Geographic Range
Tetrao urogallus
(western capercaillies) have a Palearctic distribution. They are a non-migratory
species, although hens with broods seem to use the same route to travel in search
of high-quality insects for multiple years in succession. The northern boundary of
their range reaches as far north as Scandinavia and continuously extends eastward
into eastern Siberia. Further south in temperate Europe, western capercaillie populations
occur in fragments. Prior to the second half of the 20th century, forestry practices
and efforts to improve habitat lead to range expansion and high connectivity. During
this time the connected habitats probably functioned as meta-populations. Western
capercaillie populations are now declining in most of their central European range
due to habitat deterioration and human disturbance.
- Biogeographic Regions
- palearctic
Habitat
Western capercaillies live mainly in taiga and montane ecosystems. They prefer middle
to late successional forests, but are not confined to old forests and have shown tolerance
in the face of forest management regimes. Their habitats are characterized by the
presence of coniferous trees with moderate canopy cover and ground vegetation rich
with ericaceous shrubs, especially bilberry. In winter, western capercaillies almost
exclusively feed on conifer needles. In the spring and summer their diet is supplemented
with buds, flowers, leaves and shrubs.
Western capercaillies require large, contiguous areas of forested habitat. In the
fragmented, temperate, region of central Europe, western capercaillies are exclusively
found in montane regions.
Western capercaillie broods use late succession stage forests, often near peat bogs,
and have been found to stay in natural forests rather than moving into pine plantations.
Brood movements can be extensive, moving almost continuously in 24 hours in search
of high-quality insects.
- Habitat Regions
- temperate
- terrestrial
- Wetlands
- bog
Physical Description
Western capercaillies are large
grouse
that exhibit extreme sexual dimorphism. Males are the largest of the grouse family,
ranging from 4 to 5 kg while females weigh no more than 2 kg. Males are distinguished
by their dark black body color, iridescent blue and green, and vibrant red combs over
the eye. Their fanlike tail is used for displaying to females. Female western capercaillies
are similar to female black grouse (
Tetrao tetrix
), but are larger and have warmer brown coloration on her body. Females are distinguished
by the rosy-rufous color on their throat, breast, and tail.
- Other Physical Features
- endothermic
- homoiothermic
- bilateral symmetry
- Sexual Dimorphism
- male larger
- sexes colored or patterned differently
- male more colorful
- ornamentation
Reproduction
Western capercaillies are considered promiscuous and polygynous. Females show preference
to dominant males on display grounds, and these males account for the majority of
copulations amongst females. There is variation in the literature regarding the lekking
habitat of western capercaillies. Leks are widely thought to be located in old forests.
There are records of individual displaying areas being used for as long as one hundred
years. On the other hand, a study by Rolstad and colleagues suggest that young forest
stands (26 to 46 years) can be suitable for western capercaillie displaying activity.
Mueller (1979 as cited in johnsgard 1983) found that, for fifteen years, a lek was
occupied every spring and fall. In addition, some males occupied the same territories
for as long as four years. His research suggests that the males who rank high amongst
their peers gain respect for their territory. Interestingly, only the high-ranking
males are allowed to copulate without disturbance. Males may hold this rank for several
seasons and were observed performing more than 90% of total copulations in Mueller’s
study. Once male capercaillies obtain a territory, they begin to advertise their fertility.
There are three forms of advertisement including, tree display, flight display, and
ground display. At the break of dawn, male western capercaillies begin their display
by singing from their roost. The flight display is a noisy, drumming flight down from
a perch. Variations of the flight display have also been noted; once on the ground
a western capercaillie may perform “drumming jumps” in which he springs into the air
and beats his wings. The ground display is performed with the “thinned-necked upright”
posture with an extended “beard” and fanned tail. The wings are held downward flashing
the white shoulder patches. Territory defense includes ritualized confrontations that
may lead to intense fights. Rival males may begin with a “display flight” with the
two males bill-to-bill. Neck feathers become increasingly ruffled and the males may
perform the “belching cantus.” As the confrontation becomes more intense, pecking
and blows are exchanged. Although most observed fights seem to end inconclusively,
if there is a true victor, the losing grouse retreats.
- Mating System
- polygynous
- polygynandrous (promiscuous)
Male western capercaillies gather at traditional display grounds. The onset of display
varies in different areas of their range. Territory occupation has been documented
as early as January and seems to end near the onset of molt in adult males. Usually,
nesting sites are not far from lekking grounds and are often times clumped. After
mating, females are left to incubate young on their own in shallow scrapes. Scrapes
are often created at the base of trees under thick cover. Anywhere from 5 to 12 eggs
are laid, a complete clutch takes about 10 days to lay. After the last egg is laid
the female incubates her clutch about 25 days. During incubation the female only leaves
the nest to feed, usually once in the morning and once late afternoon. Western capercaillies
exhibit synchronous hatching. After hatching, the chicks soon leave the nest to feed,
during this time the hen acts as a sentinel and keeps her brood close. Brood movements
are nearly continuous, interspersed with intense foraging in “hot-spots.” Short flight
is observed when broods are 2 to 3 weeks. The chicks fledge at 2 to 3 months old.
- Key Reproductive Features
- iteroparous
- seasonal breeding
- gonochoric/gonochoristic/dioecious (sexes separate)
- sexual
- oviparous
Once mated, females are left to raise their brood alone. Females incubate, tend, and
protect their young until they reach independence.
- Parental Investment
- female parental care
-
pre-fertilization
- provisioning
-
protecting
- female
-
pre-hatching/birth
-
provisioning
- female
-
protecting
- female
-
provisioning
-
pre-weaning/fledging
-
provisioning
- female
-
protecting
- female
-
provisioning
-
pre-independence
-
provisioning
- female
-
protecting
- female
-
provisioning
Lifespan/Longevity
Typical lifespan of western capercaillies is unreported in the literature, but in
the U.K. one wild individual lived to 3.5 years and a captive individual was recorded
living to 18 years.
Behavior
Western capercaillies can be very mobile, yearling males have been reported traveling
up to 7 km in a day visiting leks. Once broods have fledged, females and yearlings
travel and forage in loose flocks while adult males remain solitary.
They forage on the ground and in trees. Males often forage in areas 3 to 4 m below
the apex of trees.
- Key Behaviors
- terricolous
- flies
- glides
- diurnal
- motile
- sedentary
- territorial
- social
Home Range
Home range sizes of western capercaillies are not reported in the literature, but
they are capable of moving over large areas in search of food and mating opportunities.
Communication and Perception
Probably the most fascinating communication center of western capercaillies is the
lekking grounds. Male displays take three distinct forms including tree display, flight
display, and ground display. The tree display is exclusively vocal, the song includes
a metallic "gulping" sound accompanied by "clicking" sounds. During the tree display,
capercaillies assume an upright posture holding their tail low. The flight display
includes a loud flight down from a tree to the ground. The flight down is accompanied
by noisy wing drumming and vocal rattles. Once on the ground capercaillies continue
to make short vertical flights into the air known as "drumming flights." Ground displays
are performed in a "thin-necked up right" posture. The neck is stretched and the head
held upward so that the beard (a thick patch of chin feathers) is protruding. The
wings are held lowered exposing the white shoulder patch.
The song of western capercaillies, also called the "canto," is categorized into four
phases. The first phase consists of a series of doubled-noted clicks, the bill is
opened on the first and closes on the second. The tempo increases until the clicks
merge into a roll and ends in a "cork note" (the sound is similar to that when a cork
is popped from a bottle). The canto is completed when the bird utters 3 to 5 notes
that have been described as a scythe being whetted.
Food Habits
Western capercaillies depend on pine most of the year, although food habits change
seasonally. In the winter they depend on pines, primarily Scots pine (
Pinus silvestris
). They may also feed on the needles and berries of juniper (
Juniperus commuis
) sparingly. In the spring and summer their diet expands to include leaves, stems,
mosses and berries; bilberry becomes an important food plant. Western capercaillies
younger than 20 days have been observed feeding on invertebrates, including spiders,
ants and, beetles.
- Animal Foods
- insects
- terrestrial non-insect arthropods
- Plant Foods
- leaves
- wood, bark, or stems
- seeds, grains, and nuts
- fruit
- flowers
- bryophytes
Predation
The main predators of western capercaillies include carrion crows (
Corvus corone
), red foxes (
Vulpes vulpes
) and
raptors
. Although pine martens (
Martes martes
) have been documented feeding on western capercaillie eggs, their abundance does
not seem to significantly affect breeding success. Western capercaillie females, and
most female
grouse
, have cryptic plumage that blends into ground and shrub vegetation. Females and juvenile
males are mottled and barred with dull white, black and buff. This cryptic plumage
pattern aids in their ability to remain undetected by predators.
- Anti-predator Adaptations
- cryptic
Ecosystem Roles
Western capercaillies, like many ground nesting birds, fall victim to egg predation.
Common nest predators include crows, foxes, raptors, and martens. They sparingly feed
on juniper and other berries, and therefore may aid in seed dispersal, however, there
is no evidence that this dispersal is significant.
- Ecosystem Impact
- disperses seeds
Economic Importance for Humans: Positive
Historically, western capercaillies have been of hunting and cultural importance across
their range. Being the largest grouse species, they have been highly valued as hunting
trophies in central Europe. Since the 1700’s, there have been times when western capercaillie
hunts were reserved for the patricians. According to the Capercaillie Life Project,
they have traditionally been hunted in their spring lek. Highest ranking cocks were
preferred hunting trophies. Research testing the affects of spring and autumn hunts
on populations are needed.
In addition to their economic benefit as a game species, western capercaillies are
enjoyed by bird-watchers and naturalists who may engage in ecotourist activities to
see these, and other, European birds.
- Positive Impacts
- food
- ecotourism
- research and education
Economic Importance for Humans: Negative
Large, late succession forests seem to be favored by western capercaillie populations.
These forests are often cleared for farm land or lumber. If maintaining these forests
for western capercaillie populations provide profits lower than that of farms or lumber,
there is potential for a negative economic effect.
Conservation Status
In 1785 over-hunting and land mismanagement caused western capercaillie populations
be extirpated in Scotland. A Swedish stock of western capercaillie were reintroduced
in 1837, but numbers declined to around 1,000 birds in 1999. Western capercaillie
are still located throughout most of their Palearctic distribution, but serious declines
in western and central Europe have resulted in local extinctions. In the southern
reaches of the range, they are found almost exclusively in montane regions. Some major
causes of population decline include deer fences, which cause large mortalities from
collisions, predation, habitat fragmentation and loss. Western capercaillie require
large areas of contiguous forest and chicks are more sensitive to habitat changes
than adults and require higher quality habitat. The forest areas that best support
western capercaillies also benefit many other native species. For these reasons, they
are often thought of as indicator, or umbrella species. Conservation efforts that
target western capercaillie will likely benefit an array of native species.
Additional Links
Contributors
Laurel Hill (author), Northern Michigan University, Alec Lindsay (editor), Northern Michigan University, Tanya Dewey (editor), University of Michigan-Ann Arbor.
- Palearctic
-
living in the northern part of the Old World. In otherwords, Europe and Asia and northern Africa.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- temperate
-
that region of the Earth between 23.5 degrees North and 60 degrees North (between the Tropic of Cancer and the Arctic Circle) and between 23.5 degrees South and 60 degrees South (between the Tropic of Capricorn and the Antarctic Circle).
- terrestrial
-
Living on the ground.
- taiga
-
Coniferous or boreal forest, located in a band across northern North America, Europe, and Asia. This terrestrial biome also occurs at high elevations. Long, cold winters and short, wet summers. Few species of trees are present; these are primarily conifers that grow in dense stands with little undergrowth. Some deciduous trees also may be present.
- mountains
-
This terrestrial biome includes summits of high mountains, either without vegetation or covered by low, tundra-like vegetation.
- bog
-
a wetland area rich in accumulated plant material and with acidic soils surrounding a body of open water. Bogs have a flora dominated by sedges, heaths, and sphagnum.
- endothermic
-
animals that use metabolically generated heat to regulate body temperature independently of ambient temperature. Endothermy is a synapomorphy of the Mammalia, although it may have arisen in a (now extinct) synapsid ancestor; the fossil record does not distinguish these possibilities. Convergent in birds.
- bilateral symmetry
-
having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.
- sexual ornamentation
-
one of the sexes (usually males) has special physical structures used in courting the other sex or fighting the same sex. For example: antlers, elongated tails, special spurs.
- polygynous
-
having more than one female as a mate at one time
- polygynandrous
-
the kind of polygamy in which a female pairs with several males, each of which also pairs with several different females.
- iteroparous
-
offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).
- seasonal breeding
-
breeding is confined to a particular season
- sexual
-
reproduction that includes combining the genetic contribution of two individuals, a male and a female
- oviparous
-
reproduction in which eggs are released by the female; development of offspring occurs outside the mother's body.
- female parental care
-
parental care is carried out by females
- diurnal
-
- active during the day, 2. lasting for one day.
- motile
-
having the capacity to move from one place to another.
- sedentary
-
remains in the same area
- territorial
-
defends an area within the home range, occupied by a single animals or group of animals of the same species and held through overt defense, display, or advertisement
- social
-
associates with others of its species; forms social groups.
- visual
-
uses sight to communicate
- acoustic
-
uses sound to communicate
- cryptic
-
having markings, coloration, shapes, or other features that cause an animal to be camouflaged in its natural environment; being difficult to see or otherwise detect.
- food
-
A substance that provides both nutrients and energy to a living thing.
- ecotourism
-
humans benefit economically by promoting tourism that focuses on the appreciation of natural areas or animals. Ecotourism implies that there are existing programs that profit from the appreciation of natural areas or animals.
- herbivore
-
An animal that eats mainly plants or parts of plants.
- folivore
-
an animal that mainly eats leaves.
- visual
-
uses sight to communicate
- tactile
-
uses touch to communicate
- acoustic
-
uses sound to communicate
- chemical
-
uses smells or other chemicals to communicate
References
AnAge, 2014. "Tetrao urogallus" (On-line). AnAge: The Animal Ageing and Longevity Database. Accessed February 26, 2014 at http://genomics.senescence.info/species/entry.php?species=Tetrao_urogallus .
Bajc, M., M. Cas, D. Ballian, S. Kunovac, G. Zubic, M. Grubesic, P. Zhelev, L. Paule, T. Grebenc, H. Kraigher. 2011. Genetic differentiation of the western capercaillie highlights the importance of south-eastern Europe for understanding the species phylogeography. PLoS ONE , 6: e23602.
Butchart, S., J. Eckstrom, A. Symes. 2010. "LC Western Capercaillie Tetrao urogallus " (On-line). Birdlife International. Accessed February 14, 2013 at http://www.birdlife.org/datazone/speciesfactsheet.php?id=295 .
Catt, D., D. Baines, N. Picozzi, R. Moss. 1998. Abundance and distribution of capercaillie Tetrao urogallus in Scotland 1992-1994. Biological Conservation , 85: 257-267.
Dunsmore, J., T. Poole. 2004. "The Bird" (On-line). Capercaillie Life Project. Accessed March 01, 2013 at http://www.capercaillie-life.info/index.php .
EU Wildlife and Sustainable Farming Project, 2009. "Capercaillie Tetrao urogallus Fact Sheet" (On-line pdf). Accessed February 13, 2012 at http://ec.europa.eu/environment/nature/natura2000/management/docs/Tetrao%20urugallus%20factsheet%20-%20SWIFI.pdf .
Johnsgard, P. 1983. The Grouse of the World . United States of America: University of Nebraska Press.
Picozzi, N., D. Catt, R. Moss. 1992. Evaluation of capercaillie habitat. Journal of Applied Ecology , 29: 751-762.
Robinson, R. 2005. "Capercaillie Tetrao urogallus [Linnaeus, 1758]" (On-line). BirdFacts: profiles of birds occurring in Britain & Ireland (BTO Research Report 407).. Accessed February 26, 2014 at http://blx1.bto.org/birdfacts/results/bob3350.htm .
Rolstad, J., E. Rolstad, P. Wegge. 2007. Capercaillie Tetrao urogallus lek formation in young forest. Wildlife Biology , 13 (Suppl.1): 59-67.
Segelbacher, G., J. Höglund, I. Storch. 2003. From connectivity to isolation: genetic consequences of population fragmentation in capercaillie across Europe. Molecular Ecology , 12: 1773-1780.
Segelbacher, G., S. Piertney. 2007. Phylogeography of the European capercaillie (Tetrao urogallus) and its implications for conservation. Jounal of Ornithology , 148: 269-274.
Storch, I. 2000. Grouse . IUCN, Gland, Switzerland and Cambridge, UK and The World Pheasant Association, Reading, UK: IUCN Publications Service Unit.
Storch, I. 1995. Annual Home Ranges and Spacing Patterns of the Capercaillie in Central Europe. The Journal of Wildlife Management , 59/2: 392-400.
Storch, I. 2002. On Spatial Resolution in Habitat Models: Can Small-scale Forest Structure Explain Capercaillie Numbers?. Ecology and Society , 6/1: 6. Accessed February 14, 2013 at http://www.ecologyandsociety.org/vol6/iss1/art6/ .
Summers, R., R. Green, R. Proctor, D. Dugan, D. Lambie, R. Moncrieff, R. Moss, D. Baines. 2004. An Experimental Study of the Effects of Predation on the Breeding Productivity of Capercaillie and Black Grouse. Journal of Applied Ecology , 41: 3.
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