Lagopus lagopus, commonly known as willow ptarmigan, is found within North America in northern Alaska, Victoria Island, Melville Island, Banks Island, Boothia Peninsula, Southampton Island, Baffin Island, South-Eastern Alaska, central British Columbia, Alberta, Saskatchewan, Manitoba, central Ontario and Quebec and Newfoundland. They are also found in Greenland, the United Kingdom, Europe, Russia, Scandinavia and northern Mongolia. Most willow ptarmigan migrate between their winter habitat and breeding range. Although their breeding range is typically lower in both latitude and elevation than their wintering range, most willow ptarmigan inhabiting North American, European and Scandinavian regions stay within a 100 km radius of their summer habitat. It has also been observed in Alaska and British Columbia that males and females of the species often occupy different geographic regions during the winter, with the females migrating to more southerly latitudes than the males.

Biogeographic Regions: nearctic (Native ); palearctic (Native )

Other Geographic Terms: holarctic

occurs (regularly, as a native taxon) in multiple nations

Canada

Origin: Native

Regularity: Regularly occurring

Currently: Present

Confidence: Confident

Type of Residency: Year-round

United States

Origin: Native

Regularity: Regularly occurring

Currently: Present

Confidence: Confident

Type of Residency: Year-round

Global Range: Circumpolar. Alaska, northern Canada (north to low arctic islands), northern Europe, and northern Asia. In North America may wander irregularly south, during the winter, to Michigan, Wisconsin, casually to Montana, North Dakota, and Maine.

Willow ptarmigan have chunky bodies, ranging from 28 to 43 cm long, and weighing and average of 0.57 kg. They have short legs with feathered toes, a feature that enables them to navigate through loose snow. They have short tails (with an average of 118 mm or more for males and 116 mm or less for females) and wings (when folded, an average of 190 mm or more for males and 190 mm or less for females).

Willow ptarmigan molt seasonally. Their winter plumage is usually complete by November, depending on the geographic region of the population. It consists of white feathers covering the entire body except the tail, which remains black. However, the tail is often covered by other feathers on the body, making the bird appear completely white. When the spring arrives, the females’ plumage becomes a mottled brown and ochre, while the males have a rusty hazel or chestnut coloring with darker brown barring on the entire body except for the wings and tail. They also have red “combs” on their eyes which are generally more visible during the warmer months. During the fall, the plumage of both male and female willow ptarmigan becomes lighter, with the females’ plumage becoming more gray and white. The males keep the dark barring pattern but take on more ochre tones.

Red grouse populations of Scotland and Ireland do not develop the same white winter plumage as willow ptarmigan populations living in more northerly areas of Europe and North America (which are covered by snow for much of the year). Instead, they retain a plumage that is predominantly dark reddish-brown. This is likely due to the fact that populations in Ireland and Scotland are exposed to a milder maritime climate than populations living inland and further north. Some studies have also found that there is a possible trade off between white winter camouflage and the thermal benefits of keeping a darker plumage for better heat absorption. However, it has also been found that white plumage has physical characteristics that allow for slightly better insulation. For this reason, more studies must be done to determine whether or not white plumage is thermally disadvantageous.

Range mass: 430 to 810 g.

Average mass: 570 g.

Range length: 280 to 432 mm.

Average wingspan: 61 cm.

Other Physical Features: endothermic ; homoiothermic; bilateral symmetry

Sexual Dimorphism: male larger; sexes colored or patterned differently

Length: 38 cm

Weight: 601 grams

• Terrestrial

Willow ptarmigan inhabit primarily subalpine and subarctic zones. During the summer, they can be found in areas where lush vegetation and a moist environment are present, including arctic valleys, treeline areas, marshy tundra and coastal regions. In Alaska and northern Canada, they are commonly found in areas containing patches of dense vegetation, especially those in which willow (Salix species) or birch (Betula species) shrubs are present. As cooler weather approaches, willow ptarmigan seek more protected areas. This often involves moving further into valleys or more densely vegetated areas within their breeding zone. It has also been observed that during the winter, females and juveniles tend to move into boreal forests in some areas, while the males remain in a subalpine habitat.

Red grouse, Lagopus lagopus scotica, is a subspecies that inhabits Scotland and Ireland. Scottish and Irish populations of red grouse inhabit areas in which heather is readily available. This plant is not only required for shelter and nesting, but also provides a valuable source of food to the population, as it is rich in phosphorus and nitrogen. Although the general area they inhabit is referred to as 'heather moorland', this category can be further broken down into five groups based on specific characteristics of the heather present in that area: dry heath, wet heath, degraded heather, blanket bog and rough moorland grazing. Scottish and Irish red grouse populations generally inhabit areas 150 m above sea level; however, populations have been found to live around sea level and inhabit lowland blanket bogs in northwest Ireland.

Range elevation: 600 to 1980 m.

Habitat Regions: temperate ; polar ; terrestrial

Terrestrial Biomes: tundra ; taiga ; savanna or grassland

Comments: Open alpine and arctic tundra, especially in areas heavily vegetated with grasses, mosses, herbs and shrubs, less frequently in openings in boreal coniferous forest (AOU 1983). Common especially in thickets of willow and alder (National Geographic Society 1983). Nests on the ground, on tundra, beaches, or near marshes (Terres 1980). The nest is a scrape lined with grasses, moss, and feathers, usually located near a bush or tuft of grass.

Non-Migrant: Yes. At least some populations of this species do not make significant seasonal migrations. Juvenile dispersal is not considered a migration.

Locally Migrant: Yes. At least some populations of this species make local extended movements (generally less than 200 km) at particular times of the year (e.g., to breeding or wintering grounds, to hibernation sites).

Locally Migrant: No. No populations of this species make annual migrations of over 200 km.

Mainly permanent resident but somewhat migratory (Godfrey 1966). During winter may descend to intermontane lowlands; may migrate to areas south of treeline; males tend to remain in alpine fringe habitats or on arctic tundra, females tend to move into forested areas (see Johnson and Herter 1989).

Willow ptarmigan generally maintain diets that contains 13 to 18% protein, 10 to 20% fiber, 5 to 10% lipids and up to 50% carbohydrates. However, during the winter their diets consist of less protein, but is much more abundant in fiber and carbohydrates. During the summer months, willow ptarmigan consume a variety of different foods, including willow (Salix species) buds, grass shoots, flowers, seeds, berries (including Vaccinium angustifolium, Vaccinium oxycoccos, Vaccinium vitis-idaea and Empetrum nigrum), insects, caterpillars and beetles. As winter approaches and the availability of insects and caterpillars decreases, the amount of berries, seeds and buds consumed increases. During the winter, food is often more difficult to find and willow ptarmigan must dig through the snow to find sustenance. Their winter diet is composed mainly of willow buds and twigs, as well as some birch (Betula species). Up to 94% of the winter diet may come from willow, with up to 80% being from a single Salix species. The adaptation of digestive enzymes and intestinal microflora to provide nutrition and energy from a single food source is unusual; however, studies suggest that willow ptarmigan are well adapted for obtaining nutrition and energy from a variety of different diets, and that although willow is a primary food source, the species is capable of deriving their nutrition from other sources.

Contrary to North American willow ptarmigan populations, Scotland's red grouse population (a subspecies of willow ptarmigan) has a diet consisting of mainly heather. This accounts for 95% of their diet during fall and midwinter months, falling to 90% with the approach of spring. During this time, the increased availability of Erica tetralix and Erica cinerea make up the remaining 10% of the diet. It has also been found that during nesting season, red grouse tend to select for heather rich in nitrogen and phosphorus.

There are typically two peak feeding times for willow ptarmigan during the winter months. During the morning, feeding occurs for approximately 2 hours, with the same time period being followed for evening feeding. It has been observed that as the daylight hours increase, feeding time also increases.

Willow ptarmigan chicks consume mainly caterpillars, insects, flowers and seeds, largely due to their need for a diet high in protein in order to facilitate extremely fast growth. Willow ptarmigan usually feed in flocks. The size of the flock generally increases or decreases depending on the availability and amount of food present.

Animal Foods: insects

Plant Foods: leaves; wood, bark, or stems; seeds, grains, and nuts; fruit; flowers; bryophytes

Primary Diet: omnivore

Comments: Summer diet consists of leaves and flower buds of willows, alders, and birches; fruits of blueberry, cranberry, crowberry; insects. Winter diet mainly catkins and buds of trees and bushes (Terres 1980).

Willow ptarmigan are predators of various insect and beetle species. They may be infected with flukes (Trematoda species), tapeworms (Cestoda species), roundworms (Nematoda species) and ectoparasites. Many parasites are thought to decrease the body weight of willow ptarmigan, which can lead to death if the weight loss is severe enough. Although this has not been formally observed in North America, populations of red grouse in Scotland have experienced significant decline due to body parasites.

As well as being a predator to many invertebrates, willow ptarmigan are also prey to a number of larger bird and mammal species, including hooded crows, ravens, magpies, red foxes, gyrfalcons, wolverines, wolves and Arctic foxes.

Commensal/Parasitic Species:

• flukes (Trematoda)
• tapeworms (Cestoda)
• roundworms (Nematoda)
• Amyrsidea lagopi
• Lagopoecus affinis
• Goniodes lagopi
• chigger mites (Trombicula waynensis)
• fleas (Ceratophyllus garei)

Willow ptarmigan have many known predators, many of which prey solely on eggs or willow ptarmigan chicks. It has been observed that predation on willow ptarmigan nests is much more common by avian predators than by mammalian predators.

In response to predators, willow ptarmigan will often freeze in a crouching position with their heads lowered and extended. They may perform a diversionary walk to distract predators from the nest or their mate, and may also feign injury by moving slowly along the ground while quivering their wings. This movement makes the ptarmigan appear as though it is dragging itself.

One of the most important anti-predator adaptations of willow ptarmigan is plumage that seasonally changes with the environment. As the snow melts and the surroundings change color, moulting from white to mottled brown occurs very quickly to ensure optimal camouflage. The same pattern is followed for each seasonal change, with the plumage rapidly changing to match the environment. It has also been observed that female willow ptarmigan will choose feeding areas in which they will be better camouflaged rather than ones that offer better nutritional benefits.

Several known predators of North American willow ptarmigan populations include: hooded crows, ravens, magpies, red foxes, pine martens, mink, short-tailed weasels, least weasels, gulls, northern harriers, golden eagles, bald eagles, rough-legged hawks, gyrfalcons, peregrine falcons, northern goshawks, snowy owls, wolverine, wolves, Arctic foxes, lynx and polar bears.

Known predators of Scottish red grouse populations include: red foxes, wildcats, golden eagles, hen-harriers, sparrow-hawks and common buzzards and rough-legged buzzards. Merlins and short-eared owls have been known to kill young red grouse (up to the age of about 8 weeks) however, they are not important predators for adult red grouse.

Known Predators:

• hooded crows (Corvus cornix)
• ravens (Corvus corax)
• magpies (Pica pica)
• red fox (Vulpes vulpes)
• pine martens (Martes martes)
• mink (Neovison vison)
• short-tailed weasels (Mustela erminea)
• least weasels (Mustela nivalis)
• gulls (Larus)
• northern harriers (Circus ganus)
• golden eagles (Aquila crysaetos)
• bald eagles (Haliaeetus leucocephalus)
• rough-legged hawks (Buteo lagopus)
• gyrfalcons (Falco rusticolus)
• peregrine falcons (Falco peregriunus)
• northern goshawks (Accipiter gentilis)
• snowy owls (Nyctea scandiaca)
• wolverines (Gulo gulo)
• wolves (Canis lupus)
• Arctic fox (Vulpes lagopus)
• lynx (Felis lynx)
• polar bears (Ursus maritimus)

Anti-predator Adaptations: cryptic

Family group stays together until late fall; may join other groups and form winter flocks. Cycles of low/high population fluctuations occur but not well understood (Terres 1980). Female breeding territory averaged 2.5-3 ha in British Columbia. Male breeding territories ranged from 1-8 hectares in British Columbia; polygynous males had territories averaging 5.3 hectares, monogamous males had territories averaging 3.9 hectares, and unmated males had territories averaging 2.9 hectares (Hannon and Dobush 1997).

Several different communication sounds have been observed in willow ptarmigan and at least 11 different calls have been observed. Both sexes have territorial calls that are similar. However, ‘flight-song’, ‘rattle’, ‘kohwa’ and ‘aroo’ are only heard in males. ‘Koks’, ‘ko-ko-ko’ and ‘krrow’ sounds are heard in both males and females during threat, mating or territorial situations. These sounds may also be used to stay in contact with a mate or offspring. Females will ‘purr’ or ‘moan’ to communicate with their chicks. Both sexes will ‘hiss’ or ‘moan’ in defense of their offspring or mates. Males give calls that are noticeably stronger and more modulated than females' calls. Like all birds, willow ptarmigan perceive their environments through auditory, visual, tactile and chemical stimuli.

Communication Channels: visual ; acoustic

Willow ptarmigan have an extremely high population turnover, with an annual mortality rate of 60% to 72%. This high mortality rate means there is significantly less competition for territory during breeding seasons than in other species. The primary cause of death in willow ptarmigan populations is predation, whereas death due to weather conditions or food shortage is rarely observed. However, it has been observed that in Scottish willow ptarmigan (red grouse) populations, hens’ access to sprouting vegetation noticeably influences the viability of chicks. Chick weight and egg size also play a role in the survival of Scottish willow ptarmigan chicks both in captivity and in the wild. The longest survival record of a male specimen is 9 years, and the longest survival recorded for a female is 8 years (both recorded in British Colombia). These survival records are based on banded birds returning to breeding areas.

Willow ptarmigan, like many Arctic species, undergo cyclical population fluctuations, with peaks every 9 to 10 years. It is thought that weather conditions and therefore the availability and quality of food may contribute to these fluctuations.

Range lifespan

Status: wild: 9 (high) years.

Territorial establishment of the male willow ptarmigan occurs during early spring, with the males arriving about 2 weeks before the females. During the period of territorial establishment, males become intolerant of other males establishing territory. Breeding territories are usually smaller when the population is larger. Polygynous males also have larger territories than monogamous males.

There have been many different documented courtship displays by willow ptarmigan. Some of these include Tail-Fanning, Waltzing, Rapid-Stamping, Bowing and Head-Wagging. Most of these behaviors are displayed by the males of the species. When Tail-Fanning occurs, the male cocks his tail, sometimes fanning it, and lowers his wings to scrape the ground as he approaches a female. This behavior may lead into Waltzing, in which the male drapes his wings around the female and slowly circles her using short, high steps. He may also tilt his tail and angle his body towards her. Rapid-Stamping occurs when the male runs toward the female while slightly fanning his tail, arching his neck and holding his head low with the beak open. Bowing is a display in which the male raises and lowers his head while holding his body low to the ground. Head-Wagging can be performed by males or females. One member of the mating pair will crouch by its mate, while wagging its extended head side to side. This display is usually performed alternately. Females may also show submissive behavior, after which mating usually occurs.

Male willow ptarmigan develop nuptial plumage during April or May. This is a key visual signal for female attraction. The nuptial plumage is predominantly bright brown and white, and the supraorbital comb also becomes more prominent. Females are generally more attracted to males with larger territories, combs and more vigorous courtship displays. However, females may leave the territory if the display is too aggressive. Mating usually occurs after the female displays a submissive posture.

Male willow ptarmigan are usually monogamous within a season. Approximately 83 to 86% of pairs will remate the following year if both members of the mating pair survive. However, 5 to 20% of male willow ptarmigan are polygynous (mating with more than one female), and usually have larger territories than unpaired or monogamous males. Few female willow ptarmigan are polygynous. This is possibly due to the higher predation on nests of polygynous females. Monogamous females are accompanied by their mate for around 94% of their active diurnal period, which contributes to the slightly higher survival rate of monogamous females’ broods over broods of polygynous females. It has been suggested that most willow ptarmigan are generally not polygynous because hens are capable of preventing other females from settling with their mate.

Mating System: monogamous ; polygynous

Willow ptarmigans mate seasonally during the spring and summer. After mating, the ptarmigan pair excavates a bowl-shaped nest in the ground. These bowl nests are often sheltered from most sides, but one side of the nest is usually left unprotected as a means of escape. They are usually lined with leaves, grass and feathers, and range from 15 to 20 cm wide and 8 to 16 cm deep. Willow ptarmigan generally lay between 4 and 14 eggs which are usually incubated for 20 to 23 days in late June to early July. When a developed chick is ready to hatch and makes the first cracks on it's eggshell (also known as 'pipping'), eggs usually hatch within a few hours. Hatching occurs during daylight hours. Willow ptarmigan hatchlings are precocial and usually weight about 15 g. They grow extremely fast and leave the nest to feed the same day they hatch. They no longer rely on their parents for warmth after 7 to 10 days, and are able to fly within 10 to 12 days. Chicks are usually adult-sized by autumn and are able to fend for themselves by this time (about 5 to 7 months of age). Both male and female willow ptarmigan are capable of breeding as yearlings however, not all yearling males will acquire territory or a mate.

Breeding interval: Willow ptarmigan breed once a year.

Breeding season: Mating occurs during the spring and summer months.

Range eggs per season: 4 to 14.

Range time to hatching: 20 to 23 days.

Average birth mass: 15 g.

Range fledging age: 10 to 12 days.

Range time to independence: 5 to 7 months.

Average age at sexual or reproductive maturity (female): 1 years.

Average age at sexual or reproductive maturity (male): 1 years.

Key Reproductive Features: seasonal breeding ; gonochoric/gonochoristic/dioecious (sexes separate)

Willow ptarmigan are the only members of the grouse family in which males exhibit significant parental care. A male willow ptarmigan stays by the nest to defend his clutch and female during the incubation period, and will defend the brood until the chicks are independent (around autumn). Females usually brood the chicks however, lone males are capable of successfully raising chicks. It has also been observed that parents and replacement males may adopt unrelated chicks. A study in which male mates were removed from the territory after incubation had begun showed that 50% of the previously unpaired males and 5% of the paired males joined the widows to give care to and defend the chicks. Observations show that replacement males are generally younger, more vocal and sit closer to the nest than parental males. Possible motivations for unpaired males to adopt unrelated chicks include the possibility of mating with the female should the first brood be unsuccessful and gaining mating experience in order to attract a mate in subsequent breeding seasons.

Parental Investment: precocial ; male parental care ; female parental care ; pre-fertilization (Provisioning, Protecting: Female); pre-hatching/birth (Protecting: Male, Female); pre-weaning/fledging (Provisioning: Male, Female, Protecting: Male, Female); pre-independence (Protecting: Male, Female)

Breeding begins in mid-May in south to early June in north; in northern Canada, date of clutch initiation apparently is related to timing of snow melt (Hannon et al. 1988). Clutch size is 4-17 (usually 6-11) Incubation lasts 20-26 days (Harrison 1978); in northern Canada, incubated 20-22 days after clutch completion (Hannon et al. 1988). Nestlings are precocial and downy. Young are guarded by both parents, can fly at about 12-13 days, independent at about 2 months. Mature in 1 year. Monogamous.

Willow ptarmigan are extremely widespread and their conservation status is of least concern. However, as of 2009, they were classified as critically imperiled in the province of Alberta, Canada. This does not follow the general trend for Canadian willow grouse populations, as populations in other provinces are classified as being either secure, apparently secure or unknown/under review.

Several new possible threats to ptarmigan populations have emerged in recent years. Although ecotourism has not directly altered ptarmigan populations, it has inadvertently contributed to the destruction of already fragile habitats that support this species. Pollutants are also becoming a more serious problem to many Arctic populations, including willow ptarmigan.

The Scottish red grouse population has shown significant decline in recent years. This is thought to have occurred primarily due to the reduction of heather abundance (which are a main food source for red grouse), caused by afforestation and farming.

US Migratory Bird Act: no special status

US Federal List: no special status

CITES: no special status

IUCN Red List of Threatened Species: least concern

Canada

Rounded National Status Rank: N5 - Secure

United States

Rounded National Status Rank: N5 - Secure

Rounded Global Status Rank: G5 - Secure