to as Natura 2000
Natura 2000
Natura 2000 is an ecological network of protected areas in the territory of the European Union.-Origins:In May 1992, the governments of the European Communities adopted legislation designed to protect the most seriously threatened habitats and species across Europe. This legislation is called the...

.
As an EU member state, Spain has designated SPAs. The remaining Cantabrian capercaillie populations occur primarily in protected areas in Spain that are part of the Natura 2000 network (for example, Muniellos nature reserve
Muniellos nature reserve
The Muniellos nature reserve protects a wooded area in Asturias, Spain. The area of the nature reserve is 5,488 ha. The main species of tree is oak. In the 1970s timber extraction was forbidden and the woods were preserved as a hunting reserve...

). Management of natural resources by local communities is still allowed in areas designated as an SPA; however, the development of management plans to meet the various objectives of the Reserve network is required.

In November 2003, Spain enacted the ‘‘Forest Law,’’ which addresses the preservation and improvement of the
forest and rangelands in Spain. This law requires development of plans for the management of forest resources, which are to include plans for fighting forest fires, establishment of danger zones based on fire risk, formulation of a defense plan in each established danger zone, the mandatory restoration of burned area, and the prohibition of changing forest use of a burned area into
other uses for a period of 30 years. In addition, this law provides economic incentives for sustainable forest management by private landowners and local entities.

Despite recent advances in protection of this subspecies and its habitat through EU Directives and protection under Spanish law and regulation, illegal poaching still occurs (Storch 2000, p. 83; 2007, p. 96).

Other threats

Suarez-Seoane and Roves (2004, pp. 395, 401) assessed the potential impacts of human disturbances in core populations of Cantabrian capercaillie in Natural Reserves in Spain. They found that locations selected as leks were located at the core of larger patches of forest and were less subject
to human disturbance. They also found that Cantabrian capercaillie disappeared from leks situated in rolling hills at lower altitudes closer to houses, hunting sites, and repeatedly burned areas. Recurring fires have also been implicated as a factor in the decline of the subspecies. An average of 85,652 ha (211,650 ac) of forested area per year over a 10–year period (1995–2005) has been consumed by fire in Spain (Lloyd 2007a, p. 1). On average, 80 percent of
all fires in Spain are set intentionally by humans (Lloyd 2007a, p. 1). Suarez-Seoane and Garcia-Roves (2004, p. 405) found that the stability of Cantabrian capercaillie breeding areas throughout a 20–year period was mainly related to low fire recurrence in the surrounding area and few houses nearby. In addition, the species avoids areas that are recurrently burned because the areas lose their ability to regenerate and cannot produce the habitat the species
requires (Suarez-Seoane and Garcia-Roves 2004, p. 406).

Sources

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