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Willow Ptarmigan

A bird species of the grouse family, with a seasonal plumage that is brown in summer and white in winter to blend in with the surroundings, and it is commonly found in tundra and subarctic regions across North America and Eurasia.

Willow Ptarmigan

Scientific name: Lagopus lagopus

Cold and snow and predators in the Willow Ptarmigan’s arctic and subalpine habitat are all dealt with by plumage adaptations including feathered legs that help it to travel in snow, and dramatically different summer and winter plumages that provide year-round camouflage. Willow Ptarmigan typically walk, and seldom fly.

Strongly territorial during the breeding season, Willow Ptarmigan often form winter flocks, sometimes in excess of 1,000 birds. Although capable of breeding at age one, not all males are successful at obtaining a mate.

Photograph © Alan Wilson

Description of the Willow Ptarmigan

BREEDING MALE

willow ptarmigan summer aw

Photograph © Alan Wilson

The Willow Ptarmigan is a large, chunky grouse with a white eye ring and a black tail. Its plumage varies by season and gender. Reddish-brown head, neck, flanks, and back with extensively white wings.  Prominent red eye combs.

Female

willow ptarmigan summer 2
Photograph © Alan Wilson

Mottled brownish plumage with white in the wings.

Seasonal change in appearance

willow ptarmigan winter aw

Winter birds become pure white. Molting birds have an intermediate appearance.

Juvenile

Juveniles resemble adult breeding females.

Habitat

Tundra and willow scrub.

Diet

Buds, leaves, twigs, and seeds.

Behavior

Forages by walking.

Range

Resident from Alaska to eastern Canada.

Fun Facts

During extreme cold, Willow Ptarmigan will burrow into the snow to roost overnight.

Males and some females return to their breeding territories for a brief period of time in the fall.

Vocalizations

Clucking calls and growls are given.

Similar Species

  • Rock Ptarmigan are browner and less red.

Nesting

The nest is a plant-lined depression.

Number: 5-14.
Color: Red to blackish-brown.

Incubation and fledging:  
– Young hatch at 21-22 days.
– Young fledge (leave the nest) shortly after hatching but remain with the adults for some time.

Bent Life History of the Willow Ptarmigan

Published by the Smithsonian Institution between the 1920s and the 1950s, the Bent life history series of monographs provide an often colorful description of the birds of North America. Arthur Cleveland Bent was the lead author for the series. The Bent series is a great resource and often includes quotes from early American Ornithologists, including Audubon, Townsend, Wilson, Sutton and many others.

Bent Life History for the Willow Ptarmigan – the common name and sub-species reflect the nomenclature in use at the time the description was written.

WILLOW PTARMIGAN
LAGOPUS LAGOPUS ALBUS (Gmelin)

HABITS

The willow ptarmigan, with its various so-called subspecies, is of circumpolar distribution, inhabiting the arctic or subarctic regions of both the North American and Eurasian Continents. European writers know it as the willow grouse, applying the name ptarmigan to only the rock ptarmigan and its races. It seems more logical to apply the name ptarmigan to both species that assume a white plumage in winter. The British red grouse is the only species of the genus Lagopus that does not have a white winter plumage; it does, however, have the feathered toes, one of the principal characters of the genus. The following account includes the so-called Alaska ptarmigan.

Spring: As soon as the first bare spaces appear on the sunny slopes of the tundra the ptarmigan begin their spring migration from their winter quarters, among the willows in the sheltered valleys of the rivers and creeks of the interior, to their breeding grounds on the open hills or tundra. H. B. Conover says in some notes he sent to me:

On our sled trip from Nenana to Hooper Bay, this grouse was not encountered until we reached the Kuskokwim Mountains, where we found it very plentiful. It was generally encountered in flocks of 15 to 100 feeding among the dwarf willows. A few males shot on April 5 had an occasional brown feather showing on the head and neck. The morning of April 24 we left Mountain Village on the Yukon and cut across the tundra for Hooper Bay, a journey which took us four days. For the first two of these we encountered ptarmigan everywhere along the willow-bordered sloughs and creeks. The hens were still white, but the heads and necks of the cocks were about a third into the red spring plumage. As we approached the coast and the willows became scarcer, these birds were no longer seen. On May 9 the ground about Point Dall was beginning to show in spots through the snow and the first ptarmigan made their appearance. Two days later they were common. Each male now took possession of a little spot of bare ground, whence he sent out his challenges, com-ere, com-ere, go-bee, gobec. Between calls they would bob their heads as if they were pecking at the ground, or, jumping about 6 feet into the air, glide down to the earth, cackling as they descended. The hens seemed to have but one call, a cackle similar to that of a tame chicken. Often two cocks were seen chasing each other around over the tundra, but only rarely would they seem to stand and fight it out. In the evenings and early mornings these birds were especially noisy, and often It was no great stretch of the imagination, what with the calls of the waterfowl, to imagine oneself In some great barnyard. About this time the Eskimo boys began to range the tundra with their bows and arrows, and many an unwary cock and sometimes a hen were killed by the blunt shafts of these 8 and 10 year olds.

Courtship: Edwyn Sandys (1904) gives the following very good account of this performance:

The love-making of the ptarmigan is not unlike that of the Canada Grouse, or “spruce-partridge.” The males, with their plumage changing from white to the handsome summer dress, strut with all the pomposity of their kind. The red combs over the eyes are swollen and very conspicuous, as the bird struts with head thrown far back, tail raised and spread, and wings trailing. Presently he leaps into the air, raises himself higher and higher with a vigorous flapping, then sails on set wings through a descending spiral, which brings him back to his starting-point. While thus a-wing, he utters a curt, gruff challenge oft repeated, a defiance to all rivals. Again be struts, and again goes into the air, frequently to see male after male arise from near-by stations. While so occupied the birds make considerable noise, the bark-like challenge of other calls being heard for some distance. Meanwhile, the females loiter about in the cover, admiring the efforts of the males, and gradually acknowledging their charms. The Inevitable battles follow: spirited encounters, in which many hard knocks are given, and much pretty plumage marred, until the weaker have been well whipped.

Nesting: Herbert W. Brandt has sent me some elaborate notes on this species. He says of its nesting:

The willow ptarmigan at Hooper Bay is not at all particular as to the location of its home site, for it dwefls impartially from the drift-strewn sea beach to the higher altitudes on the mountains. Down under the protection of a drift log, a clump of grass, a small hush, a mossy hummock, or any screeny object, she scrapes out a cavity to fit her requirements. This she lines more or less with any material at hand, and here she deposits daily her rich crimson egg. During the period of egg accumulation, we found the nest to be covered with surrounding material, because the bird does not begin to incubate until the full complement Is satisfied. When the first eg~ is laid there is but little form to the nest, but as the set progresses, the birds mold it into the proper shape, and by the time incuhation has progressed, the eggs snuggle together in a well-cupped basin. The brooding female spreads her feathers as does the sitting domestic hen, and when her frail body is examined in hand, it seems almost incredible that she can cover so large a clutch of eggs. The extreme measurements of 12 nests examined are: Total height, 5 to 7 inches; inside diameter, 6 to S inches; depth of cavity, 3 to 6½ inches.

F. Seymour Hersey collected some 10 sets of eggs for me between St. Michael and the mouth of the Yukon. They were mostly in fairly open situations on the tundra. One nest was a depression in the tundra moss at the base of a small clump of grass; it was lined with dry grass, leaves, and a few feathers; it measured 7 inches across and 5 inches deep. Another was in a hollow under a dwarf willow on a raised mound on the tundra. Another was in a deep hollow in a wet place on the border of a marsh, a very open situation; it had a heavy lining of dry grass and a few feathers. Some of the hollows were very shallow, not more than 2 inches deep. There were three sets of 7, two of 8, one of 9, three of 10, and one of 11 eggs. George G. Cantwell writes to me of a nest he found on Copper River, Alaska, that was “placed on an open river bar among a light growth of willows, close to a growth of mountain spruce.” Stanton Warburton. jr., tells me of a nest he found late at night, saying:

To preserve them intact for a photo in the morning, I put my khaki coat over the nest and eggs, completely covering them. Then (as an experiment) I wrinkled up the collar by the eggs so that it formed an opening not over 2 Inches high. This slight opening did not expose the eggs to view from any angle, as they were still completely covered. Early the next morning when I approached, the female flushed from right beside my coat. All 11 eggs were now in a new nest just outside my coat, not over a foot from the original location. Durilig the night the female had made a new nest, moved all 11 eggs into it, and recommenced incubation.

MacFarlane mentions in his notes another case where the eggs were probably removed by the birds. 1-le “had reason to know that some, at least, of the nests were used by pt.armigan several seasons in succession.”

Joseph Dixon (1927) writes:

The male ptarmlgan spends the day hiding in little thickets, keeping within 50 or 100 feet of the nest. He has a definite form or nest of his own which he occupies when roosting. One reason for his staying so close Is tile danger of Short-billed Gulls finding the piarmigan’s nest. These egg thieves work in organized gangs, usually three together. One will swoop (lown at the female, trying to make her shift about on the nost so as to oxpOse the eggs. The second or third gull following tries to sill) in and grab an egg. As soon as the gulls appear, the hen ptnrmigan gives a peculiar call for help. Upon hearing tills the cock ptarmigan bursts forth like a rocket and charges the thieving gulls. He doesn’t boat around tile bush but tiles directly at the intruders, knocking them down with the impact of his body . An average cock Willow Ptarmigan at this season weighs 507 grams, while one on the Short-billed Gulls which was shot weighed 358.2 grams. In addition to being one-third heavier than the gull the cock ptarmigan flies much the faster of the two, and when he hits a gull it is almost like a Duck Hawk striking a duck.

Eggs: The willow ptarmigan lays commonly 7 to 10 eggs; as many as 17 have been found in a nest, and in late or second sets there may be only 5 or 6. An egg is laid each day, and incubation does not begin until the last egg is laid. For a description of the beautiful eggs, I can not do better than to quote Mr. Brandt’s remarks, as follows:

Eggs in the same set follow the same type of ground color, the same style of markings, and are nearly uniform In size; and as each bird seems to lay a type of egg individually its own, it may be noted that scarcely any two sets of eggs of this interesting species are exactly alike. In shape the egg is almost always ovate, lint in rare instances tend to short ovate with stubby ends, and nearly elongate ovate ~vhen more slender in shape. The surface of the shell is smooth and often somewhat greasy like that of a duck egg; the texture is hard; and the egg sturdy like its parents. The luster is rather shiny and this apparently increases somewhat as incubation progresses. A study of the coloration of the egg of the willow ptarmlgan is of special Interest because of its change of hue after it is laid. When the egg first appears, the markings are from “ox-blood red” to “scarlet red,” and the whole surface is vivid and moist, and it appears as if it has been dipped in fresh, red paint. This undried pigment is very easily rubbed off, and in consequence, it is unusual that an egg does not show the sign somewhere on the surface of being brushed by the feathers of the parent bird. In fact, these rubbed spots may show distinctly the individual barbs of the ptarmlgan feather that scratched it when it was wet. As the moisture from the egg dries, however, the pigment sets rapidly, and at the same time so darkens, like congealed blood, that by the time the eggs are a few hours old, the brilliant reds turn to blackish brown. Once the pigment becomes dry and sets, it is very durable, and eggshells that have lain out in the weather from the previous year still retain their bold markings.

The ground color is usually inconspicuous, because it Is seldom that more than half of it is visible, and often it is all practically hidden by the overlying spots. The only place that the ground color is prominent is on those areas where there is an aforementioned rubbed spot. This ground color exhibits many variations of the pale creamy tints, such as “ivory yellows’ and “seafoam yellow,” but a few sets are further decorated with paler reds, making the ground color “orange-crimson” to “vinaceons-tawny” and “pecan brown,” while the ground color of one egg is even “ocher red.”

The markings are the richest of any egg we collected at Hooper Bay and are irregularly flecked in profusion all over the surface. In size, these spots range from the finest pepperings to blotches thumb-nail in size, and are all more or less confluent, some so much so that they cover the surface and almost envelope the ground color. When the spots are large, the ground color is often well defined, and then the most handsome effects are produced. If the ground color is distinctly reddish, the surface markings are usually not nearly so numerous, and because of their sparse distribution, the egg then approaches in appearance that of the spruce grouse (Ooaachite.s caaadensis). The markings show almost uniformly blackish brown with a reddish suggestion, yet where the pigment has been scratched very thin, the color is often as light as “maroon” and eve ii garnet brown,” while the deeper colors are “warm blackish brown” and “blackish brown (1).”

The measurements of 250 eggs, in the United States National Museum, average 43 by 31 millimeters; the eggs showing the four extremes measure 48 by 31.5, 46.5 by 34, 39 by 30, and 39.5 by 28 millimeters.

Young: Mr. Conover says in his notes:

Newly hatched young were first found on June 22. The incubation period seems to last about 22 days. A nest found on June 2 with 11 eggs had 12 on June 3, and on being visited on June 25, it was found to be empty. The chicks are very precocious. One day a hen was flushed from a nest containing two eggs and eight youngsters still damp. Hardly had she left when every downy chick scrambled weakly from the nest and attempted to hide in the grass. The minute they were replaced, out they would gu again, until finally they became tired out and stayed in the nest. Toward the end of June broods were constantly encountered about the tundra. Both parents were always with them and the cock was especially combative, although discretion always got the better part of valor. The young after running a few feet would suddenly disappear, whereupon the hen would join the male in threats and attempt to lead one off. It was amusing to imitate the peeping of a chick and watch the cock go into a frenzy, ruffling himself up, making short dashes here and there, and in unmistakable language telling you just what he was going to do if you didn’t get away from his children. After a few minutes of this, both birds would be worn out and would retire a short way to watch for your next move. By July 22 the young were about a third grown and had begun to shed their first brown primaries and grow their new white ones. The adults were then in the midst of shedding their toenails.

Dixon (1927) noticed that the family party traveled as follows:

First came two or three chicks in the thick grass, then the mother surrounded by the other chicks; the cock sometimes led and at other times brought up the rear. I timed them and found that they covered a lineal distance of 45 feet in five minutes. Following this there came a period of rest of five minutes, during which the mother hovered her brood of young. We never saw the cock hover the young; but when one of the chicks became entangled in a network of twig.s he was right there and helped it get free. By noon the ptarmigan family had wandered out in the low bushes 100 yards from where they had hatched. The chicks were now nearly 24 hours old, and all of them were strong and lusty, each able to run about with agility and to secure food for itself. At Copper Mountain, about 4 o’clock In the afternoon of July 12, a family of Willow Ptarmigan came feeding along through the dwarf willows near camp. There were six young about the size of quail. The cock kept a lookout for enemies from elevated positions while the hen herded the young along through the willows. The hen kept up a running conversation with the young as did also the cock. This liaison note was a loud ke-oe,ck, repeated at intervals of from five to ten seconds. The cock’s call was somewhat coarser than that of the hen. I had difficulty in hearing the thin peeping of the chicks at a distance of fifty feet, but it served to keep them together. The young were very active, Jumping up into the willows and catching insects over a foot off the ground.

Plumages: In natal down the young willow ptarmigan has a large patch of “burnt sienna,” bordered with black, on the center of the crown and occiput; the rest of the head and the underparts are colonial buff “; the upper parts are variegated with ” colonial buff,” black, and “cinnamon-rufous,” the last mainly in the center of the back and rump, bordered by broad black bands; a small spot on the lores, a larger auricular spot, and a narrow line behind the eye are black.

The juvenal plumage comes in first on the wings, then on the scapulars and back, while the chick is very small; the juvenal remiges are ” sepia,~~ bordered with ” cinnamon-buff ~ or buffy white, and their coverts are tipped with buff; the last of the down to disappear is on the chin, neck, and belly. The juvenal plumage is at its height before the young bird is half grown, at which time the postjuvenal molt begins, early in August. In full juvenal plumage the feathers of the mantle are black, edged, notched, or barred with “ochraceoustawny,” and with triangular white tips; the breast and flanks are “ochraceous-tawny,” heavily barred or spotted with black or dusky; the belly is buffy or grayish white; the remiges are as stated above, except the outer two, which are the last to appear and are white; the rectrices are black, edged, spotted, or barred with “ochraceoustawny.”

When the young bird is nearly half grown the postjuvenal molt begins by shedding the juvenal remiges, which are replaced by white ones. This is a more or less incomplete molt, involving most of the wings (except the two pairs of outer white primaries), the tail, and a varying extent of the body plumage. In this intermediate, or preliminary, winter plumage, the sexes are still practically alike. There may be only a few scattered, reddish brown, finely vermiculated, or mottled feathers, or the renewal may be nearly complete; the helly, flanks, and legs become white.

This intermediate plumage is worn for a very short time while the young bird is getting its growth; for, during the last of August and through September, a supplementary, partial molt takes place, which completes the change into the white winter plumage. Young birds can be distinguished during their first winter by the outer primaries, which are more worn and often speckled at the tips.

Both young and old birds have a partial prenuptial molt in spring, the date varying greatly with the latitude and season. The reddish spring plumage begins to appear on the head and neck of the male at about the time that bare ground begins to appear in its summer home, from the first to the middle of April in Alaska. The females molt a month or more later. Dr. Jonathan Dwight (1900) says:

Females may now be distinguished with certainty from males for the first time by plumage characters, the barring being coarser and extending to the head, throat, and breast, the feathers of which in the male are reddish brown, chiefly with narrow, dusky terminal bands, and often tipped, on the chin especially, with white. It should be observed that parti-colored feathers basaUy or terminally white, may be assumed at this moult on the internal borders of the sternal band, just as in juvenal dress, the abdominal wedge, flanks, legs, and feet, retaining as a rule the white feathers of the winter plumage. The white remiges and their coverts are always retained and often much of the rest of the wing plumage, the median rows of coverts being the ones renewed if any are. The tail-coverts may be renewed, but the fourteen black rectrices remain.

Dwight says of the next molt:

Even before the nuptial dress is fully acquired the postnuptial moult sets in, beginning a little prior to the postiuvenal and resulting in an intermediate plumage partly white and partly reddish brown which may hardly be told from that of young birds at the same season. It should be observed that the moult of the remiges now includes the two distal primaries which are retained in young birds. Adults, however, seem to be somewhat grayer with finer mottllng or vermiculation, the throat being of a deeper red-brown with less barring than that of young birds. PracticaUy young and old, both males and females, are all indistinguishable except by inconstant differences when clothed by the preliminary winter dress, but their age and sex may usually be told by the left over tell-tale feathers of an earlier plumage.

A supplementary molt early in fall, September and October in Alaska, completes the change into adult winter plumage. Females are indistinguishable from first winter birds, the feathers of the crown being basally gray, whereas in the adult male these are basally black. Winter adults in high plumage often have a decidedly rosy tint, which soon fades in the dry skin.

Food: During summer ptarmigan feed on the tender leaves and flower buds of the willows, birches, and alders, with a fair percentage of berries, such as mountain cranberry, crowberry, blueberries, arbutus, and kinnikinnick. They also eat what insects they can find. Turner (1886) writes:

During the winter these birds subsist on the past year’s twigs of the willow and alder or other bushes. I have cut open the crops of many of these winterkilled birds and found them to contain only pieces of twigs about one-third of an inch long, or just about the width of the gape of the posterior, horny part of the bill, as though this has been the means of measurement in cutting them off. The flesh at this time is dry and of a peculiar taste. In the spring the Ptarxnigans congregate in great numbers on the willow-bushes and eat the tender, swelling buds. The flesh then acquires a bitter, but not unpleasant, taste. As open weather advances they find berries that have remained frozen the entire winter, and tender grass shoots, and later, insects. The young are Insectivorous to a great degree In their youngest days, They consume great numbers of spiders that are to be found on the warm hillsides.

Dixon (1927) sent the stomach of a 5-day-old chick to Washington for examination; it contained 17 yellow caterpillars, 1 spider, 15 Tky8anura, and 15 other insects and larvae, but no vegetable matter.

He says:

The eight chicks foraged In a loose flock covering an area about five feet wide and sIx feet long. They pursue small insects and mosquitoes which they run down or reach up for and pick off the grass. I watched one chick catch a cranefly and after hammering and pecking at it awhile he concluded that it was too tough, gave It up as a bad Job, and left it

Dr. Joseph Grinnell (1900) writes:

Occasionally a few spruce needles were also foond. The gizzards of the birds obtained, Invariably contained a quantity of small polished pieces of clear quartz, this probably being the hardest substance for the purpose obtainable by the birds. A bare place on a sand-bar in the river, kept clear of snow by the wind, was wont to be frequently visited by the ptarmigan and I have seen them scratching over tile gravel In such places, even In the coldest mid-winter weather.

In addition to many of the above items, E. A. Preble (1908) lists mushrooms, tops and seeds of grasses, and leaves, seeds, and berries of various other plants.

Behavior: Ptarmigan rise from the ground with a loud whir of wings and harsh cackling notes. When scattered about feeding among the willows they do not all rise at once but jump up singly or a few at a time, and there are generally a few laggards. When well under way their flight is strong, swift, direct, and often prolonged for a long distance. They alight readily on trees and bushes, where they are skillful at balancing. In winter it is difficult to see them on the snow-laden branches, where they look much like balls of snow. On bright, sunny days their shadows show up plainly on the snow-covered ground; but on dull, hazy days they are very hard to see, unless the black bill or eye is in motion. The black tail is entirely concealed, except in flight, when it serves as a very good direction mark for other members of the flock to follow. They seem to understand the value of protective coloration, for, if the ground remains bare after the white plumage is assumed, they are very shy; but, after the snow comes, they become very tame. Grace A. Hill (1922) calls attention to the fact that, while the ptarmigan are molting into the white winter plumage, they frequent the open tundra at the time the cottongrass (Eriophorumi polystaeAion~) is bearing its white cottony plumes, which aid the birds in their protective coloration.

Ptarmigan are evidently monogamous and make quite devoted couples. Dixon (1927) tells of a male, perched on the top of a spruce, standing guard over his mate while she was feeding. Mr. Dixon writes:

The bird gave a couple of warning calls as I approached the tree, and then it dawned on me that he was probably standing guard while his nesting mate fed. So I hunted around; and sure enough, I found the female ptarmigaii feeding in some dwarf willows about twenty feet from where I stood. As soon as I started after the female the male ptarnilgan flew down from the tree top and ran off ahead of me, trying in various ways to decoy me away from his mate.

I bad been told of an instance where a cock willow ptarmigan had attacked and routed a large grizzly bear that happened to stumble upon his nest. But even after seeing the ptarmigan drive off the gulls I did not fully appreciate the furiousness of the attack until June 23, when I came across an old hen ptarmnigan with her brood of small young which were just able to fly. I rushed after the young, trying to catch one. Just as I ~vas about to grab a chick, a willow bush in front of me exploded and the cock ptarmigan flew directly into my face, knocking my glasses to one side as he slapped my face with his beating wings. He then dropped to the ground, hut instead of retreating flew directly into my face ugain; but this time I was ready for him and caught him wilh my bare hand when he became mixed up with my mosquito head net. The bird Ihen tried to bite and to flap his way to freedom. As I started off with the cock under my arm the hen rtarmigan left her young and came rushing at me and then crawled feebly about at my feet as though in mortal agony. When I started away she rushed frantically about flapping my heels with her wings at every step. Every time she rushed at me she hissed. When the male found he could not escape he uttered a few creaking notes and the hen left me at once and went back to her chicks.

Voice: Dixon (1927) says that the warning cackle of the male “sounds like running a nail over a stiff comb.” He mentions three notes of the female: A harsh ke-ouk, ke-ouk is a warning of danger; a soft purring keer-er-eerk is a hush-a-bye, when hovering young; and a clucking cuck-cuck is a note used to call the chicks to her.

The downy chicks give a soft ckeep-ckeep-cheep when in distress or when separated.

Enemies: Gulls, jaegers, hawks, owls, foxes, wolverenes, and other predatory birds and animals levy heavy toll on the ptarmigan and their eggs and young. Ptarmigan are so plentiful that they furnish the principal food supply for many of these creatures, as well as for the human natives. Dixon (1927) writes:

After the young ptarmnigaa are out of the shell they are menaced by Blackbilled Magpies as well as by the foxes. Thus on June 24 a family of four young and two adult magpies was found systematically working the willows In the Savage River bottom for ptarmigan chicks. When these magpies located a pair of adult ptarmigan they would retire stealthily and hide in the willows near by, until the ptarmigan chicks began to run about. Then the magpies swooped down and grabbed the chicks before they could hide, and then carried them off and ate them. A cock ptarmigan that I watched put one magpie to flight, hut where there were sIx and In another case nine magpies working together against two adult ptarmlgan the odds were overwhelming. As a result of this persecution by the magpies we found that by July 10 many families of young ptarmlgan had been reduced to only one or two Individuals. Gyrfalcons also levy continuous toll on ptarmlgan; and since these large falcons are relatively numerous in the Mount McKinley district, the aggregate number of ptarmlgan killed by them Is considerable. It is thus easy to see why the hen ptarmlgan lays from 6 to 12 eggs. If only one or two eggs were laid each season the species would soon become extinct.

Game: Were its haunts not so far removed from the centers of civilization, the willow ptarmigan would be a popular game bird. Our experience with the Aleutian ptarmigan taught us that these birds possess excellent game qualities. Except during the breeding season, when they are very tame, they are wild and wary enough to give good sport. Their flight is strong and swift and sufficiently prolonged to give one all the exercise he wants. Their thick winter plumage is somewhat shot resisting, so that they have to be hard hit, with a close-shooting gun; it often requires considerable chasing to get within effective range. Edwyn Sandys (1904) gives a thrilling account of a winter ptarinigan hunt, with its hardships and dangers. The flesh of the old birds in winter is apt to be dark and dry and to have a bitter flavor, as a result of a steady diet of willow buds, but that of young birds in fall, fed on fresh foliage, berries, and insects, is lighter colored and very good to eat.

Dr. E. W. Nelson (1887) writes:

Among the Alaskan natives, both Eskimo and Indian, especially those in the northern two-thirds of the Territory, this bird is one of the most important sources of food supply, and through the entire wint~r It is snared and shot In great abundance, and many times it Is the only defense they possess against the ever-recurring periods of scarcity and famine.

The Eskimo of the Kaviak Peninsula have a curious way of taking advantage of the peculiarities of this bird in their migrating season. Taking a long and medium fine-meshed fishing-net they spread it by fastening cross-pieces to it at certain distances; then taking their places just at sunset in early November or the last of October, on a low open valley or “swale,” extending north and south, they stretch the net across the middle of this highway, with a man and sometimes two at each cross-piece, while the women and children conceal themselves behind the neighboring clumps of bushes. As twilight advances the net is raised and held upright. Ere long the flocks of Ptarmigan are seen approaching skimming along close to the snow-covered earth in the dim twlligh~ anil a moment later, as the first birds come in contact with the obstacle, the men press the net down upon the snow sometimes securing 50 to 60 birds. While the men throw themselves upon the net and hold it down, the women and children rush forward and kill the birds by wringing their necks or by biting their heads. On some evenings several flocks are thus intercepted, and the party of natives return to their houses heavily laden with spoils. In winter the birds are snared in their haunts by placing fine nooses attached to low bushes close to the ground. Sometimes small brush fences are built with snares at the passage-ways purposely left open. In spring, as the snow begins to leave the mossy knolls here arid there, the natives shoot a male bird and stuff it roughly with straw, and, mounting it on a small stake, place this effigy upon one of the bare knolls in a conspicuous position; then they surround It with a fine sinew net held in place by slender stakes. The hunter then conceals himself close by and Imitates the challenge cry of the male. All around can be heard the loud cries of the pugnacious birds, and attracted by the decoy notes of the native some of them are almost certain to bestow their attention upon the decoy; they approach swiftly, and either fly directly at their supposed rival or alight and run at him in blind rage. In either case their jealousy is fatal, as they are at once hopelessly entangled in the net of the hunter, who disposes of them and repeats the maneuver indefinitely, generally returning home well laden.

Ptarmigan are subject to great fluctuation in numbers from year to year, and during periods of scarcity they may be nearly or quite absent from regions where they were once abundant. Alfred M. Bailey (1926) says:

In 1919, ptarmigan were very scarce throughout the territory; in December, en a trig to within a short distance of the source of the Copper River, we snw but one bird. In 1920 the birds began to return, and In 1921 they were reported abundant at all points where they usually occur. I am unable to explain the cause of this scarcity at intervals, for, so far as I know, no disease has been reported among them.

Mr. Warburton, who spent the summer of 1929 about the mouth of the Yukon, writes to me:

The scarcity of these birds was a great disappointment. I had expected to find them as plentiful as I had seen theta about Nome and Teller, Alaska, In the summers of 1924 and 1925. At that time on the Seward Peninsula they were most plentiful, together with many rock ptarmigan. This year the exact reverse was true, at least about the Yukon mouth.

Fall: Turner (1886) says that the ptarmigan migrate to the interior late in the fall. He writes:

When the snow has pretty well covered the ground in late November the Plarmigans assemble In Immense flocks, often numbering thousands. I was once out on the higher grounds just south of the Crooked Canal. I ascended a slight hill and came, unexpectedly, on one of these large flocks that covered acres of ground. I was among them before either was aware of it. They flew, and made both the air and earth tremble. There must have been over five thousand birds in this one flock. They flew beyond a neighboring hill-range. Approaching night and a heavy snow falling prevented me from following them.

Winter: When the snow covers the tundra so deeply that no food or shelter can be found there, the ptarmigan are forced to migrate into the interior valleys, river bottoms, and creek beds, where they can find shelter among the willows, alders, or spruces and can feed on the bugs and twigs of these trees or such berries and fruit as still remain above the snow. They often congregate at that season, and, particularly when migrating, in enormous numbers. Mr. Brandt says that, where the railroad crosses the Continental Divide in Alaska, he saw large flocks arising “like snow drifts in motion, alongside the snorting engines, and whirling away over the great white hills.”

In winter the ptarmigan’s feet are thickly covered with long, hairlike feathers, resembling the foot of a hare, which serve as snowshoes and enable the bird to walk on soft snow. Sandys (1904) Writes:

And Nature, as If realizing the perils of the ptarmlgan asleep, has taught It to plunge beneath the cold drifts to escape the cold, and to fly at, not walk to, the chosen drift, so that there will be no telltale trail for some keen nose to follow to the sleeping-place. And this the bird Invariably does, going at speed and butting Its way into the snow, leaving never a print to betray its retreat, from which it flies in the morning. The game of life and death is interestingly played up North: where the weak white snow-shoers are ever biding from the strong white snow-slicers forever searching over a field of baffling ice-bound white.

Doctor Nelson (1887) describes another method of roosting:

On November 25, 1877, they were numerous, in large and small flocks, along the bushy gullies and bill slopes on the shore of Norton Sound, but were shy. In many places where they had stopped the night before, their sleeping-place was well marked. In each instance they had occupied a small clear spot in the midst of a dense thicket, and in no case had the birds approached on foot, but had flown in over the top and plumped down into the soft snow, where they had remained during the night, each bird thus making a mold of itself In the snow. In some instances there were fifteen to twenty of these molds in the snow in an area of a few feet. In leaving their stopping-place the birds arose and flew directly from their “forms,” as was shown by the marks of the wings on each side as they touched the snow in rising, so there were no tell-tale tracks to or from these places; the open places were undoubtedly chosen to allow the birds an unobstructed escape in case they were surprised by prowling foxes, which hunt these thickets for food.

DISTRIBUTION
Range: The range of the willow ptarmigan as a species is circumpolar, extending in the Old World from Scandinavia, Russia, and Siberia south to Turkestan, and in North America from Greenland, Newfoundland, the Arctic Archipelago, and Alaska south through Canada. The species is casual in winter in the Northern United States.

Breeding Range: The North American breeding range extends north to Alaska (Gape Lisburne, Wainwright, Point Barrow, Smith Bay, Delta of Colville River, Camden Bay, Humphrey Point, and Demarcation Pointy; Franklin (Bay of Mercy, Port Kennedy, Felix Harbor, and Igloolik) ; northern Quebec (Fort Chimo) ; and Labrador (Okkak). East to Labrador (Hamilton Inlet) and Newfoundland (Raleigh and St. Johns). South to Newfoundland (St. Johns and south coast) ; west-central Quebec (Carey Island) ; northern Ontario (40 miles south of Cape Henrietta Maria); northern Manitoba (50 miles north of York Factory and Fort Churchill); southern Mackenzie (Artillery Lake and Fort Resolution) ; central British Columbia (Moose Pass, Icha Mountains, and Ninemile Mountain); and southeastern Alaska (San Juan Island). West to Alaska (San Juan Island, Kruzof Island, Glacier Bay, Nushagak, Nelson Island, Igiak Bay, Askinuk Range, Pastolik, St. Michael, Nome, Mint River, Cape Blossom, and Cape Liaburne).

Winter range: Although willow ptarmigan perform a very definite migration, individual birds are frequently found in winter almost at the northern limits of the breeding range. At this season they have been recorded north to Alaska (Nunivak Island, Nulato, Kutuk River, and Miller Creek) ; Mackenzie (mouth of the Dease River, Fort Rae, and Fort Reliance); northern Manitoba (Fort du Brochet); Franklin (Igloolik); Quebec (Great Whale River, and Piashti); and Newfoundland. Normally they are found south to southern Quebec (Lake St. John); central Ontario (east of Cochrane and Martin Falls); central Manitoba (Norway House and Grand Rapids) ; Saskatchewan (Cumberland House and Fort Carleton) and southwestern Mackenzie (Fort Simpson).

Migration: Though the bulk of the willow ptarmigan move south each fall or winter, and return north in spring, the movement has no regularity and is directly correlated with the food supply and, in consequence, with the fall of snow.

First fall arrivals are: Quebec, Lake Mistassini, October 25, and Godhout, November 2; Ontario, Martin Falls, about October 20; Manitoba, Lac du Brochet, November 4, Grand Rapids, November 12, and Winnipeg. January 12; Mackenzie, Fort Rae, October 1; and Alberta, Fort Chippewyan, October 11.

In spring, late departures for the north are: Quebec, Lake Mistassini, about May 1; Ontario, Coclirane, March 20; Manitoba, Winnipeg, March 21; and southwestern Mackenzie, Fort Simpson, March 12. They have been observed to arrive at Fort Resolution, Mackenzie, on June 28, and at Demarcation Point., Alaska, as early as April 6.

Casual records: The willow ptarmigan has been reported a few times in the Northern United States. Among these records are: Maine, one at Kenduskeag, April 23, 1892; New York, one at Watson, May 22, 1876; Wisconsin, two taken near Racine in December, 1846; Minnesota, one taken at Sandy Island, Lake of the Woods, April 20, 1914; and Montana, three taken in Glacier National Park in the winter of 1913: 14. Prof. IV. B. Barrows believed that they occasionally occurred on Keweenaw Point, Mich.

The range as above described is for the entire species, which in the 1931 edition of the American Ornithologists’ Union Check List is subdivided into five subspecies. According to this authority the subspecies albus is found in North America from the eastern Aleutian Islands, northern Mackenzie, northern Banks Island, and central Greenland south to central British Columbia, central Alberta, central Ontario, and southern Quebec. The Ungava ptarmigan (L. 7. ungavus) occurs from northern Quebec west probably to the eastern shore of Hudson Bay. Allen’s ptarrnigan (L. 7. alleni) is confined to Newfoundland. The Alaska ptarmigan (L. 7. alascensis) is found on the Alaskan mainland (except the southeastern coast), northern Yukon, and eastward for a distance not yet determined. Alexander’s ptarmigan (L. 7. alexandrac) occurs on Baranof Island, Alaska, and adjacent islands, west to the Shumagin Islands, and south to Porcher Island. This race also may occupy a narrow strip on the mainland from Glacier Bay to central British Columbia.

Egg dates: Northern Alaska: 08 records, May 25 to July 10; 34 records, June 6 to 25. Arctic Canada: 37 records, June 2 to July 7; 19 records, June 10 to 21. Labrador Peninsula: 18 records, June 1 to 30; 9 records, June 6 to 23. Newfoundland (alieni) : 11 records, May 12 to June 30; 6 records, June 8 to 12. Southern Alaska and British Columbia (alexartdrae): 3 records, May 28, June 25 and 26.

LAGOPUS LAGOPUS ALLENI (Stejneger)
ALLEN’S PTARMIGAN [Current A.O.U. = Willow Ptarmigan]

HABITS

The willow ptarmigan of Newfoundland was originally described by Dr. Leonhard Stejneger (1884) as similar to the common willow ptarmigan, “but distinguished by having the shafts of both primaries and secondaries black, and by having the wing-feathers, even some of the coverts, marked and mottled with blackish.” He examined only 14 specimens in all, all of which presented the above characters. In a large series (I have 75 in my own collection and have examined a great many in other collections) these characters appear to be none too constant in Newfoundland birds and to crop out occasionally in willow ptarmigan elsewhere, with a great range of individual variation. In this connection it is interesting to note what Harry S. Swarth (1924) has to say about the willow ptarmigan of southern Alaska:

It is of interest to note in aleJefl(lrOC the frequent presence of black shafts on the io’imaries, sometimes on secondaries and greater coverts. This character has been considered an important foature of the Newfoundland subspecies (C. 1. alIen I), hut obviously it can not be used as a feature characteristic of that race alone. In an immature female from Prince of Wales Island, which has acquired the winter flight feathers, not only are primaries and secondaries with distinct black shafts, hut there are large, tear-shaped spots of black near the tips of all the primaries and most of the secondaries. Furthermore, the primaries have a black “freckling” over much of their surface, and the greater coverts are also marked with black, though to a lesser degree.

Dr. Hart Merriam (1885) has called attention to the great variation in the extent of black in a large series of wings sent to him by Napoleon A. Comeau from Godbout, Quebec. Perhaps the range of alleni should be extended to include eastern Quebec.

Allen’s ptarmigan are very abundant and widely distributed in Newfoundland, on the upland tundra in the central and northern parts of the island and on the mountains. J. R. Whitaker tells me that from about the middle of April to September 20 they live on the high tundras and rocky ridges; he has never seen a bird on the lower ground during summer. F. S. Hersey, who went to Newfoundland in September and October, 1913, to collect ptarmigan for me, found these birds more abundant than he ever found willow ptarmigan anywhere in northern Alaska. He found them on the hills a few miles back from Cape Ray, and some were seen in the Lewis Hills near the west coast; but they were most numerous about Gaiftopsail in the interior of the island. This is a rough, barren, open region, tending to run into low, rocky ridges, separated by lower marshy areas, carpeted with thick mosses, often knee-deep, and dotted with small ponds. There is a little cover here, although a few sheltered spots may have low shrubs and nearly prostrate blueberry bushes, and in a few places there are small areas of exceedingly tough, thick, dwarf spruces.

Courtship: Referring to the courtship of this ptarmigan, Mr. Whitaker says in his notes:

The cock bird nearly always mounts a rock to utter his challenge; having gained this vantage point he looks all around, then with bead erect and breast expanded bleats out his notes; they will start a little before sunrise and continue for a short time and are seldom heard during the day. After the young are hatched crowing appears to cease entirely. The crow of this bird is very like the British red grouse; it Is quite easy to imitate, but to me, very difficult to write. The first part sounds like the low trembling bleat of a nanny goat, or, If one can imagine it, a very coarse drum of a snipe; there are usually five quickly uttered tremulous notes run off er-er-er-er, followed by three or four Dobeck-gobeok-gobeck; these latter may be copied perfectly by partly closing your throat and uttering gobeck harshly through the nose.

Nesting: Whitaker says that nesting begins in May and the young are usually full grown by the end of August. The nest Is usually placed at the base of some bowider and Is well hidden; the hen bird sits very close indeed. Some years ago a nest was found close to a watershoot by the railway; several trains a week stopped here for water; the train hands used to take passengers to see the bird sitting, which did not upset her in the least.

A set of eight fresh eggs, collected for me near Gaiftopsail on June 16, 1912, was taken from a hollow in the tundra moss, under a little bush, but in plain sight, between two hummocks. Mr. Hersey found a nest from which the young had hatched, which he describes in his notes, as follows:

The site was on the side of a ridge some 20 feet above the marshy ground at its base. A few bushes grew at this point, and under the overhanging branch of one of these the nest had been made. Even at this season the hollow that had been the nest was well defined. There was no indication of lining beyond the accumulation of fine dry leaves, bits of sticks, moss, and other vegetation that made up the general ground cover on the whole ridge. The eggshells from which the young had hatched were still in the nest.

Eggs: The eggs of Allen’s ptarmigan are indistinguishable from those of its mainland relative. The measurements of 54 eggs average 42.5 by 33 millimeters; the eggs showing the four extremes measure 45.6 by 32.3, 44.1 by 37.6, 40.1 by 30.2, and 43.4 by 28.7 millimeters.

Plumages: The molts and plumages of this race are similar to those of the willow ptarmigan. It is interesting to note, however, that many of the wings sent by Mr. Comeau to Doctor Merriam (1885) taken from birds killed in November, were “deeply tinged with a delicate and very beautiful shade of rose-pink which is more pronounced than in a freshly killed Roseate Tern.” Mr. Whitaker has also noted this, as he says in his notes: “They are in full white winter plumage by the end of October; I say white, but there is a beautiful faint pink flush; I first noticed this when having shot several one winter day and laid them on a block of pale blue ice, this showed up the pink to advantage; the pink quickly fades after death.” Food: In his notes on food, Whitaker writes:

As the snow melts off the high ground many berries that have lain snug all winter provide plenty of food; after these are shriveled up by the sun and before the new crop matures, the birds feed on the tips of a low-growing plant, which looks like a very dwarf heather. In August the blueberries and partridgeberries. (Mitchella repens) ripen and on these they feed almost exclusively. When the snow begins to pile up on the highlands during October the bulk of the birds move down to lower levels and begin feeding principally on buds of scrub birch; they also eat buds of blueberry and pussy willow.

The birds shot by Mr. Hersey had their crops filled with blueberries and their leaves, the small green berries from a dwarf evergreen resembling a cedar, or with small white seeds.

Behavior: Mr. Whitaker says in his notes:

These ptarmigan are very tame in the wilder districts, where they are not disturbed or shot, and seem loath to take wing; they will run in front of you stopping every few yards and spread their tails with a quick jerky motion; they often utter a suppressed little grunting note, possibly a protest against your intrusion In their midst.

Mr. Hersey’s notes contain the following observations:

The birds were usually found in parties of 3 or 4 to as many as 11 and 13 and, as many were in a plumage that indicated they were birds of the year, it was assumed that these flocks usually represented a pair and their grown-up young. They were very erratic in the matter of taking flight, on some occasions not flying until nearly stepped on and again taking wing before a person was within 100 yards of then,. As a general thing they were wildest on sharp frosty days or when rather windy and would lie closest when the day was warm and calm. I do not believe the ptarmigan range over any large area at this season. Birds were seen day after day on the same ridges and were believed to be the same flocks. On bright clear days a flock often spends several hours crouched on the ground In the sunlight, particularly In rocky places, where one or two males will take up a position on the rocks. These sentinels perform their duty poorly, as it is usually rather easy to approach and shoot such birds and then secure one or more from the flock with the remaining barrel as the birds take wing. At times the birds on the ground will not flush, even after one or more has been shot. I remember one very warm day when I came upon three birds crouched on the ground a few feet apart. I was unable to flush any of them and was obliged to back away and shoot them on the ground one at a time, the remaining birds paying no attention to the sound of the discharge. When flushed they are frequently silent, but some birds give voice to a harsh cack-cock-oack-cack-caek. On foggy mornings birds, presumably males, may be heard “crowing,” at least the notes sound very similar to the spring call of the male. They may be a signal or flock call.

Game: Allen’s ptarmigan, known also as “willow grouse” or “partridge,” is by far the most important game bird in Newfoundland. It is jealously guarded during the close season, only to be shot in enormous numbers during the open season. Many parties of gunners go out every fall from St. Johns, as well as visiting sportsmen from Canada and the United States, camp along the line of the railway, and kill hundreds of the birds. Fortunately for the birds, there are immense tracts of uninhabited country, inaccessible by branch lines or by roads, where they are not disturbed, so that they do not. seem to be in any immediate danger of extermination.

Winter: Mr. Whitaker says that “quite a number perish during the winter; they make a shallow scratching in the snow in which to roost and are frequently buried by drifts and are imprisoned; I have often found them dead in the spring when the snows are melting, and once saw seven within a few feet of each other that had met this fate.”

LAGOPUS LAGOPUS ALEXANDRAE (Grinnell)
ALEXANDER’S PTARMIGAN [Current A.O.U. = Willow Ptarmigan]

HABITS

The willow ptarmigan of the humid coast region of southern Alaska and nortbern British Columbia is darker and more richly colored than birds from northern Alaska and Labrador, with a smaller and narrower bill, the general color above being deep, rich chestnut thickly vermiculated with black. The most typical birds of this race are found on the islands along the coast of southern Alaska; the type came from Baranof Island. It was described by Dr. Joseph Grinnell (1909) and was named in honor of Miss Annie M. Alexander. Alfred M. Bailey (1927) first saw them on Willoughby Island in Glacier Bay, of which he says:

This island is about five miles in length, with scant vegetation, other than alder and stunted spruce. The knobs are devoid of l)lSflt life, and it is only on the terraces that soil can hold. The southern and eastern slopes are rather densely clothed with alders, however, and form ideal cover for Ptarmigan. Seven other birds were seen by Young, one of which was still in the winter white. The pair collected were breeding birds.

He suggests a good reason why they should prefer to live on the islands:

The lack of predatory animals was very noticeable, for with the exception of two Eagles, none was noted, and no signs of predatory animals were seen on the outer Beardslee Island, although those nearer shore must have some carnivorous mammals. We saw a fox in June and August on the mainland shore, and wolves are abundant. It is rather apparent then that the birds are free from molestation, in direct contrast to their life upon the mountain slopes of the mainland. Ravaged by fur-bearers, it is possible the Ptarmlgan first used the Islands for protection, and having found both food and comparative safety, have continued to live under such conditions. On the other hand, the islands might be considered as being alpine in nature, with timber-line conditions, as the spruce are small and willow and ‘alder predominate, with the characteristic profusion of small growths. The soil is scant, the glacial sands and moraine d~brls being exposed, while the windward shores of the outer Beardslee are precipitous; the glacial winds sweep down channel, Icebergs line the shores, and taking all into consideration, the region is probably the coldest of southeastern Alaska.

Harry S. Swarth (1924) noted that the birds he collected in the Skeena River region of northern British Columbia “are intermediate in color between lago pus of the interior and alex~aindrae from the islands; the average is nearer alexandrae.” He says of their haunts there:

Ptarmigan are said to occur occasionally in the lowlands of the Hazelton region In midwinter, but during most of the year they are restricted to the Alpine-Arctic mountain tops. We found them in limited numbers on the timberless summit of Nine-mile Mountain. There are miles of open country on the two converging ridges that form the top of this mountain, barren of trees save for occasional thickets of dwarfed or prostrate Alpine conifers, and here, at long Intervals, we encountered ptarmigan.

Nesting: We know very little about the nesting habits of Alexander’s Ptarmigan. According to Doctor Grinnell (1909), Joseph Dixon “records that at Coppermine Cove, Glacier Bay, July 10 to 20, the feathers and bones of a ptarmigan were found near a nest of broken eggs on the summit of the mountain, 2,100 feet. The nest was under a stunted hemlock. All the feathers were white, so the ptarmigan must have laid early.”

There is a set of 11 eggs in Col. John E. Thayer’s collection, taken by John Koren on Kodiak Island, Alaska, on June 25, 1911. The eggs were fresh and “were placed on a bed of moss in a two-foot groove in an elevated part of the tundra “; both parents were present. The eggs are not distinguishable from those of the willow ptarmigan. The measurements of 20 eggs average 42.8 by 30.9 millimeters; the eggs showing the four extremes measure 45.9 by 31.5, 43.6 by 32, 41.5 by 30.8, and 42 by 30.2 millimeters.

Young: Swarth (1924) writes:

In all, ten broods of willow and rock ptarnaigan were encountered (the species were not always to be differentiated) and about five or six single birds in addition. The broods ranged from three to twelve in number; the aggregate of young birds seen was about fifty. The chicks grew rapidly. Some seen on July 25, and a day or two later, were down-covered and unable to fly. At that time they were accompanied by the female parent only, and the male birds were flushed separately. By August 10 the young ptarmlgan were the size of quail and larger, and were strong on the wing. The old males were then associated with the families. In some of the larger broods seen the difference in size among the young was so marked as to suggest the junction of two families. It might happen that upon the death of a hen her offspring would seek the companionship of another family.

Food: These ptarmigan probably feed on as varied a diet as other ptarmigan, but the following note by Alfred M. Bailey (1927) is all that I can find in print on the subject:

A flock was flushed from a bed of wild strawberries, at an altitude of scarcely thIrty feet, when I had expected to find them above timber line. There were several pairs of adults, as well as many young, and a good series was taken. An examination of crop contents proved the birds had been feeding entirely on strawberries and pea-vine, no alder or willow buds being found. After finding ptarmlgan In such a low altitude, I searched all the points along the east mainland shore, and did not fall once to find them, where there were berries. It was noticeable they preferred the points where they could feed close to a fringe of alder.

Behavior: Mr. Bailey (1927) says also:

I returned to Glacier Bay again from October 10: 14, and observed the Ptarmigan under still different conditions. A stop was first made on the outer Beardslee, upon which the young bird had been taken. I was Interested to see it the Ptarmigan had left the island, as it was now drab and dry looking, and the birds were assuming their winter’s white. I found them very abundant, and over forty were flushed from the dense alders, and a few taken. They were extremely wild, as they should have been, for their changing plumage: entirely white below with many white feathers in head and neck, made them extremely conspicuous among the leafless alders. I had little chance to observe them, due to their wildness. Stops were made at two other points, not visited in August. and ptarmlgan were noted in both places. At Sandy Cove, October 14, I collected another series from the point they were first found in August. These were as conspicuous as the others, but when the flock was broken up, the birds did not flush so readily, and I had a chance to study them at close range. They ran over the ground the same as in August, evidently believing themselves inconspicuous, and when closely pressed, crouched quail-like and depended upon their “protective coloration.” When the birds became scattered several climbed into alders, about a foot from the ground, and sat hunched in some convenient crotch, where they were more evident than ever. I was surprised to find strawberries still abundant here, hidden under the mosses in shady places, and each of the specimens taken had Its crop full.

Doctor Grinnell (1909) quotes from Chase Littlejoha’s notebook as follows:

While searching for eggs of the glaucous-winged gull on one of the small islands on the east side of Glacier flay on July 14, I suddenly came upon a flock of ptarmigan in a little opening among some spruce, hemlock, and alders, which covered the ground In dense masses in spots; the remainder of the area supported a thick growth of grass interspersed with patches of moss and low-growing flowering plants. There were about eighteen birds all told, young and old, and as near as I could determine there were four or five old birds present. They would not fly after they were first flushed, but kept dodging about on the ground, sheltered by the thick cover; several times I saw them, but so near that a shot would have ruined them as specimens.

LAGOPUS LAGOPUS UNGAVUS (Riley)
UNGAVA PTARMIGAN [Current A.O.U. = Willow Ptarmigan]

HABITS

The ungawus race of the willow ptarmigan was described by 3. H. Riley (1911) as “like Lago pus lago pus aUnts, but with a heavier bill,”. based on a series of 20 birds collected at Fort Chimo, Ungava, in the northwestern portion of the Labrador Peninsula. He gives as its probable range “from Ungava and probably the eastern shore of Hudson Bay south.” Birds that I have seen from the eastern and southern parts of the Labrador Peninsula do not seem to have appreciably large bills. If this form is worthy of recognition at all, which I very much doubt, it will probably prove to be a northern race and perbaps identical with alasoertsis Swarth, about which I have expressed my views elsewhere. But, pending further investigation, we may as well consider all the willow ptarmigan of the Labrador Peninsula as referable to this form.

In this connection it might be well to consider what Lucien M. Turner has to say, in his unpublished notes, concerning the great degree of individual variation that he noted in the hundreds of Ungava ptarmigan that he handled. He found a great “individual variation in the size and shape of the head and beak.” He collected a large series of skulls and found them to vary from “1.94 to 2.30 inches in length from occiput to tip of bill.” He says that “the large beak is especially noticeable in some of the birds, more especially in the males,” but that “some only of the males alone have a noticeably large beak.” He concludes by saying that this character should not, in his opinion, warrant the separation of this form.

I found and collected a few specimens of willow ptarmigan near Hopedale and at Ukjuktok Bay, on the northeast coast of Labrador, in 1912. They were. breeding there, and I was told that they also breed on some of the islands. They are said to migrate farther south in October and return in April and May. I have a set of eggs taken near Okkak in July. Dr. Harrison F. Lewis (1928) said that this species t’ bred in the summer of 1928 on the large island at the Bluff Harbor where they had nested every year since 1925. It seems probable that two pairs of willow ptarmigan nested on this island in 1928.”

0. J. Murie has sent me the following interesting notes:

Along the Nastopoka River, on the shore of Hudson Bay, lies a region that is apparently an ideal nesting ground for willow ptnrmigan. On the morning of May 11, my Eskimo guide leJi me inland over the granite hills about parallel with the river. The immediate coast in this vicinity is bare and treeless. Ten or 15 miles inlan(l we found scattered patches of small stunted spruces, covering some of the lower hills. Here we found a ptarinigan paradise. The birds were everywhere, apparently all paired, the males thrilled with the energy of the mating season. It was invariably the male that flew out first when we approached small clumps of willows. He went off lIke a rocket, with a clattering racket of harsh notes and whir of wings. He would generally take a commanding position on a rocky point (Jr tnT) of a hill and, strutting pompously with head high and tail raised and spread, let out his clattering crow.

At this time of the year I was struck by the perfect blending of the plumage with the surroundings, especially in the case of the female. Her plumage at this time is speckled irregularly with various shades of brown and white, in varying stages of molt, which was difficult to distinguish on a ground of mosses, lichens, rocks, and willows, sprinkled with remnants of snow. The females showed a tendency to keep well under cover. The males were not particularly difficult to see, and I spied them readily at a considerable distance en occasion, but sometimes I thought I could detect a principle of protective coloring. The body was pure white, with a brown head and neck, apparently the usual plumage in the season of courtship, judged from this one spring. Usually the white plumage flashed out prominently, but the brown neck was lust’ on the background of rock and moss. Although the eye xvas attracted at once, there was a tendency to pass it over without recognizing the headless body as a bird. Similarly, on a snowbank, the brown neck was seemingly detached, and not a part of a bird.

Nesting: Turner says in his notes:

In the vicinity of Fort Chimo, nesting of this species begins during the latter part of May. The nest is usually placed in a dry spot among the swamps or on the hillsides where straggling bushes grow. The nest is merely a depression in the mosses and contains a few blades and stalks of grass together with a few feathers from the parent bird, which is now In the height of the molt from the winter to the pre-aestival p]umage. The first eggs obtained were two on June 1, 1884.

Young: Ref erring to the period of incubation, Turner writes:

It is a rather difficult matter to determine, as the female Is compelled, during a stress of severe weather, to sit upon her eggs to prevent them being lost by cold or rain. It Is not unusual for severe snow and sleet accompanied by cold rains and even a severe freeze to occur during the early half of June at Fort Chimo. Some of the most dismal days of the year occur In early June. The parent bird, during such weather, may be two or three days on the nest after the first young bird has appeared and thus prevented from giving such attention to the young as these tender creatures require. It Is not rare to find a nest containing two or three eggs and near by to find one or more young which have perished while the mother has perhaps wandered off with three or four young which were able to follow her.

He says that “the Indians consider the downy young of the ptarmigan a special delicacy. Even taken from the shell the bird serves in lieu of an oyster.” lie frequently saw them eating the embryos taken from eggs that they were blowing. He says that they make special excursions to collect the small chicks for food. One party that he saw returning from such a hunt had more than 250 of these helpless young.

At Ukjuktok Bay, on August 3, 1912, I surprised a family party of willow ptarmigan in a boggy, grassy hollow. The young, which were about half grown, rose with a startling rush of wings and went whirring off like a flock of quail. The old birds did not flush. The female feigned lameness, in spite of the fact that the young had all flown; I could not make her fly, and she finally walked away. The mile walked boldly out into the open marsh, looking at me, too close to shoot, then ran behind some spruces and flew away to join the young. I followed them up and flushed the male first; then three of the young rose singly. I could not find any more of the young, but a little later I found both old birds in the exact spot where I had first seen them.

Voice: O. S. Murie has sent me the following good description of the ptarmigan’s notes:

The call of the Ptarmlgan Is very striking. It consists of a rattling lcrrrr-r-ruk,-uk-uk-itk-s&k, followed by a more deliberate, low-toned, throaty puk-que’-o, puk-qae’-o, puk-que’ -0. I thought It fitted well with the surrounding hills of rough granite and the scant growth of ragged, twisted spruces. Sometimes the female was heard responding with a peculiar whirring sound, a nasal ssyek, nyelc, somewhat similar to some notes of other members of the grouse family. By Imitating this note we frequently drew the male to us In a headlong flight. He would drop on a knoll near by and send out his startling call. The Eskimos take advantage of this trait and decoy the birds to he shot

Winter: Ptarmigan are great wanderers in winter, but very erratic in their movements, appearing in enormous numbers dur ing some seasons on the eastern and southern coasts of the Labrador Peninsula, and being nearly or quite absent during other seasons. Dr. Harrison F. Lewis writes to me:

This species often appears on the coast in great numbers in autumn, but the numbers so appearing vary from year to year. When these birds come out from the interior to the coast they seem to come as far out as possible, and often go in numbers to the very outermost islands which, of course, are often united to the mainland by ice at that time of year. Large numbers are taken by the residents for food. No matter how plentiful the ptarmlgan may be in the fall and the month of December, comparatively few are usually to be seen after the 1st of January, although a few may often be observed from time to time until spring.

But vast numbers remain in northern Ungava, at least during some winters, according to Turner. One day he saw a gyrfalcon flying across the Koksoak River, and writes:

We had seen hundreds of ptarmigans on the left bank among the thickets. The hawk plunged among these birds which began to rise as soon as the hawk was sighted. I am certain that not less than 1,800 ptarmigans rose before that hawk; and, as the latter did not reappear, we suspected there was at least one less ptarmigan. The air fairly trembled as these birds arose. In my notes I find the following for the locality of Fort Chimo and date of December 7, 1882. Hundreds of this species of ptarmigan have made their appearance In this vicinity during the past week where two weeks ago not a dozen birds of the kind could be found during a tramp of an entire day.

The traffic in ptarmigan feathers is very heavy and will give an idea of the enormous number of birds killed. The Indians save only the clean feathers from the breasts and backs of the bircfs, pack them in bags, and trade them to the Hudson’s Bay Co. They are then packed in barrels and shipped to England. Mr. Turner estimated that it required the feathers of 16 ptarmigan to make a pound and says that 31 barrels of feathers were shipped from Fort Chimo during the year ending June 1, 1883. As the average weight of feathers in a barrel was 51 pounds, he calculated that it accounted for 25,296 ptarmigan killed. And to this must be added a very large number of birds killed whose feathers were soiled or for some other reason were not saved. Fortunately the ptarmigan is a prolific breeder; otherwise, the wholesale slaughter of chicks and old birds would soon exterminate the species.

LAGOPUS LAGOPUS ALASCENSIS (Swarth)
ALASKA PTARMIGAN [Current A.O.U. = Willow Ptarmigan]

HABITS

Harry S. Swarth (1926) gave the name alascensie to this supposed subspecies, which he characterized as “slightly larger than albus. A large billed race; bill slightly smaller than in ungavus, much larger than in albus. In summer plumage, generally more reddish-colored than either ungavus or albu8, a difference that is most conspicuous in females in the barred breeding plumage.”

He gives as its range “the Alaskan mainland except on the southeastern coast, northern Yukon Territory (specimens from vicinity of Forty-mile), and eastward for an undetermined distance.”

At least three authors have attempted to subdivide the willow ptarmigans of North America, with decidedly confusing and somewhat unsatisfactory results. Austin H. Clark (1910) after studying 115 specimens, 20 from Newfoundland, 60 from Labrador, 3 from central arctic North America, 18 from the mainland of Alaska, 2 from Kodiak Island, and 12 from the Shumagin Islands, says:

All those from Labrador and central arctic America, with others from Point Barrow, Kotzebue Sound, Cape Lisbourne, Kowak River, Yukon River, and near St. Michaeis, belong to a well-differentiated race, with the beak very large, high and stout, the culmen strongly arched, and usually with a prominent ridge from the inferior corner of the maxilla to in front of the nostriL They are identical among themselves, it being impossible to tell from the examination of any one specimen whether It was taken in Alaska or in Labrador.

J. H. Riley (1911) evidently differed from him, for when he described and named the Ungava bird, from the Labrador Peninsula, he gave as the range of albus “from the west side of Hudson Bay, west through northern Alaska to eastern Siberia.” Thus two investigators, with practically the same material for study, have arrived at quite different conclusions.

Now Swarth (1926) comes along with a still different theory, based on a study of a large series of western birds in California collections, together with 3 from the west coast of Hudson Bay and 10 from Fort Chimo, Ungava. He writes:

Comparison of these birds with the series in this museum convinced me of the existence of the following recognizable subspecies of the willow ptarmigan on the North American mainland: (1) Lago~us Zagopus ungav,~s from the region east of Hudson Bay, as defined by Riley; (2) Laiyo pus iagopus albus from the west shore of Hudson Bay westward to the coast ranges of northern British Columbia, and for an undetermined distance northward; (3) an undescribed subspecies from the Alaskan mainland and extending for an undetermined distance eastward in the extreme north.

Swarth (1926) then goes on to describe the northern bird, alascensts, as quoted above and says further:

Conditions in these western races of willow ptarmigan parallel to some extent those found in the rock ptarmigan. In each species the northern Alaskan subspecies is an extremely ruddy-colored bird compared wiih the others, and in each the British Columbia subspecies seems to reach an extreme of grayness. In each species, too, the Labrador birds are much more grayish than are those from Alaska. Thus the Labrador willow ptarmigan (unga’vu8) and the British Columbia bird (albus) are much alike as regards color but differ in size of hill. The Labrador bird and the northern Alaskan bird (aiascen-sis) are both largebilled forms, but differ in coloration.

Perhaps, after studying the three papers referred to above, the reader may get a clear idea of the subject; but I must confess that I can not. In my opinion, all the willow ptarrnigan of the North American mainland (excluding alexandrac and perhaps allend) are of one subspecies, Lago pus lago pus albus. The differences pointed out above are 100 slight and too variable to be worthy of recognition in nomenclature.

About the Author

Sam Crowe

Sam is the founder of Birdzilla.com. He has been birding for over 30 years and has a world list of over 2000 species. He has served as treasurer of the Texas Ornithological Society, Sanctuary Chair of Dallas Audubon, Editor of the Cornell Lab of Ornithology's "All About Birds" web site and as a contributing editor for Birding Business magazine. Many of his photographs and videos can be found on the site.

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