Second part: Behaviour

 

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Steve Garvie
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Tom Grey
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Patrick Ingremeau
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René Lortie
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Tom Merigan
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Yves Thonnérieux
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Ingo Waschkies
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Text and illustrations by Nicole Bouglouan

Sources :

HANDBOOK OF THE BIRDS OF THE WORLD Vol 2 by Josep del Hoyo-Andrew Elliot-Jordi Sargatal - Lynx Edicions - ISBN: 8487334156

THE HANDBOOK OF BIRD IDENTIFICATION FOR EUROPE AND THE WESTERN PALEARCTIC by Mark Beaman, Steve Madge - C.Helm - ISBN: 0713639601

FIELD GUIDE TO THE BIRDS OF NORTH AMERICA - National Geographic Society - ISBN: 0792274512

BIRDS OF THE GREAT BASIN – by Fred A. Ryser - Univ of Nevada Pr -ISBN: 0874170796

THE COMPLETE BOOK OF BRITISH BIRDS – Written by “Royal Society for the Protection of Birds” experts - Préface de Magnus Magnusson - Michael Cady- Rob Hume Editors - ISBN: 0749509112

BirdLife International (BirdLife International)

The IUCN Red List of Threatened Species

Nature Works

Wikipedia, the free encyclopaedia

Animal Diversity Web (University of Michigan Museum of Zoology)

ARKive (Christopher Parsons)

Audubon

 

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FAMILY TETRAONIDAE

Grouses

This natural and fairly homogeneous group belongs to the order Galliformes, and is sometimes regarded as a subfamily within the Phasianidae.  
But, in spite of several morphological and behavioural characteristics shared with the Phasianidae, the Tetraonidae species show some differences such as feathered nostrils and tarsi, lack of spur, toes with scales on the sides adapted to climbing branches and walking on snow, shorter tail and heavier body with duller plumage.

During the breeding season, the Tetraonidae males develop prominent, bright red or yellow comb above each eye, but not red bare face or wattles like in pheasants. They also have coloured patches of bare skin on the neck sides, inflated and used as air-sacs that produce low-pitched booming calls while males are displaying. This is another difference between Tetraonidae and Phasianidae.

However, these families are closely related and unfortunately, both are famous gamebirds and hunted in numerous countries.    

The family Tetraonidae includes 18 species and 7 genera. Males have more contrasted plumage and are larger than females which have very cryptic colours, making them almost invisible while incubating on the ground. The chicks usually have typical brown to chestnut crown often bordered by black line, and the juvenile is a smaller version of the adult female.

SPECIES' DESCRIPTION

The genus Falcipennis includes two species, the Siberian Grouse (F. falcipennis) and the Spruce Grouse (F. canadensis).
Both species are very close to each other especially by plumage, courtship behaviour, ecology and some other behavioural aspects. Males are bulky, with almost same pattern on brown or grey upperparts and the black underparts are conspicuously spotted or barred white. The black bib is bordered with narrow white stripe, and head, neck and upperbreast form a long grey or brown hood finely barred whitish, similar in both species. The red comb is followed by a short post-ocular white stripe. The only difference is found on the uppertail on which the Spruce Grouse shows white tips on tail-coverts and rectrices.

Spruce Grouse

F. canadensis

Male

Both females have cryptic plumage, but the Spruce Grouse female has tawny neck and breast and has darker plumage overall than female Siberian Grouse.

The Siberian Grouse has restricted range in far eastern Russia where it occurs in coniferous forests with spruce and fir, and frequents thick understorey of moss and berries. This species is evaluated as Near Threatened, due to intensive exploitation of forest by humans.

The Spruce Grouse is found from Alaska, through Canada to Nova Scotia, and in some northern regions of the United States.
This species frequents boreal coniferous forests and during summer, it can be seen in rich understorey of low shrubs and blueberries. This one is evaluated as Least Concern and populations are not globally threatened.  

Both species are largely sedentary in their respective ranges, only performing short dispersion.

Spruce Grouse

F. canadensis

Female

The Dusky Grouse (D. obscurus) is the only member of the genus Dendragapus. This one is larger with a length of 47-57 centimetres. The male is mostly grey to slate-grey and has long, square tail with grey terminal band. During the displays, we can see the yellow combs becoming orange to bright red. This species has airs-sacs on neck sides, buff-yellow in coastal races and purplish in inland males. On the underparts, the undertail-coverts are spotted white and exposed when the tail is cocked. 

The female is browner with white barring on top of head, nape, scapulars, breast and flanks.

The Dusky Grouse is found in the highland regions of North America, from SE Alaska and NW Territories, S to California, Arizona, New Mexico and Colorado. It is visible from sea-level up to 3600 metres of elevation. It frequents subalpine coniferous forest with firs, and breeds in drier and more open forested habitat. It moves to higher elevation in winter.
This species is common throughout its range with variable densities according to urbanisation and cultivated areas. It is evaluated as Least Concern.    

Dusky Grouse

Dendragapus obscurus

Inland male displaying

The genus Lagopus includes today four species such as the Willow Ptarmigan (L. lagopus), the Rock Ptarmigan (L. mutus), the White-tailed Ptarmigan (L. leucurus) and the Red Grouse (L. scotica).     
Except for the Red Grouse which is today a full species and formerly a subspecies of Lagopus lagopus, males in winter are all white. However, both males Willow Ptarmigan andRock Ptarmigan have black outermost rectrices conspicuous in flight, whereas the male White-tailed Ptarmigan is all white, including the tail.  

© Nicole Bouglouan
Tail pattern

During winter, the three all white species have white plumage overall with some different head pattern. White-tailed and Willow Ptarmigans show narrow red eye combs and white head in both sexes, whereas the Rock Ptarmigan male has a black line from bill through the eye, and red comb too. The female has white head.
All have black bill, dark eyes and feathered white legs and feet.

Rock Ptarmigan - L. mutus

Male in winter -----------------------------------------------------------------------Male in late summer

The breeding plumage is mottled and barred blackish, brown or rufous on upperparts, head, neck, upperbreast and flanks. Usually, belly and vent are white, as part of the upperwing. The red comb is becoming brighter and larger in males.
The male Willow Ptarmigan has more rufous colours than other species.

Willow Ptarmigan

L. lagopus

Male in summer

Spring and fall are moulting seasons, giving the birds a patchy white appearance. Females have duller and more cryptic plumage than males, with much smaller combs.

Rock Ptarmigan

L. mutus

Female in summer

The Willow Ptarmigan is found mainly in the Arctic tundra, and in wooded areas with alder and willow trees, and berry-bearing shrubs, grasses and mosses. It ranges from Alaska, throughout northern Canada and Newfoundland, Scandinavia and Siberia. According to the place, this species is sedentary (Scandinavia) and partially migratory elsewhere, moving into forested areas.
It is widespread and common or abundant in its wide distribution. Some fluctuations have been reported due to unfavourable habitat in some parts of the range. However, the Willow Ptarmigan is not currently threatened.

Willow Ptarmigan

L. lagopus

Male in late spring

The Rock Ptarmigan can be found in all countries circling the North Pole, including Greenland, Iceland, Europe, Asia and N America, and in mountainous areas of S Europe. It frequents rocky country in tundra with sparse vegetation, and further south, it is found in mountains. During winter, it may move to less snowy areas in order to find some ground vegetation to feed.
The Rock Ptarmigan’s populations make limited movements, mainly altitudinal and related to local weather conditions and food resources. Arctic populations perform greater movements but mainly nomadic. This species is not currently threatened.

Rock Ptarmigan

L. mutus

Male in spring

The White-tailed Ptarmigan is locally common on rocky mountain slopes and high meadows. It is found in the mountainous areas of W United States, Canada and Alaska where it is permanent resident, only performing limited altitudinal movements in winter. The species is not currently threatened.

The Red Grouse has reddish-brown plumage all year round, including during winter, but the underparts show white streaks and spots more conspicuous than above. Legs and feet are covered in whitish feathers. The short tail is blackish.

Red Grouse

L. scotica

Male

The Red Grouse occurs in treeless tundra and open upland heather moors in Scotland, Wales, England and Ireland, and may frequent farmland during harsh winters. This species only performs some altitudinal movements and is mainly sedentary in its range.
In spite of some declines due to changes in its habitats, the Red Grouse is not currently threatened.  

Red Grouse

L. scotica

Female

The genus Tetrao contains four species, two capercaillies and two black grouses.
The capercaillies are the largest ones. We can find two species, the Western Capercaillie (Tetrao urogallus) and the Black-billed Capercaillie (Tetrao parvirostris).
The two other species are the Eurasian Black Grouse (Tetrao tetrix) and the Caucasian Black Grouse (Tetrao mlokosiewiczi). Both are smaller than capercaillies, with peculiar lyre-shaped tail. Formerly, they were placed in genus Lyrurus.  
The Tetrao species are forest dwellers. They differ mainly in male’s plumage, tail shape and body size.

The Western Capercaillie is the largest species of the family Tetraonidae with a length of 80-115 centimetres. The male has dark plumage with blackish head and neck. The body is dark grey to blackish with variable white spots or streaks and the breast shows dark glossy blue-green tinge.
Unlike black grouses, the tail is relatively long and rounded with light whitish patches often exposed during the displays.

The bill is creamy-white. The eyes are brown with red comb above. The legs and feet are dark grey with feathered tarsi.

Western Capercaillie

Tetrao urogallus

Male

The female is smaller. She has cryptic plumage overall, barred and mottled black, grey and buff. We can see a rusty breast patch. The tail is rounded and rufous.

The Western Capercaillie is found in the northern parts of Europe, W and C Asia and some isolated populations in N Spain and C Europe. It has been introduced in Scotland. It frequents coniferous forest and woodland and isolated broad-leaved forests according to the range. During summer, it is mainly found in dense forest with fruit bushes, whereas in winter, it prefers more open forest.    
This species is sedentary in its wide range, only performing local movements according to the food resources. 

Populations appear stable but serious declines are reported in W Europe. However, this species is not currently threatened.

Western Capercaillie

Tetrao urogallus

Female

The Black-billed Capercaillie is slightly smaller with a length of 68-97 centimetres. This one is darker than previous, with blackish plumage and black underparts. But there are white tips to secondaries and white markings on scapulars, greater wing-coverts and uppertail-coverts.
The black bill is smaller than in Tetrao urogallus. The eyes are brown with small red combs above. Legs and feet are dark brown with feathered tarsi.

Black-billed Capercaillie

Tetrao parvirostris

Male

The female is much smaller. She is almost similar to the previous but darker with more heavily barred pattern.

The Black-billed Capercaillie breeds in larch taiga forest in E Russia and parts of N Mongolia and China. It can be found both in plains and in mountains, and according to the region, it can be found in mixed deciduous-coniferous forests.
It usually prefers more open landscapes than Tetrao urogallus.
Populations have decreased, and formerly, this species was very common and much hunted in some areas. But currently, it is not globally threatened.   

The other species of genus Tetrao, formerly genus Lyrurus, have different appearance with glossy black plumage overall and lyre-shaped tail.

The Eurasian Black Grouse is the largest, with a length of 60 centimetres. It has glossy black plumage with blue-green tinge, and white rounded carpal patch, wingbar and undertail-coverts. The long black, lyre-shaped tail has long outer rectrices, curved outwards. It has red combs above the eyes.
Neck and head feathers become barred or mottled brown and there is some white on throat in late summer.

Eurasian Black Grouse

Tetrao tetrix

Male

The female is largely brown and barred black. She has shorter, slightly forked tail when closed. 

The Eurasian Black Grouse is sedentary and breeds across northern Eurasia, mostly in boreal regions. Some regular winter migrations are reported in parts of Siberia.
It frequents variable habitats, but it usually occurs between forest and open areas, both in lowlands and mountains up to 2000-3000 metres of elevation. It breeds in areas with high variety of plants.
In spite of several declines due to deforestation and agriculture expansion, and to local excessive hunting and disturbances, this species is not currently threatened.

Eurasian Black Grouse

Tetrao tetrix

Male

Eurasian Black Grouse

Tetrao tetrix

Female

The Caucasian Black Grouse is slightly smaller with a length of 53 centimetres. It has glossy green-black plumage overall. It lacks the white markings on wings and undertail, and appears darker than previous. The long, lyre-shaped tail shows shorter outer rectrices curving downwards. It has red eye-combs too.

The female is grey-brown and narrowly barred black. She has almost square-shaped tail.

The Caucasian Black Grouse is found in Great and Little Caucasus Mountains, S into NE Turkey and NW Iran. It only performs altitudinal movements.  
It lives between the upper limit of mountain forests with spruce and birch, and subalpine meadows with rhododendron and thickets.
This species is currently evaluated as Near Threatened. Increase of hunting, grazing, fragmentation of the habitat and wood cutting are the main threats for this grouse.  

The genus Bonasa had formerly only one species, the Ruffed Grouse (Bonasa umbellus) from N America. Now, two Eurasian species are included, the Hazel Grouse (Bonasa bonasia) and the Severtsov’s Grouse (Bonasa sewerzowi). Both were placed in the genus Tetrastes and are very similar with more primitive features than the Ruffed Grouse. This one is quite distinct with thick ruff on neck sides, and may represent a recent divergence.
They are today in the same genus Bonasa.

Ruffed Grouse

Bonasa umbellus

Female

The Ruffed Grouse is fairly large with a length of 43-48 centimetres. The cryptic plumage makes the bird almost invisible on the ground. This grouse has short crest, erectile black ruffs on neck sides, and fan-shaped tail with dark subterminal band in male. On the head, the small red eye combs become larger in spring. Like other Tetraonidae species, tarsi are feathered.  

Ruffed Grouse

Bonasa umbellus

Male

The female is fairly similar, but eye combs are not developed. She has shorter tail with less visible dark band.  

The Ruffed Grouse is resident in its wide range, from W Alaska, across Canada to Appalachian Mountains.
It frequents several types of woodlands, from boreal forest to Pacific coast rainforest, and dry deciduous and mixed woodlands. The brood needs thick vegetal cover such as brushy areas.  
In spite of the usual threats such as fragmentation of the habitat and hunting, this species is not currently threatened.   

Ruffed Grouse

Bonasa umbellus

Female

The Hazel Grouse is smaller with a length of 35-40 centimetres. Male has brownish-grey plumage with fine dark barring on crown, neck, mantle and uppertail-coverts. The underparts are barred or spotted, dark grey to ochre. The chin is black, forming a bib, bordered with white. The tail is fairly long and rounded, with white fringe and black subterminal band. The central pair of rectrices is brown.
On the head, there is a small erectile crest. We can see a small red eye comb. The bill is greyish-black.   

The female is brownish and has whitish throat, finely barred brown and edged white.

The Hazel Grouse is sedentary in its wide range. It breeds throughout N Eurasia to Hokkaido, and in C and E Europe. Females and young birds usually disperse farther.
Populations are declining, probably due to changes in the habitat, and in spite of heavy hunting in several regions, the species is evaluated as Least Concern by BirdLife International.

Hazel Grouse

Bonasa bonasia

Male

The Severtsov’s Grouse is very similar to the previous but slightly smaller with a length of 34 centimetres. It has more conspicuously barred black upperparts and chestnut on upperbreast. Feathers are finely edged white, giving scaling effect on underparts. The head is chestnut and the black bib is slightly larger than in hazel Grouse. Small red eye combs are present.

The female shows almost similar plumage pattern that Hazel Grouse, but she is more rufous and darker overall.

The Severtsov’s Grouse is probably sedentary in its restricted range in mountains of C China, NW Yunnan and N Sichuan.
It frequents montane forests with birch and coniferous trees and cover of moss on forest floor above 1000 metres, but up to 4000 metres of elevation in Tibet.
This species inhabits in very restricted range, suffering habitat loss, egg collecting and hunting. The Severtsov’s Grouse is classified as Near Threatened.

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The four remaining species of genera Centrocercus and Tympanuchus are closely related to open habitats with herbaceous vegetation, often in dry temperate climates. They are found in North America, whereas in Eurasia, the similar niches are occupied by the Otididae.

The genus Centrocercus contains only one species, the Sage Grouse (Centrocercus urophasianus). This is a large bird with a length of 66-76 centimetres, and the male is much larger than the female. It has greyish-brown plumage with white-edged feathers making fine scaling on the upperparts. On the underparts, the male has black throat and foreneck, separated by V-shaped white line. There is a large white ruff on breast, and two large greenish cervical sacs, mainly visible when the bird is displaying. It has fairly small yellow eye combs. The belly is black. The long tail is made with stiff and pointed feathers. Undertail-coverts are dark with white tips. Underwing-coverts are white.
The stout bill is black. The eyes are brown. Legs are feathered to the toes, giving scaled effect on tarsi.

Sage Grouse

Centrocercus urophasianus

Male

The female is smaller. She has similar plumage but she lacks the white ruff and the air sacs, and throat and foreneck are greyish-brown streaked dark brown. She has black belly too.

The Sage Grouse lives mainly in Sagebrush (Artemisia tridentata) of genus Artemisia. This plant belongs to the grouse’s world, providing food, shelter, protection and nesting cover to the bird. It frequents foothills and plains.

Sage Grouse

Centrocercus urophasianus

Female

This species is mostly sedentary in its range in Western North America, from SE Alberta and Saskatchewan and SW North Dakota, S to E California, Nevada, Utah and W Colorado. They only perform altitudinal movements.  
Sage Grouse’s populations are threatened by habitat loss due to fragmentation and degradation. Human developments degrade the sagebrush ecosystem with in addition, the degradation due to overgrazing, fires, herbicides and farming. Industrialisation is also an important threat for the species which is now listed as Near Threatened.

Sage Grouse

Centrocercus urophasianus

Male

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The genus Tympanuchus contains three species, the Greater Prairie-chicken (Tympanuchus cupido), the Lesser Prairie-chicken (Tympanuchus pallidicinctus) and the Sharp-tailed Grouse (Tympanuchus phasianellus).  
The first two species are very similar in appearance, the Lesser Prairie-chicken being a smaller version of the Greater Prairie-chicken, with slightly paler plumage and reddish cervical sacs opposed to yellowish in the Greater. Both are mainly brown and heavily barred white, black and buff on upperparts and underparts. The short tail is blackish and rounded.
On the head, the long feathers on neck sides or “pinnae” are about 70mm long in male Greater, and shorter in male Lesser. Both have yellow to orange eye combs. Cervical sacs are golden yellow in Greater, and orange-red in Lesser.

Greater Prairie-chicken

Tympanuchus cupido

Male

Females are smaller than males, with shorter pinnae and considerably smaller and paler eye combs and air sacs. They have barred tail instead blackish.

The Greater Prairie-chicken is mostly sedentary in its restricted range in Kansas, Nebraska and South Dakota (race pinnatus), and some places in SE Texas (race attwateri).
It frequents open grasslands and oak woodlands. This species prefers the scrub oaks mixed with prairies, and sandy soils. They need areas of native plants for breeding and roosting. The leks are situated in short-grass areas, on elevated grounds. They nest in tall grassy places.
The Greater Prairie-chicken is threatened by habitat loss and hunting, and introduced mammals. They live in fragmented and restricted habitats where prairies are replaced by croplands. Hunting continues in four states.

The species is classified as Vulnerable on the IUCN list.

Greater Prairie-chicken

Tympanuchus cupido

Male

The Lesser Prairie-chicken is largely sedentary in its range in SE Colorado, SW Kansas, Oklahoma, E New Mexico and NW Texas.
It frequents sandy dry grasslands, sagebrush (Artemisia filifolia) and short-grass prairie areas with shinnery oaks for acorns (Quercus havardii). Displays take place in short-grass prairie country with ridges or other elevated places.
They nest in low shrubs or among clumps of grass. Later, the brood moves to thicker cover.
The Lesser Prairie-chicken’s populations have declined, and although stabilization and even some increase since 1995, they remain threatened by effects of drought and habitat loss.
This species is currently retained as Vulnerable by BirdLife International.

Lesser Prairie-chicken

Tympanuchus pallidicinctus

Two males and one female

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The third species of genus Tympanuchus, the Sharp-tailed Grouse (Tympanuchus phasianellus), formerly placed in genus Pediocetes, is clearly related to the two previous species. They share features of plumage and also courtship behaviour.
This grouse frequently interbreeds with the Greater Prairie-chicken, giving fertile hybrids.
It is probably the most primitive form of these three birds in terms of morphology.

The Sharp-tailed Grouse is a medium-sized bird, with a length of 40-48 centimetres. It is similar in plumage to prairie-chickens, mainly on upperparts, but it has paler speckled underparts, not barred. Undertail-coverts are pure white.
We can see a small erectile crest and small yellow eye combs. Cervical sacs are pinkish or purple.
But the tail shows different shape, giving the bird its name. The tail is pointed. The central pair of rectrices extends far beyond the others, well visible when the tail is cocked. Prairie-chickens have rounded tail.

Female is similar to prairie-chicken’s. She has less obvious eye combs and cervical sacs, and she is smaller than male.  

The Sharp-tailed Grouse frequents mainly open environments such as grasslands, sagebrush, woodland edges and river canyons. The displays take place in more open and elevated sites with good visibility.
This species has wide range in South-Western Canada and Northern United States. where it is uncommon to fairly common. It is often hunted, and it may be threatened by changes in its habitat, but currently, the species is not endangered or globally threatened.

All these grouses have fairly similar behaviour during courtship, feeding and breeding. To study their habits will be the subject of the second part of this article.

Second part:

Behaviour

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