Western capercaillie

Western Capercaillie Tetrao urogallus distribution map.png

Range of the western capercaillie[2]

Western Capercaillie Tetrao urogallus distribution in Europe map.png

Distribution in Europe[2]

The western capercaillie (Tetrao urogallus), also known as the wood grouse, heather cock, or just
capercaillie /ˌkæpərˈkeɪli/, is a Eurasian member of the grouse family and its largest member. The
largest known specimen, recorded in captivity, had a weight of 7.2 kg (16 lb). Found across Europe
and Asia, this ground-living forest bird is renowned for its mating display. The species shows extreme
sexual dimorphism, with the male twice the size of the female. The worldwide population is in the
category "least concern" of the IUCN,[1] although the populations of Central Europe are declining
and endangered, or already extinct.

Taxonomy

The species was first described by Linnaeus in his Systema naturae in 1758 under its current binomial
name.[3]

Its closest relative is the black-billed capercaillie, Tetrao parvirostris, which breeds in the larch taiga
forests of eastern Russia and parts of northern Mongolia and China.

Male and female western capercaillie can easily be differentiated by their size and colouration. The
cock is much bigger than the hen. It is one of the most sexually dimorphic in size of living bird
species, only exceeded by the larger types of bustards and a select few members of the pheasant
family.

Cocks typically range from 74 to 85 cm (29 to 33 in) in length with wingspan of 90 to 125 cm (35 to 49
in) and an average weight of 4.1 kg (9.0 lb).[5][6][7] The largest wild cocks can attain a length of 100
cm (39 in) and weight of 6.7 kg (15 lb).[8] The largest specimen recorded in captivity had a weight of
7.2 kg (16 lb). The weight of 75 wild cocks was found to range from 3.6 to 5.05 kg (7.9 to 11.1 lb).[7]
The body feathers are dark grey to dark brown, while the breast feathers are dark metallic green. The
belly and undertail coverts vary from black to white depending on race (see below).

The hen is much smaller, weighing about half as much as the cock. The capercaillie hen's body from
beak to tail is approximately 54–64 cm (21–25 in) long, the wingspan is 70 cm (28 in) and weighs 1.5–
2.5 kg (3.3–5.5 lb), with an average of 1.8 kg (4.0 lb).[7] Feathers on the upper parts are brown with
black and silver barring; on the underside they are more light and buffish yellow.
Both sexes have a white spot on the wing bow. They have feathered legs, especially in the cold
season, for protection against cold. Their toe rows of small, elongated horn tacks provide a snowshoe
effect that led to the German family name "Rauhfußhühner", literally translated as "rough feet
chickens".

These so-called "courting tacks" make a clear track in the snow. The sexes can be distinguished very
easily by the size of their footprints.

There is a bright red spot of naked skin above each eye. In German hunters' language, these are the
so-called "roses".

The small chicks resemble the hen in their cryptic colouration, which is a passive protection against
predators. Additionally, they wear black crown feathers. At an age of about three months, in late
summer, they moult gradually towards the adult plumage of cocks and hens. The eggs are about the
same size and form as chicken eggs, but are more speckled with brown spots.

The capercaillie is a non-migratory sedentary species, breeding across northern parts of Europe and
western and central Asia in mature conifer forests with diverse species composition and a relatively
open canopy structure.

At one time it could be found in all the taiga forests of northern and northeastern Eurasia in the cold
temperate latitudes and the coniferous forest belt in the mountain ranges of warm temperate
Europe. The Scottish population became extinct, but has been reintroduced from the Swedish
population; in Germany it is on the "Red list" as a species threatened by extinction and is no longer
found in the lower mountainous areas of Bavaria; in the Bavarian Forest, the Black Forest and the
Harz mountains numbers of surviving western capercaillie decline even under massive efforts to
breed them in captivity and release them into the wild; in Switzerland, they are found in the Swiss
Alps, in the Jura, in the Austrian and Italian Alps. The species is extinct in Belgium. In Ireland it was
common until the 17th century, but died out in the 18th. In Norway, Sweden, Finland, Russia and
Romania populations are large, and it is a common bird to see in forested regions.

The most serious threats to the species are habitat degradation, particularly conversion of diverse
native forest into often single-species timber plantations, and to birds colliding with fences erected
to keep deer out of young plantations. Increased numbers of small predators that prey on
capercaillies (e.g., red fox) due to the loss of large predators who control smaller carnivores (e.g.,
gray wolf, brown bear) cause problems in some areas.

Status and conservation

Footprints of western capercaillie in the Czech national nature reserve Kladské rašeliny
This species has an estimated range of 1,000,000–10,000,000 km2 (390,000–3,860,000 sq mi) and a
population of between 1.5 and 2 million individuals in Europe alone. There is some evidence of a
population decline, but the species is not believed to approach the IUCN Red List threshold of a
population decline of more than 30% in ten years or three generations. It is therefore evaluated as
Least Concern.[1]

As reported by the Spanish researcher Félix Rodríguez de la Fuente in his "Fauna" series, the
northwestern Spanish subspecies T. u. cantabricus—an Ice Age remnant—was threatened in the
1960s by commercial gathering of holly fruit-bearing branches for sale as Christmas ornaments—a
practice imported from Anglo-Saxon or Germanic countries.

In Scotland, the population has declined greatly since the 1960s because of deer fencing, predation
and lack of suitable habitat (Caledonian Forest). The population plummeted from a high of 10,000
pairs in the 1960s to fewer than 1000 birds in 1999. It was even named as the bird most likely to
become extinct in the UK by 2015.

In mountainous skiing areas, poorly marked cables for ski-lifts have contributed to mortality. Their
effects can be mitigated by proper coloring, sighting and height alterations.

Behaviour and ecology

The western capercaillie is adapted to its original habitats—old coniferous forests with a rich interior
structure and dense ground vegetation of Vaccinium species under a light canopy. They mainly feed
on Vaccinium species, especially bilberry, find cover in young tree growth, and use the open spaces
when flying. As habitat specialists, they hardly use any other forest types.

Western capercaillies are not elegant fliers due to their body weight and short, rounded wings. While
taking off they produce a sudden thundering noise that deters predators. Because of their body size
and wing span they avoid young and dense forests when flying. While flying they rest in short gliding
phases. Their feathers produce a whistling sound.

Western capercaillie, especially the hens with young chicks, require resources that should occur as
parts of a small-scaled patchy mosaic: These are food plants, small insects for the chicks, cover in
dense young trees or high ground vegetation, old trees with horizontal branches for sleeping. These
criteria are met best in old forest stands with spruce and pine, dense ground vegetation and local
tree regrowth on dry slopes in southern to western expositions. These open stands allow flights
downslope, and the tree regrowth offers cover.

In the lowlands such forest structures developed over centuries by heavy exploitation, especially by
the use of litter and grazing livestock. In the highlands and along the ridges of mountain areas in
temperate Europe as well as in the taiga region from Fennoscandia to Siberia, the boreal forests
show this open structure due to the harsh climate, offering optimal habitats for capercaillie without
human influence. Dense and young forests are avoided as there is neither cover nor food, and flight
of these large birds is greatly impaired.

The abundance of western capercaillie depends—as with most species—on habitat quality. It is
highest in sun-flooded open, old mixed forests with spruce, pine, fir and some beech with a rich
ground cover of Vaccinium species.

Spring territories are about 25 hectares (62 acres) per bird. Comparable abundances are found in
taiga forests. Thus, the western capercaillie never had particularly high densities, despite the legends
that hunters may speculate about. Adult cocks are strongly territorial and occupy a range of 50 to 60
hectares (120 to 150 acres) optimal habitat. Hen territories are about 40 hectares (99 acres). The
annual range can be several square kilometres (hundreds of hectares) when storms and heavy
snowfall force the birds to winter at lower altitudes. Territories of cocks and hens may overlap.

Western capercaillie are diurnal game, i.e., their activity is limited to the daylight hours. They spend
the night in old trees with horizontal branches. These sleeping trees are used for several nights; they
can be mapped easily as the ground under them is covered by pellets.

The hens are ground breeders and spend the night on the nest. As long as the young chicks cannot fly
the hen spends the night with them in dense cover on the ground. During winter the hens rarely go
down to the ground and most tracks in the snow are from cocks.

Diet

The western capercaillie lives on a variety of food types, including buds, leaves, berries, insects,
grasses and in the winter mostly conifer needles. One can see the food remains in their droppings,
which are about 1 cm (0.39 in) in diameter and 5–6 cm (2.0–2.4 in) in length. Most of the year the
droppings are of solid consistency but, with the ripening of blueberries, these dominate the diet and
the faeces become formless and bluish black.

The western capercaillie is a highly specialized herbivore, which feeds almost exclusively on
blueberry leaves and berries with some grass seeds and fresh shoots of sedges in summertime. The
young chicks are dependent on protein-rich food in their first weeks and thus mainly prey on insects.
Available insect supply is strongly influenced by weather—dry and warm conditions allow a fast
growth of the chicks, cold and rainy weather leads to high mortality.

During winter, when a high snow cover prevents access to ground vegetation, the western
capercaillie spends almost all day and night in trees, feeding on coniferous needles of spruce, pine
and fir as well as on buds from beech and rowan.
To digest this coarse winter food, the birds need grit: small stones or gastroliths which they actively
search for and devour. With their very muscular stomachs, gizzard stones function like a mill and
break needles and buds into small particles. Additionally, western capercaillie have two appendixes
which grow very long in winter. With the aid of symbiotic bacteria, the plant material is digested
there. During the short winter days the western capercaillie feeds almost constantly and produces a
pellet nearly every 10 minutes.

Courting and reproduction

Female in Bavarian Forest, Germany

Tetrao urogallus urogallus—eggs

The breeding season of the western capercaillie starts according to spring weather progress,
vegetation development and altitude between March and April and lasts until May or June. Three-
quarters of this long courting season is mere territorial competition between neighbouring cocks or
cocks on the same courting ground.

At the very beginning of dawn, the tree courting begins on a thick branch of a lookout tree. The cock
postures himself with raised and fanned tail feathers, erect neck, beak pointed skywards, wings held
out and drooped and starts his typical aria to impress the females. The typical song in this display is a
series of double-clicks like a dropping ping-pong ball, which gradually accelerate into a popping
sound like a cork coming of a champagne bottle, which is followed by scraping sounds.

Towards the end of the courting season the hens arrive on the courting grounds, also called "leks",
Swedish for "play". The cocks continue courting on the ground: This is the main courting season.

The cock flies from his courting tree to an open space nearby and continues his display. The hens,
ready to get mounted, crouch and utter a begging sound. If there is more than one cock on the lek, it
is mainly the alpha-cock who engages in sexual intercourse with the hens. In this phase western
capercaillies are most sensitive to disturbances. Even single human observers may cause the hens to
fly off and prevent copulation in this very short time span where they are ready for conception.

There is a smaller courting peak in autumn, which serves to delineate the territories for the winter
months and the next season.

Egg-laying
About three days after copulation the hen starts laying eggs. In 10 days the clutch is full. The average
clutch size is eight eggs but may amount up to 12, rarely only four or five eggs. Brooding lasts about
26–28 days according to weather and altitude.

At the beginning of the brooding season, the hens are very sensitive to disturbances and leave the
nest quickly. Towards the end they tolerate disturbances to a certain degree, crouching on their nest
which is usually hidden under low branches of a young tree or a broken tree crown. As hatching
nears, hens sit tighter on the nest and will only flush from the nest if disturbed in very close
proximity. Nesting hens rarely spend more than an hour a day off of the nest feeding and as such
become somewhat constipated. The presence of a nest nearby is often indicated by distinctively
enlarged and malformed droppings known as "clocker droppings". All eggs hatch in close proximity
after which the hen and clutch abandon the nest where they are at their most vulnerable.
Abandoned nests often contain "caeacal" droppings; the discharge from the hens' appendixes built
up over the incubation period.

Hatching and growth

After hatching the chicks are dependent on getting warmed by the hen. Like all precocial birds, the
young are fully covered by down feathers at hatching but are not able to maintain their body
temperature which is 41 °C (106 °F) in birds. In cold and rainy weather the chicks need to get warmed
by the hen every few minutes and all night.

They seek food independently and prey mainly on insects, like butterfly caterpillars and pupae, ants,
myriapodae, ground beetles.

They grow rapidly and most of the energy intake is transformed into the protein of the flight
musculature (the white flesh around the breast in chickens). At an age of 3–4 weeks they are able to
perform their first short flights. From this time on they start to sleep in trees on warm nights. At an
age of about 6 weeks they are fully able to maintain their body temperature. The down feathers have
been moulted into the immature plumage and at an age of 3 months another moult brings in their
subadult plumage; now the two sexes can be easily distinguished.

From the beginning of September the families start to dissolve. First the young cocks disperse, then
the young hens. Both sexes may form loose foraging groups over the winter.

Predation and hunting

Mammalian predators known to take capercaillie include Eurasian lynx (Lynx lynx) and gray wolf
(Canis lupus), although they prefer slightly larger prey. Meanwhile, European pine martens (Martes
martes), beech martens (Martes foina), brown bears (Ursus arctos), wild boars (Sus scrofa) and red
foxes (Vulpes vulpes) take mostly eggs and chicks but can attack adults if they manage to ambush the
often wary birds.[9][10][11][12][13] In Sweden, western capercaillies are the primary prey of the
golden eagle (Aquila chrysaetos).[14] Large numbers are taken by northern goshawks (Accipiter
gentilis), including adults but usually young ones, and Eurasian eagle-owls (Bubo bubo) will
occasionally pick off a capercaillie of any age or size; they normally prefer mammalian foods.[15][16]
White-tailed eagles (Haliaeetus albicilla) are more likely to take water birds than upland-type birds
but have been recorded preying on capercaillie around the White Sea.[17]

A traditional gamebird, the capercaillie has been widely hunted with guns and dogs throughout its
territory in central and northern Europe. This includes trophy hunting and hunting for food. Since
hunting has been restricted in many countries, trophy-hunting has become a tourist resource,
particularly in Central European countries. In some areas, declines are due to excessive hunting,
though this has not generally been a global problem. The bird has not been hunted in Scotland or
Germany for over 30 years.[18]