Closely related to T. urogalloides, with which very commonly hybridizes where ranges overlap in Siberia (in one area up to 12% of males reported to be hybrids); hybrids with Lyrurus tetrix also common and with Lagopus lagopus occasional. Subspecies name crassirostris replaces major, as latter preoccupied. Many of proposed races clearly intergrade, and sometimes reduced essentially to two, with nominate in N and taczanowskii in S & E (these two differ clearly, latter having largely white belly, vent and “trousers”, but they are linked by very extensive intergradation zone#R); this option, however, probably oversimplistic. At least three phylogeographical studies found evidence for treating Pyrenean and Cantabrian populations as a separate evolutionary unit from all other populations#R; one of these studies, however, found close similarities between populations from Balkans and Pyrenees#R. Otherwise, rudolfi sometimes included in crassirostris, obsoletus in kureikensis, and lonnbergi, karelicus and pleskei in nominate. Other proposed races are lugens (from Finland), a presumed variant of nominate; hiomanus (thought to be from Baltic countries), included in crassirostris; and grisescens (extreme SE European Russia), subsumed within uralensis. Full review desirable. Thirteen subspecies currently recognized.
80–115 cm, female much smaller, 59–64 cm#R; male 3300–4300 g#R (up to 6500 g), female 1440–2500 g#R. Large, very dark grouse, mostly slate-grey with fine vermiculations, blackish on head and neck; breast glossy greenish black; wings dark brown; white carpal patch, and variable amount of white on upper and undertail-coverts and underparts; tail long and rounded (18, sometimes 24 rectrices)#R; bill ivory white; combs scarlet. Female barred and mottled black, grey and buff; larger than that of Lyrurus tetrix, with longer, more rounded rufous tail, and a large rusty patch on breast. First-winter male similar to adult but smaller, with narrower tail feathers and smaller and duskier bill; in both sexes first-winter has outer two primaries speckled reddish brown. Juvenile plumage (short-lived)#R shows pale buff shaft-streaks on mantle, shorter and duller rufous tail with buff tip, and female is more coarsely barred with black than same-age male, which has grey wash to uppertail-coverts#R. Races separated mainly on darkness of plumage and size, but there is extensive individual and clinal variation, e.g. in extent and saturation of white on belly in males and degree of pale upperparts spotting in females#R. Race lonnbergi is small and males are especially dark, whereas race karelicus is paler and has well-developed white on belly. Race cantabricus is smaller than most, except aquitanicus, which differs from other W populations in that females are much darker and more heavily barred, with buff (rather than whitish) underparts; race major has extensively dark underparts and upperparts, while males have warm brown wings, but male rudolfi is even darker, and shorter-tailed, while race pleskei is difficult to distinguish from any of the races whose ranges it directly abuts; race obsoletus is one of the largest and darkest taxa, with dull, dark brown upperparts in male, which also has very small white patches on belly; race kureikensis is generally duller than the last subspecies, with cold grey neck, rump and flanks (like next race), paler back, and smaller size; male of race volgensis has quite pale grey head, neck and rump, and extensive grey-and-white marbling on posterior underparts; race uralensis is palest race; and male of race taczanowskii also has many pale grey feathers on head, neck and rump, and extensive white on belly, while female is yellowish brown above and whitish below#R.
Song (given by male) rather complex and audible over up to 200 m, commences with low-intensity ‘tapping phase’ during which emits disyllabic clicks, “TE-lip”, “Kikop”, followed by ‘drum-roll’ during which clicking notes are given much more rapidly, terminating in the ‘whetting phase’, characterized by a strangled, pig-like squealing sound, rendered “tzjiTHEthethetzjiTHEthethe”#R, the whole performance lasting c. 5·5 seconds and given frequently, up to 7–8 times per minute during twilight and first hour or so of daylight. Western birds (nominate race, major, aquitanicus and cantabricus) also make popping sounds (audible over 300 m) during central drum-roll#R, which develops with age, being loudest in old males and weak and indistinct in first-year males. The birds also perform a ‘flutter-jump’ display, starting on ground and reaching up to 2 m above it, during which very low sounds are produced (infrasound), but it is currently unknown whether these are used in communication between males and females#R. Male also utters harsh “koorKRERKkoroor” during ‘face-off’ with other male or when otherwise agitated, e.g. by human presence, as well as hissing or grunting clicks towards rivals or perceived predators#R. Although displays and therefore vocal sounds are concentrated in spring (as early as Feb)#R, males can display during summer, autumn and even winter, albeit in latter season only very rarely. Female only really vocal during spring, when gives weak braying sound on arriving at male display area, a steadily repeated, chuckling “kok-kok”, likened to that of Ring-necked Pheasant (Phasianus colchicus), in response to male’s display, and various other calls including when with young#R. Chicks give rather plaintive mewing sounds.
Forest and woodland, mainly coniferous (especially Pinus sylvestris, but also Picea, Abies, etc.) or mixed coniferous deciduous; also isolated broadleaved forests (e.g. beech Fagus sylvatica and oak Quercus petraea, Q. pyrenaica)#R#R in some parts of range, e.g. Cantabrian Mts (N Spain) and S Urals. Studies in Scotland have shown that some areas of otherwise suitable Pinus sylvestris forest are rendered unsuitable by high monoterpene levels in the needles, the production of which is genetically controlled, apparently due to counter-feeding by herbivores such as T. urogallus#R. Prefers extensive areas of old, shady forest, often with damp soil and interspersed by bogs, areas of peat or glades, and a dense undergrowth of ericaceous plants (Vaccinium, Calluna), but with the canopy neither too open or closed#R. Old-growth forest with strong component of bilberry (Vaccinium myrtillus) in understorey is strongly selected in Norway#R. In winter, may select more open forest, at least in N, while during summer denser forest with abundant fruit bushes are preferred, especially by broods and moulting birds. Reaches 2000 m in Pyrenees#R, where females regularly frequent alpine scrubland with their nestlings#R, as well as in Altai Mts, and 1500 m in Carpathians.
Food and feeding
During winter mainly pine needles, constituting up to 100% in Finland and Scotland, mostly P. sylvestris, but also P. cembra, P. uncinata, P. sibiricus, etc., in N Europe favouring the needles of diseased trees around bogs because of their higher nutritional value (more protein and less calcium), and up to 90% in Black Forest#R, but autumn-released birds in Harz Mts, Germany, fed on spruce (Picea abies, 34%), bilberry (Vaccinium myrtillus, 26%) and herbs (20%), whereas the diet of tose released in spring was dominated by spruce (56%) and grass (20%)#R. Tree-feeding in winter can last for five months in N, until absence of snow permits feeding on ground. In S, winter diet more varied (although at a microscale may be still dependent on just one or two species), e.g. in Cantabrian Mts, holly (Ilex aquifolium) leaves and birch (Betula) buds during snowy periods, with rockrose (Halimium lasianthum) a major food resource in autumn and winter and eaves, buds and acorns of Pyrenean oak (Quercus pyrenaica) throughout year#R, as well as buds of blaeberry/bilberry (Vaccinium myrtillus) and heaths (Erica)#R#R. Daily food intake in winter is typically c. 440–550 g for males (fresh weight). Inter-seasonal and summer food includes needles, leaves, stems and berries of a variety of plants, including juniper (Juniperus communis), blaeberry, whortleberry (Vaccinium uliginosum), crowberry (Empetrum nigrum), cloudberry (Rubus chamaemorus), cow wheat (Melampyrum pratense), woodrush (Luzula), sedges (Carex), horsetails (Equisetum), mosses, etc., with high protein foods especially important to females for egg production; in Black Forest, beech (Fagus) was main food plant in May (68%), leaves and stems of bilberry were also important in spring (maximum 29% in Jun), in summer and early autumn, they mostly fed on herbs (63% in Jul), grasses (one-third in Sept and Oct) and berries (20% in Jul and Aug)#R. Insects (mainly Formicidae, Coleoptera, Diptera and especially Lepidoptera larvae in Scotland)#R#R#R, spiders and molluscs are important only for chicks, until an age of c. 4 weeks#R, with animal matter always constituting less than 10% of adult diet, even during moulting period. Among more unusual foods are apples in an orchard#R. Forages mainly on ground between spring and autumn, but in tree canopy during winter#R. Females and young form groups of up to 50 in autumn and winter#R (formerly 100, with male-only flocks of up to 70 in past), at which time birds may roost in holes in the snow like some other grouse#R, provided it is deep enough, even consuming snow, apparently for its water content, while at other times of year the species is said to drink twice daily. In places where the species occasionally enters the tundra, e.g. N Urals and Kola Peninsula, it sometimes feeds in company of Lagopus muta.
Lays mainly in May (mid Apr to mid Jun); varies with latitude#R. Promiscuous (or polygynous, with some males briefly possessing harems)#R; males form ill-defined leks (mean number of males per lek 4·8 in Estonian study)#R spaced at c. 1·9–2·4 km intervals#R, at which activity peaks for just 3–4 weeks, although overall display period can last from midwinter to late summer#R (see Voice). Areas used for lekking may be of long-standing tradition, even for 100s of years. Each arena measured 12–67 ha in Estonian study, depending on number of males present#R, exceptionally up to 150 ha elsewhere, while in French Pyrenees males attend smaller leks in more fragmented forest landscapes#R and in SE Europe arenas are typically just 1–2 ha, with up to five males, whereas largest leks in present day may be attended by up to 70 males. Females principally visit leks between dawn and 09:00 hours, although males lek at any time of day#R. The number of males present at an individual lek appears to be determined by habitat quality#R; at Estonian leks trees mostly 81–126 years old, with a height of 10–18 m, and usually no undergrowth#R. Male activity and behaviour at leks shows a hierarchial structure according to age#R. Nest (constructed by female)#R is shallow depression, scantily lined with grass, pine needles, twigs or some feathers, in dense cover, often at base of tree (rarely in old nest of large bird, placed low in tree)#R. In Bavarian Alps, females strongly favour edge habitats for nesting#R. Normally 6–9 pale yellow eggs with some buff to reddish-brown markings#R (4–12, mean 6·5 eggs in Pyrenees and 7·25 in Soctland#R, while elsewhere reports of 12–18 eggs may indicate two females laying in same nest)#R#R, laid on alternate days (exceptionally up to four days apart), mean size 57·3 mm × 41·5 mm#R, mass c. 53 g, but size varies with age of female and regionally; incubation c. 24–29 days by female alone#R, starting before last egg is laid; eggs hatch synchronously#R; downy chick (hatch weight 36–45 g) lacks brown on crown and has characteristic black V mark on forehead; flight feathers and upperwing-coverts start to develop at hatching, and chick able to fly reasonably well at 2–3 weeks (and short distances when just 4–5 days old); first primary shed at 14–17 days#R. Sex ratio on hatching favours females in several regions, including Scotland#R, N & E Finland#R and Russian Karelia, but males frequently outnumber females among adults, even in some of same areas, and percentage of males increases with annual breeding success#R. Nests occasionally parasitized by Phasianus colchicus and Lyrurus tetrix#R. Clutches that hatch, 84–94% for different years in Finland, where another study revealed 60% of eggs laid produced chicks that were still alive in late Aug; in Bavarian Alps, eggs in two-thirds of nests hatched, and one-fifth of chicks survived until autumn#R. In Spanish Pyrenees, multi-year study at various sites found that breeding success varied from 0·2–1·8 young per female per annum#R, while in Scotland 11 of 20 nests were predated at egg stage, most by pine martens (Martes martes)#R. Females may be extremely sensitive to disturbance during incubation period#R and rates of nest predation by mammals can be very high (66%) but are lowest in edge habitats#R. Re-nesting common, with up to 87% (mean 36% in one study) of females that lost first clutch electing to attempt second nest, and this can have a significant effect on overall breeding success#R. Heavy and prolonged rainfall during hatching period, when chicks still small, can be highly detrimental to their survival#R, while wild boars (Sus scrofa) locally can very negatively effect nesting success#R. Sexual maturity in first year, but males do not mate before three and it is probable that only first-year females above a certain mass threshold actually nest. Annual survival rate c. 60% in adult males, but lower in adult females, and mortality of adults of both sexes highest in winter, with lower peak for males in spring#R; one survived to 13·5 years old; in captivity may live up to 18 years or more. Captive-bred young released in autumn were heavily predated by foxes, martens and Northern Goshawks (Accipiter gentilis)#R and efforts to reestablish populations in Germany through translocations and introductions have failed to become self-sustaining#R.
Mainly sedentary, but often with local movements in winter in response to feeding requirements, e.g. in N Siberia, regularly moves from deciduous to coniferous woodland for winter, sometimes in the past in flocks of up to 50 individuals, although also singly. Females tend to move farther than males, as well as moving earlier (older birds first, young females second, thereafter old males and young males last), and same pattern is replicated among young in Scotland, where females also disperse longer distances (up to 30 km), in either their first autumn or next spring#R. Also some irregular movements, occasionally in excess of 1000 km, which may be eruptive; wandering birds even enter villages, where recorded crashing into buildings. However, even in S Siberia, the species is much less mobile, with longest movement in this region just 40 km and most birds being resighted within 6 km of their original locality, with few individuals of either sex wandering far from the lek. In NW Russia, there is evidence that youngsters emigrate from native forest to logged areas, with a reverse movement of mature males attempting to join large leks in native forest areas#R. Brood home ranges in Bavarian Alps averaged 148 ha, between hatching and late summer#R, that of males in the early breeding season c. 47·5 ha#R, while in Russia and SE Norway males moved on average 2·3 km from their leks once display had ceased, only exceptionally travelling further than 6–7 km#R.
Status and conservation
Not globally threatened (Least Concern). Cyclical population changes probably linked to food quality, specifically bilberry (Vaccinium myrtillus) production in some areas#R. Still occupies most of original range, although serious declines in W & C Europe have led to extinctions in Ireland (principally due to habitat loss)#R, Great Britain (lost from England by 1665)#R, Belgium, most of France and Germany, Hungary, locally in Czech Republic#R and many other areas, with dramatic declines registered in Poland#R, Estonia (4500–5000 individuals in mid 1990s)#R, Lithuania and Slovakia; now mainly confined to mountains and N. Race cantabricus of Cantabrian Mts considered Endangered in Red Data Book of 1978/79 and remain so#R; c. 600 males in 1980, but between 1981 and 2003 the range of this population shrank by 66% (from 2,070 km² to 693 km²) and the population by c. 70% (based on counts of singing males)#R, while at least some subpopulations are apparently becoming genetically isolated#R and smaller forest patches have been abandoned during recent decades#R#R; however, improved and more complete censusing methods suggests overall population is currently c. 1000 birds#R. Genetic isolation is also increasing elsewhere through the species’ W & C European range#R, thus maintaining habitat connectivity between subpopulations in regions such as the Alps is vital#R. Largest numbers in Russia, where as many as 6,000,000–7,000,000 were said to survive in 1960s, subsequently just 1,500,000 in 1970s, but the total has since been revised upwards again, to c. 2,200,000–3,000,000 individuals. In Pyrenees, c. 2500 birds estimated in Spain (and population at least locally stable, e.g. within Parc Natural de Cadí-Moixeró)#R, and c. 4000 males in France#R; in Alps, recently extinct in France (elsewhere in the country c. 340 adults in the Jura#R, 200 males in the Vosges and a reintroduced population in the Cévennes of at most 50 individuals)#R, at most c. 1100 males in Switzerland (where population has halved during last c. 30 years)#R, 6500–9000 birds in Italy, and c. 10,000 males in Austria; just 750–1200 territories in Germany#R. In Scotland, where extinct from 1785 (and now Red-listed)#R, successfully reintroduced in 1836–1837#R and by 1862 there were 2000 individuals on the same estate in Perthshire#R; however, countrywide numbers estimated at c. 2200 birds in early 1990s#R#R, c. 1075 in winter 1998/99#R, 1980 birds in winter 2003/04#R#R, c. 1285 in 2009/10#R and is still declining, with just 202 lekking males in 2012#R and 193 in 2013#R, and climate change postulated to be playing an important role in this process#R, although changes in silviculture, increased predation, overshooting, overgrazing by red deer (Cervus elaphus) and collisions with fences are also implicated#R. Also declining, though generally more secure, in Fennoscandia, where during 1960s there were an estimated 300,000–400,000 males in Norway (but had declined by c. 35% by 2000s, especially in S & N)#R and 600,000 in Finland. There are an estimated 10,000 birds in Romania#R. In Ukraine, occurs in Poles’ye zone and Carpathians, where common and fairly numerous at end of 19th century; has declined rapidly since 1960s, due to deforestation, amelioration of climate, pesticides, disturbance and poaching; total population now reckoned to number 4400–5300 birds; protected in three nature reserves and suffers no hunting pressure. In addition to Scotland, the species has also been successfully reintroduced to N Kazakhstan, in 1965, following its extirpation there in the early 20th century. Many estimates of densities available, ranging from 0·1 males/km² in primeval forest of Bialowieza (NE Poland) to 24 birds/km² in natural pine forests in Scotland, but most typically c. 0·5–1·0 males/km². Still commonly hunted (even during breeding season)#R, except in SW & C Europe#R, with annual bags as follows: in Sweden c. 16,000 birds in 1978–1980; in Norway c. 13,000 in 1980, but c. 40,000 around 1960; and in Finland 14,500–33,000 in 1969–1976, but up to 104,000 during early 1960s (numbers taken are usually cyclical, with more being hunted during periods of greater abundance due to breeding success, etc.)#R. Main threat is destruction or alteration of woodland, with fires being especially deleterious, as pines burn more easily. Afforestation has locally fostered some recoveries, but modern forestry practices generally detrimental#R. Other factors possibly involved in declines include excessive shooting, disturbance (e.g. development of ski facilities#R and other winter recreation activities#R), collisions (especially of juveniles) with high-tension powerlines#R and in some areas fences#R, predation (e.g. by foxes)#R, pollution (acid rain) and climatic changes, e.g. in Scotland#R and former East Germany#R. Suggested guidelines for buffer zones around leks and breeding areas have been made in the UK#R and it has been suggested that this capercaillie would make a candidate umbrella species for the conservation of taiga forests#R.
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Winterlosung einer Auerhenne Tetrao urogallus im Alpstall [Dropping of a Capercaillie female Tetrao urogallus in a cattle shed].Orn. Beob.
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