All three subspecies of the Greater Prairie-Chicken have had population declines. The Heath Hen is extinct, the Attwater’s Prairie-Chicken is endangered, and the Greater Prairie-Chicken is declining over much of its range. Its inflatable, colored air sacs and its springtime courtship displays on communal booming grounds have made it an iconic symbol of the tallgrass prairie.
The large, meaty prairie-chicken is a tempting meal for many hawks, as well as foxes and coyotes. But habitat loss is likely more of a problem for the species, with agricultural conversion, intense grazing, and fragmentation of habitat through power lines, wind turbines, and other tall, vertical structures all playing a role in diminishing populations.
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Description of the Greater Prairie-Chicken
The Greater Prairie-Chicken is a grassland grouse, with mostly brownish plumage that is heavily barred on the underparts. It has long, erectable pinnae on the head.
Males have inflatable golden neck sacs and yellow eye combs. Length: 17 in. Wingspan: 28 in.
Females lack neck sacs and eye combs.
Seasonal change in appearance
Juveniles resemble adult females.
Greater Prairie-Chickens inhabit tallgrass prairie.
Greater Prairie-Chickens eat seeds, insects, acorns, and leaves.
Greater Prairie-Chickens forage on the ground, and occasionally in trees and shrubs.
Greater Prairie-Chickens are resident locally in remaining tallgrass prairie areas of the Midwest and eastern Great Plains. The population has declined dramatically in recent decades.
Bent Life History
Visit the Bent Life History for extensive additional information on the Greater Prairie-Chicken.
Male Greater Prairie-Chickens have a courtship display in which they inflate their air sacs, give a series of resonant hoots, and stamp their feet.
There are three recorded subspecies of Greater Prairie-Chicken, the extinct Heath Hen of the Atlantic Coast, the widespread subspecies of the central U.S., and the endangered Attwater’s Prairie-Chicken of coastal Texas.
The song consists of a series of moans or hoots. Cackles and clucking calls are also given.
The Greater Prairie-Chicken’s nest consists of a shallow depression lined with plant materials.
Number: Usually lay 10-12 eggs.
Color: Olive or buffy with darker markings.
Incubation and fledging:
The young hatch at about 23-25 days, and leave the nest shortly after hatching, though associating with the adults for some time.
Bent Life History of the Greater Prairie-Chicken
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 Greater Prairie-Chicken – the common name and sub-species reflect the nomenclature in use at the time the description was written.
TYMPANUCHUS CUPIDO AMERICANUS (Reichenbach)
The prairie chicken ranks first among the game birds of the prairies of our Middle West. It is to the prairie what the rutted grouse is to the wooded sections of the country. As intensive agriculture pushed to all sections of the range of the prairie chicken and as interest in hunting increased, this fine game bird at one time seemed in grave danger of following the course taken by the heath hen, to extinction as a game bird. In fact, it is gone from much of its former range, and its original numbers have been greatly reduced in practically the entire area of its distribution.
Because market hunting has been made a thing of the past since the beginning of the twentieth century and also because of the increasing restrictions on hunting by State departments, as well as various effective conservation programs, the prairie chicken is now holding its own and is increasing its numbers in many sections of its present range. Another hopeful sign is the fact that it has been extending its range to the northwest, and to-day the species is well represented on the prairies of Manitoba and is gradually spreading westward through Saskatchewan and Alberta, where formerly it did not exist.
The State Department of Conservation of Wisconsin has undertaken a comprehensive investigation of the prairie chicken to ascertain all the facts that affect its life, with the expectation that the department will be able to carry on a more effective program of conservation. Until the fundamental facts in the biology of our game birds are clearly known, conservation commissions will be handicapped in handling questions of game legislation and game management.
Prairie chickens, in common with other grouse, go through definite cycles of numbers. The problem of fluctuations in numbers of various species of wild life is not yet definitely solved, but work on it in relation to the ruffed grouse is being undertaken by many institutions and individuals in different parts of the country; hence there are excellent prospects of this work being brought to a successful conclusion.
The weather condition during the nesting season, especially during the height of the hatching period, is so important that it is frequently the determining factor in the number of young birds available for the next hunting season. A series of torrential cloud-bursts fol lowed by long, cold, rainy spells during the first two weeks of June will cause hundreds of broods to perish.
During severe winters, especially when deep snows cover the ground, the birds are severely pressed to obtain enough food. A successful attempt has been made in Wisconsin to relieve this con~ dition by the establishment of winter feeding stations. Crops of buckwheat and other grains are planted and left in the field to provide food to tide over the birds during these severe times.
One of the major problems involved in the conservation of the prairie chicken is the menacing fires that have swept the prairie regions during the nesting season of the birds. A fire at this time will destroy hundreds of nesting birds and their nests and eggs and in the course of a few hours undo the work of years of conservation work. Fires in fall destroy quantities of prairie-chicken food and the much-needed cover, without which the birds are left exposed to predators. The encroachment upon the breeding and feeding area by agriculture has long been recognized as a factor that has affected the status of the prairie chicken in the Middle West. This unfavorable situation is being relieved somewhat by the establishment of large State game preserves, on which the birds are given absolute protection and where conditions are systematically improved for the birds. The maintenance of winter feeding stations has been especially helpful in tiding the birds over the times when deep snows cover most of the normal food supply. Intensive hunting has done much toward decimating the numbers of prairie chickens. The automobile and the fine modern roads have all been in favor of the hunter and against the birds.
Predators in their relation to game birds are important, but the value of vermin control is frequently overestimated. The wholesale killing of all hawks and owls, for example, should be rigidly avoided, for in the past this practice has actually acted as a boomerang to the objective of conservation of game birds.
Diseases and parasites of birds have not been well known in the past, but they are now becoming to be recognized as important factors in the life of our game birds. Under ordinary conditions, diseases and parasites may be of minor importance, but just as soon as the vitality and normal resistance of the birds are lowered by a series of adverse conditions, such as severe weather and scarcity of food, diseases and parasites manifest themselves and become of prime importance. It is the exceptional bird that is not parasitized, and hence this menace is ever present. There is also danger of infectious diseases, such as blackhead, which has been found to affect the prairie chicken and which figured in the decline of the heath hen. It is highly probable that the cycles in the grouse population are primarily dependent on some disease, either in itself or in combination with other factors. The evidence points to the conclusion that the vast majority of the parasites and diseases of our game birds have been introduced through poultry and exotic game birds. It is apparent that the adaptation of the prairie chicken to the conditions imposed by civilization is not a simple matter. In this adjustment, birds such as the prairie chicken will require much organized assistance on the part of conservation commissions, sportsmen, and bird lovers.
Courtship: The courtship of the prairie chicken generally begins during the first warm days that lay bare the open fields of the winter’s accumulation of snow. Though an early beginning may be made, the courtship does not reach its maximum until the latter part of April or the first of May, when companies of prairie chickens may be .seen collected in favorable, often traditional, spots of the open fields throughout the prairie-chicken country.
0. M. Bryens, of Luce County, Mich., reports the first booming, or ‘crowing,” as it is generally termed in the Middle West, as March 22, 1925; March 13, 1926; April 17, 1928; and March 27, 1929. According to Prof. W. W. Cooke, the booming of the prairie chicken was from March 7, at Caddo, Okla., to March 24, at Barton, N. Dak.
The courtship season continues through the month of May, but the vigor of its execution diminishes and the number of individuals that take part decreases as the sets of eggs hidden in the grasses of the prairie are completed and the domestic duties of incubation on the part of the female begin.
A few birds were still booming on the prairies near Hancock, Wis., when I arrived there the first week of June, 1929, and birds were also booming the second week of June, 1930, in various parts of Wood and Waushara Counties, central Wisconsin. I heard no booming and obtained no authentic accounts of birds booming after the second week of June.
The courtship of the prairie chicken is similar to that described for the heath hen, but since these performances are such an important part of the behavior of this bird a number of interpretations as made by other observers are of great interest.
Dr. Frank M. Chapman (1908) has given us a very vivid account of the prairie chicken as he observed it in the sandhills of Nebraska: At short range tbo bird’s note suggested the mellow resonant tone of a kettledrum, and when bird after bird, all still unseen, uttered its truly startling call, the very earth echoed with a continuous roar. As a rule, each bird had itu own stand separated by about ten yards from that of its neighbor. The boom Is apparently a challenge. It is preceded by a little dance in which the bird’s feet pat the ground so rapidly as to produce a rolling sound. This cannot I)e heard for a greater distance than 30 yards. It is Immediately followed by the inflation of the great orange air sacks at the side of the neck, which puff out as quickly as a child’s toy balloon whistle; the tail is erect and widely spread, the wings drooped, the neck tufts are raised straight upward, giving the bird a singularly devilish look, then with a convulsive movement of the lowered head, the boom is lerked out and at its conclusion the air sacks have become deflated.
One might imagine after so violent a performance the bird would feel a certain sense of exhaustion or at least quiescent relief, but his excess of vitality seeks still other outlets; uttering hen-like calls and cacks he suddenly springs a foot or more straight into the air, whirling about as though he were suffering from a combined attack of epilepsy apd St. Vitus dance. But all this activity Is only a prelude to the grand finale of actual combat. Like a strutting turkey cock, the neighboring birds go towards each other by short little runs, head down, the orange eye-brow expanded and evident pouch inflated, neck tufts, and tail straight up, and looking like headless birds with two tails. Their mating is followed by no make-believe duel but an actual clash of wings. Uttering a low, whining note they fight as viciously as game cocks; and the number of feathers left on the ground testifies to effective use of bill and claws.
First bird called at 4.40 and by seven o’clock the performance was practically over.
A prairie cock when in the lists is a strikingly conspicuous creature; he wears no adornment which cannot be concealed at a moment’s notice. The sight of a passing hawk changes the grotesque beplumed, be-oranged bird into an almost invisible squatting brownish lump, so quickly can the feathers be dropped and air sack deflated. With woodland birds so great a change is unnecessary, hut the prairie hen can hide only under its own feathers.
H. L. Stoddard (1922), in notes from southern Wisconsin, says that “the ‘cooing ground’ at the sandy west end of Sauk Prairie has been used each spring for over 30 years, the birds always using the same knoll whether in rye, stubble, or grown to grass.” Cooing started early in March and continued well into June. The birds arrived early; some were on the grounds before daylight, but on other occasions the bulk came shortly after daylight. The cooing is a “resonant C-A-O-O-O-O-O, H-O-O, H-O-O, rising and in the same tones as do re mi of the musical scale.” This note carries a long distance. “I have heard it over water when the nearest land was nearly 2 miles away.” Two cackling calls were like that of roosters, “one a loud Ka-Ka-Ka-Ka-a-a-a-a and the other a longdrawn q-u-a-h.”
Alexander Sprunt, Jr., says in his notes:
I witnessed a dance one afternoon of five pairs of the birds which came to a sudden end in a strange manner, and one which would thrill the heart of an ornithologist. Lying ensconced behind a log, I was reveling in the eavesdropping act of witnessing the ludicrous antics of five males, who, with air sacs inflated, tails spread, and wings drooping, were bobbing up and down like corn in a hopper, about an admiring group of hens. The booming was intense and incessant, all having something to say at once. Suddenly, without a moment’s warning, a huge snowy owl appeared from behind a low ridge at the far edge of the dancing ground and on widespread wings shot low over the dancers at a height of about 3 feet. Like so many feathered bombs, the chickens scattered to right and left, and In an instant the dancing ground was deserted. No attempt was made by the disturber to follow any of the revelers.
Nesting: The nest of the prairie chicken is invariably on the ground, but the character of the vegetation in which it is built reveals considerable individual variation. Generally the nesting site is among grasses and weeds or low shrubbery in very open situations, but sometimes it may be adjacent to trees and woodlands and in rare instances may be surrounded by trees of considerable size. The vege. tation about the nest is usually very thick and effectively conceals the eggs and the incubating bird from view. It also serves as a protection from extremes of temperature. There are sometimes killing frosts during the nesting season, in May, and there are many days in June when the heat is great enough to kill the embryos if left exposed to the direct rays of the sun for any great length of time.
The nest is placed in a natural hollow of the ground, or a slight excavation may be made by the bird by scratching out the loose earth and then molding the cavity to conform to the size and shape of the body. In this cavity the bird places a scant quantity of nesting material, in some instances the nest lining being little more than the bent-over blades and weeds growing about the structure.
The following descriptions serve to represent the character of the nesting site as well as the nature and construction of the nests built in three different types and situations located in central Wisconsin: A prairie-chicken nest containing 17 eggs was found 4 miles southeast of Bancroft, Portage County, on June 4, 11)29, in a small clearing of a jack-pine grove, the trees of which ranged from 35 to 50 feet in height. The trees of the clearing had been cut the year before and piles of brush left in place. Some of the brush was more or less hidden by the rank growth of grass and weeds which had sprung up around it. The nest, built in a very shallow depression, 4 centimeters deep and 18 by 20 centimeters in diameter, was near one of the piles of brush. Some of the smaller branches were arched over the nest when found. It was protected by the brush on one side, but on the other it was well exposed to view, a condition very favorable for observations and photography from the blind that was later placed in position. Several of the pine trees were so near that. they provided shade for the nest during certain hours of the day. Although this nest was not built in the usual surroundings, it is interesting to note that there were extensive marshlands all about the site.
In the drainage area of Wisconsin there are isolated areas of high ground locally called “islands,” a name originating from the days before the drainage ditches, when they were in reality “islands” during the rainy period of the year. One such island of about 100 acres near Bancroft, Wis., is called Prairie Chicken Island, because these birds have always lived and nested there in unusually large numbers.
Since agriculture has been encroaching on the original habitat of the prairie chicken, many of the birds have adapted themselves with more or less success to conditions created by farming activities. A nest of the prairie chicken was found on June 24, 1929, on a farm 10 miles northeast of Friendship, Adams County, Wis. The nest was on rather high meadowland and was completely surrounded with a luxuriant growth of clover, timothy, and other grasses. The eggs were well concealed by a beautiful canopy of red-clover blossoms. The bowl of the nest measured 15 by 19 centimeters in diameter but only 4 centimeters in depth. The lining of the nest consisted of grasses and weed stems, all apparently picked up from the vicinity of the nest. The clover field was bordered on one side by a low, wet marsh, in which various sedges and rushes prevailed; on the other side was a cleared area being used as a potato field.
Other individuals cling to the more remote prairie districts away from farms, often in situations occupied in common with the sharptailed grouse. A nest containing 11 eggs was found in such a situation in Portage County on June 17, 1929. It was placed among tufts of sedge (Carea’ stricta). The nesting cavity was 18 by 21 centimeters in diameter and 8 centimeters deep. The lining of the nest was made up entirely of sedge, among which were a few feathers of the incubating bird. Near the nesting site was a low, wet marshy area, and on the other side there were thickets of small willows and poplars. No farmland or farm buildings were within 3 miles of this location.
Eggs: The background color of the eggs varies from a “dark olive-buff” to a “grayish olive” tint. Most of the eggs are dotted with many fine and a few larger spots of “sepia.” The spotting varies considerably in different eggs from those with scarcely any marks to those with many fine dots and 20 or more well-defined spots ranging from 1 to 2 millimeters in diameter. Sets of eggs found in open prairie regions seemed to have less spotting and the color of the spots was a “vinaceous-buff” rather than the dark markings of sepia present on eggs found in nests located in the wooded sections of the State. This difference in coloration, however, is probably a mere coincidence and is not to be correlated with a consistent difference in habitat.
Nine sets comprising 100 eggs, all found in central Wisconsin during the summer of 1929, were weighed and measured. The average long diameter was 44.86 millimeters; average short diameter, 33.59 millimeters. The eggs showing the four extremes measured 49.1 by 34.1, 45 by 35.3, 40.5 by 32.9, and 40.8 by 30.2 milLimeters.
The laying of the eggs as determined by studies of captive birds extends over a period of days equal to nearly twice the number of eggs in the set. Apparently approximately the same ratio holds for birds living under natural conditions. In a nest in Adams County, Wis., the first egg was found on May 5, 1929; on May 12 there were 5 eggs; and the set of 11 eggs was completed on May 22. The time required to complete a set of eggs depends on a number of factors, such as the condition of the weather, the health of the bird, and the available food supply. The laying of the eggs is not necessarily on alternate days, but more apt to be very irregular. Certain eggs of the set are laid on successive days to be followed by a lapse of two days before the next egg is deposited. During the laying period of one individual under observation the bird covered the rather exposed eggs with nesting material before she left them. This instinctive habit may be for the purpose of concealment or for protection from extremes of temperature or for both.
The number of eggs layed by the prairie chicken based on studies made of 40 nests, in which the number of eggs was presumably complete, varied from 7 to 17. The average number was 11.5 eggs per set. A nest containing 21 eggs has been reported, but such unusual sets probably represent the eggs of two females using the same nest.
The dates when the nests were found are of some interest to those who may desire to know the probable date when a nest of the prairie chicken may be found. Of 41 nests in which the date of finding was recorded, 20 were found in May, 18 in June, and 3 in July. The earliest date was of a nest found on May 5, when it contained a single egg, and the latest was a nest found July 10 that still contained eggs on July 15, the last time it was visited. The average date when the 41 nests were found is June 3.
Judging from these results we may say the last week of May and first week of June are the times when one may expect to find the largest numbers of nests of the prairie chicken in central Wisconsin.
Young: The incubation of the eggs is performed by the female. Soon after the courtship season the male goes into retirement and undergoes the ordeal of molting. I have never seen the male bird near the nest, nor have I ever observed him participating in the care of .the young.
Incubation begins soon after the set of eggs is completed, but in certain cases where we have found live embryos in eggs left in the nest after the rest of the brood had departed it was evident that incubation had started before these eggs were laid. Sometimes an unusual disturbance about the nest may delay the start of incubation for several days, thus making the determination of the incubation period under normal conditions in the field a difficult task. This accounts for discrepancies in the determinations made by different observers, which vary from 21 to 28 days as the incubation period for the prairie chicken. In the case of a nest under continuous observation, incubation began on May 22 and the eggs hatched on June 14, establishing an incubation record of 23 days. The incubation period for the eggs of the closely related heath hen, as determined by Dr. George W. Field, is 24 days.
After incubation is started the prairie chicken, under normal conditions, remains faithful to her duties through the vicissitudes of weather, storms, and dangers of attacks from enemies. lIJnless flushed from the nest she leaves only for very short intervals to feed, usually at dawn or late in the afternoon about sunset, times when the eggs will not be exposed to extreme heat. Excessive heat is very destructive to the embryos, and great care must be exercised in flushing birds away from nests in which the protecting vegetation has been removed for purposes of photography. The birds that left the nest normally slipped off it quietly and made no attempt to cover the eggs, as they did during the laying period when the nest was abandoned for a longer time. One bird, after nervously surveying her surroundings, sneaked off the nest and walked briskly in a crouched position until she had gone several yards from the nest. There she hesitated, elevated her head, and looked about as if to determine whether her movements had attracted any attention. She then casually nipped at the grasses as she walked along, and finally when about 25 yards away from the nest she arose with a loud whir of wings and disappeared in the scrub pines, where she probably found at least a part of her meal. After an absence of half an hour she flew into view, circled the nest, and alighted in the tall grass 15 to 30 yards away from the nest. At first she crouched in the grass completely concealed from view, but after being assured all was well she walked along stealthily though not directly toward the nest. It was not possible to keep her in view at all times, but now and then she would come to an open place and from this vantage point more carefully scrutinize the surroundings. Sometimes she completely encircled the nest and blind with a wide radius and frequently retraced her steps to make a careful inspection in order to satisfy herself that no spying enemy was near. One could not be sure whether this behavior was prompted by apprehension of harm to herself, fear of revealing the presence of the nest, or to both. After these maneuvers, which usually took about 20 minutes, were completed, she appa,rently was assured, and then without hesitation she walked quickly and directly to the nest. The position she assumed on the nest varied, and one could never be sure whether she would be facing toward the blind or away from it. The ruffed grouse, which builds its nest at the base of a stump, a log, or tree, invariably faces away from the side of the nest thus protected from the sneaking approach of some prowling enemy. Prairie chickens, which nest in thick vegetation, will usually face toward the side that is opened up for purposes of photography, a very desirable position for the photographer. Many of the prairie chickens studied exhibited extreme restlessness and made much more of a task of incubation than does the ruffed grouse. One continuously shifted her position during the course of the day; at other times she would pick aimlessly at the nesting materials, and not infrequently she would reach far out of the nest at some unsuspecting grasshopper or other insect that chanced to alight on the tall grass. Certain females revealed a sensitive, nervous temperament and quickly responded to any stimulus, whether it was the caw of a crow or the hum of a distant tractor; and even the shadow of a passing cloud was sometimes sufficient to make her respond. When startled she would frequently elevate her head to command a wider view of her surroundings. If she caught sight of the source of her alarm, such as the passing of a dog, she would retract her head, become perfectly motionless, and retain a “frozen~’ position until the source of danger passed. If the animal ventured too near she would fly off with a violent “whirrrr” of her wings, which was sure to attract, if not startle, the intruder. She would drop into the grass a short distance away, utter a sharp distressing cry, and feign a wounded bird. After several repetitions of the performance, until she had attracted the enemy away to a safe distance from the nest, she would sail gracefully away, leaving the bewildered creature behind. This behavior is common to many birds, hut the deception is remarkably well executed by the prairie chicken. I have seldom seen a prairie chicken try these tactics with a human being; apparently they have learned that it is best in such cases to get out and away without the least delay. That the leaving of the nest when flushed by a human being is a quick performance is revealed by moving pictures. The usual speed is 18 frames per second in the ordinary moving pictures. It requires only three or four frames of the picture to show the bird until she is away from the scene. This means that it requires only threesixteenth to one-fourth of a second for the bird to leave the nest.
As it is not possible in most cases definitely to establish the cause of nest destruction, it is necessary to depend on circumstantial evidence. In 1929, out of 12 nests studied in Wisconsin only three reached the period of hatehing. The adult birds of two of the nests were killed and the eggs destroyed, presumably by coyotes, as the tracks of the animals were found in the sandy soil around the nests, and the mass of feathers left behind seemed to indicate the work of such animals. One incubating prairie chicken was killed by a horned owl; the eggs of another located in an open situation were destroyed by crows, and in two other cases the eggs disappeared without a trace of the intruder. One nest was accidently destroyed by a farmer while plowing, and the eggs of two others were deserted where it was problematical whether the incubating birds were taken by some predacious bird or mammal or were merely frightened away in some manner.
During the summer of 1930, 28 prairie chicken nests were found, of which 17 reached the period of hatching and the others failed for one reason or another. Indirect evidence indicates that crows were responsible for the destruction of three nests of eggs. One was probably broken up by a dog, four were drowned out by floods, in one case the bird was killed, presumably by a mink, within a few feet of the nest, in another case the embryos in the eggs were killed by exposure to the heat of the sun, and one nest was deserted. Combining the records of 1929 and 1930, we have 40 nests of which 3 hatched in 1929 and 17 in 1930, 20 in all, or an average of 50 per cent for all the nests observed.
Every egg hatched of the three nests that reached the stage of hatching in 1929. The 17 nests that succeeded in reaching the hatching stage in 1930 contained 208 eggs. Twenty-nine eggs, or approximately 14 per cent, failed to hatch. Only 8 of the 17 birds succeeded in hatching every egg, and in 9 nests there were one or more sterile eggs, eggs with dead embryos, or both. Of 29 eggs that failed to hatch, 6 were sterile and 23 contained dead embryos. The latter were killed by excessive heat of the sun or by failure of the eggs to hatch in time before the old bird left the nest with her young.
During the summers of 1929 and 1930, the date of hatehing was noted for 23 sets of eggs in the field or by special incubation, Of these, 3 hatched in May, 17 in June, and 3 in July. The earliest date of which we have a record was May 29, 1930, and the latest July 7, 1929. Nest No. 12 contained eggs on July 15, 1929, but it was not possible to record its hatching. The average date of hatehing of the 23 nests was June 10. These records indicate that the majority of the nests hatch during the first two weeks of June. The condition of the weather at this time is a most important factor in the determination of the number of birds to be expected the following season. A long continuous cold rainy spell with cloud-bursts, such as is sometimes experienced in the Middle West during the first part of June, is certain to have a disastrous effect on the broods of young birds.
The first eggs are pipped on about the twenty-second day after incubation starts. The shell is slightly raised at the point on the circumference between the blunt and pointed ends, but slightly nearer the larger end. The embryo inside the egg can be heard to peep ” at this time, an event that greatly excites the mother and at once becomes a great stimulus in her behavior. It causes her to reach under her breast feathers to turn the eggs every few minutes and to exhibit a great deal of nervousness in her response to various other stimuli.
On the morning following the day on which the eggs are slightly pipped, all the eggs destined to hatch are pipped. In those eggs where there was but a slight elevation in the shell the day before, there are now well-defined openings through which the tip of the bill of the embryo can be clearly seen. At this stage the mother bird may frequently roll the eggs with her body in addition to turning them with her bill. I have even observed the old bird pick out bits of shell from a pipped egg, as if attempting to facilitate the process of hatching. After a few hours the calcareous shell is cracked for its entire circumference, but the shell membrane may remain intact for a longer time. The struggles of the young, however, soon fling the cap open with a part of the shell membrane on one side serving as a kind of hinge. After the embryo has kicked itself out of the. prison shell, the tension of the drying shell membrane pulls the cap back in place and thus prevents the youngster from being cupped by its own shell.
The time of hatching of the various eggs of a set is remarkably uniform, and in some instances the time elapsed from the time of hatching of the first to the last egg is less than an hour. In cases where incubation started before the last one or two eggs were laid, the latter may be delayed, and in several cases under observation. during the summer of 1930, they failed to hatch in time, and the contained young were left behind when the brood left the nest.
The precocious young of the prairie chicken are ready to leave the nest as soon as their down is dry, and the mother bird often has difficulty in preventing the first young from leaving the nest before the last to hatch are prepared to go. The brood may leave in a few hours after hatching, but if the hatching takes place late in the afternoon the old bird unless disturbed, will brood them on the nest during the night, but leaves the next morning just as soon as the temperature and weather conditions are suitable for the chicks to move. The eggahells are never removed from the nest by the old bird.
The chicks, as well as embryos in the eggs, are very sensitive to extremes of temperature. Young left exposed to the cool damp night air will quickly perish, and brooding at such times is absolutely necessary. Though the young quickly perish if cold, especially if wet, they will make rapid and remarkable recoveries from nearly paralyzed conditions. Young that seem almost lifeless can be quickly restored to an active condition by merely holding them in your hand and blowing your breath over them for 10 or 15 minutes. In one instance a mother bird was frightened away from the nest at night and failed to return. The young, still damp from the fluids of the eggs, seemed destined to die. They were taken inside the blind, revived, and then kept alive and contented all night by placing them against my body inside of a flannel shirt. At daybreak the old bird appeared on the scene and claimed the youngsters one by one as they were released under the burlap of the blind.
The language of the prairie chicken is readily understood by the young, even when first hatched. If the young are taken inside of the blind at the time of hatching they are indifferent to various sounds and notes of other birds calling outside of the blind, but as soon as the adult prairie chicken appears and begins calling they respond at once. If she gives her brirrrb: brirrrb call, they struggle all over one another trying to get out, but if the old bird becomes alarmed and suspicious she gives a sharp shrill call of caution and immediately each little chick cows down and “freezes” to a perfectly motionless pose. Chicks set free at the edge of the blind made no effort to go to the mother bird unless called. As soon as the call note is given there seems to be an irresistible impulse on the part of the young to follow that call, although they can not see the old bird. When the chicks wander from the nest at hatching time that same call brings them back, and hence this response is important in their preservation and doubtless is a matter of evolutionary development. Though the character of the call has a distinct meaning to the birds, one can with little effort imitate it and completely deceive the adult or young. I have often made use of this fact by inducing the bird to come near enough to the blind to obtain large portrait pictures of her. One can so excite the bird that is nearing the hatching time that she will exhibit an unusual behavior, such as turning the eggs over and over or twisting and squirming about the nest. She will, if the call is well imitated and continued, leave the eggs to search for the young. The bird at one nest under observation circled the blind again and again and even attempted to get under the burlap to reach what she apparently supposed was a young in distress. The old bird seems to have but little resourcefulness in aiding a young in unusual situations. In one case where the bird was taking her young away from the nest, two of the young accidentally fell into a deep horse track. The young called desperately, but the mother seemed helpless. She raced around the opening several times and then settled down in the grass to call for them. These two young would have perished without my intervention. An experiment of placing young in a hat near the nest also proved the bird’s lack of resourcefulness to cope with an unusual situation.
Few things in nature have a greater human appeal than a family of gallinaceous birds. The whole scene from the hatching of the first young to the departure of the brood is one brimming with thrilling incidents. The motherly interest of the old bird when the first youngster pokes its head through the breast feathers and gives a contented peep, as it picks at its mother’s bill or her eye, is an event never to be forgotten. Then the unexpected poking of a downy head through the plumage, first at the side, then through a rear window, and perhaps two youngsters surprising each other as they appear simultaneously, all are experiences that make a long vigil in the blind well worth the effort. As more of the young hatch they become more daring and may vigorously compete for a position on the mother’s back. They make repeated attempts to scale the slippery feathered dome, and finally when one does succeed he has an unmistakable look of triumph. All these things seem to have a truly human aspect, and surely the most skeptical can not help but take an anthropomorphic attitude toward their behavior.
Ordinarily a few hours after the young are hatched the old bird leaves the nest, allowing the young to follow after her. She generally goes a few yards, settles in the grass and then continues calling until all the young are gone. This procedure is repeated until she is well away from the nest. During this time the young are brooded a great deal, but before many hours, especially if the day is warm, they become active in searching for insects and other food.
Plumages: In a prairie chicken a few hours old the chin and lores are “primuline yellow,” sides of head, including down on eyelids, “naphthalene yellow,” throat and breast “wax yellow,” remainder of underparts “barium yellow,” and down on tarsus “straw yellow.”
The lighter areas above are yellowish, strongly tinged with “cinnamon,” which approaches “Mikado brown” on the rump, and the entire upper parts are marked with numerous irregular black spots and patches. There is a small black patch hack of the eye and three irregular shaped black spots in the auricular region. The iris is “dark Quaker drab,” the base of the upper mandible ïis “pecan brown ” and tipped with a lighter color, and the upper surface of the mandible is black, which extends for a distance equal to two-thirds the length of the bill. The lower mandible is pale “flesh color,” tipped with “straw yellow.” The posterior part of the tarsus, not covered with down, is “yellowish citrine.” The upper surface of the toes is “honey yellow,” the undersurface “mustard yellow,” and the nails “flesh color.”
The young of the sharp-tailed grouse is similar to that of the prairie chicken, but with the following minor differences: The yellow of the underparts, not so deep or so bright as in the prairie-chicken young. Upper parts with much less black, especially in the region of the back and rump. The brown of the rump is a paler shade. The black on the upper mandible extends down only one-half the distance of the length of the bill. When the young of the prairie chicken and sharp-tailed grouse are compared with young of the ruffed grouse, they are seen to be much yellower and much less reddish brown, so characteristic of the day-old ruffed grouse.
The average weight of 17 young prairie chickens that hatched in a nest near Bancroft, Portage County, Wis., was 15.9 grams. The average weight of the eggs 2 days before hatching was 19.4 grams.
[AUTHORS NOTE: As with all young grouse, the wings and scapulars begin to grow soon after the chick is hatched, and juvenal plumage is rapidly acquired, long before the young bird attains its growth. In this plumage the crown is “hazel,” spotted with black; the feathers of the back, scapulars, tertials, and wing coverts are boldly patterned with “ochraceous-tawny,” black, and “snuff brown,” many feathers, especially the scapulars, having broad white shaft streaks spreading out into a white tip; the primaries are spotted with pale buff on the outer web; the pointed tail feathers are barred or patterned with the colors of the back; the chin is white, and the underparts are dull white, washed with buff and spotted with dusky on the breast and flanks.
Before the young bird is fully grown, in July, the postjuvenal molt begins with the primaries. This is a complete molt, except that the outer pairs of juvenal primaries are retained for a whole year. Otherwise the first winter plumage is practically adult. Adults have a partial prenuptial molt, about the head, in March and April, and a complete postnuptial molt in August and September.]
In addition to the normal plumages of the prairie chicken there are unusual types that have attracted the attention of sportsmen and ornithologists. In certain individuals of the prairie chicken there is a prevalence of rufous or reddish brown, which is due to an excess of red pigment in the feathers, a condition known as erythrism. The red phase of the plumage is a common occurrence among ruffed grouse, but as yet it has been noted in comparatively few cases of pinnated grouse. The following cases, which have come to my attention, are of interest:
George N. Lawrence (1889) described a specimen in which all of the light markings were tinged with light, bright rufous and the entire underparts, throat, and neck tufts were deep rufous (reddish brown).
William Brewster (1882 and 1895) called attention to four specimens exhibiting the red phase in which there was but little variation with respect to the depth and extent of the reddish brown or chestnut coloring. The upper parts of the birds were strongly suffused with reddish brown, while most of the underparts were clear reddish or rusty chestnut and the usual blackish chestnut bars were nearly or quite wanting on the sides. All four specimens examined by Brewster were males.
Cases of albinism, in which there is a lack of pigment, resulting in a white plumage, have been frequently noted. Some of the albinos are not pure, but may have a little pigment in certain of the feathers, giving them a dusky appearance.
An albinistic specimen collected March 6, 1898, near the Missouri River, Iowa, is in the collection of the Museum of Comparative Zoblogy, Cambridge, Mass. This bird is white, with the exception of minor, pale, rusty brown crossbars and markings.
J. A. Spurrell (1917) records a very interesting case of an albino in Sac County, Iowa, which attained quite a local reputation because the bird was so clever that it eluded all attempts to trap it. The hunters in the vicinity made a point to spare the “white chicken” in the hope that it might be captured alive.
Hybrids between the prairie chicken and the sharp-tailed grouse have been noted by many observers. In Wisconsin, where the ranges of the two species overlap, it is a common experience to see them associated together at all seasons of the year, and it is not at all surprising that they frequently interbreed.
J. H. Gurney (1884) described a male hybrid between the sharptailed grouse and the prairie chicken in which the pinnae were present, but only one-fourth of an inch long. The tail was a hybrid gray between the brown of the sharp-tailed grouse and the white of the prairie chicken. The sides of the toes were only slightly feathered and the general coloration was intermediate between the two species.
A hybrid between the prairie chicken and the sharp-tailed grouse in which the elements of the prairie chicken predominated has been reported by F. C. Lincoln (1918). The author has examined a hybrid specimen in the possession of Mrs. H. M. Hales, of Hancock, Waushara County, Wis., which also resembles the prairie chicken in most of the characters of its plumage.
William Bowan (1926) figures and describes two female hybrids of the sharp-tailed grouse and the prairie chicken collected in Alberta, Canada. One individual collected near Edmonton resembles the pinnated grouse more closely than the sharp-tailed grouse, while the other, shot at Gough Lake in the southern part of the Province, resembles more nearly the sharp-tailed grouse. The ovary of the Edmonton bird, according to Mr. Rowan, was normal. These are the only Alberta hybrids known to him, but he states that hybrids between the prairie chicken and the sharp-tailed grouse are frequent in Manitoba, where the pinnated grouse is more numerous than it is in Alberta.
Glenn Berner, of Jamestown, N. Dak., writes that he killed a hybrid grouse in 1923 in which the back, head, and tail resembled the prairie chicken, whereas the breast, legs, feet, and under tail parts were like those of the sharp-tailed grouse. The breast was not barred as in the prairie chicken but spotted as in the sharp-tailed grouse. The bird when flying had the characteristic cackle of the sharp-tailed grouse.
According to 0. ‘A. Stevens, Fargo, N. Dak., there is a hybrid in the collection at the North Dakota State Agricultural College.
I have examined a female and three male hybrid specimens of the prairie chicken and sharp-tailed grouse in the Museum of Comparative Z&5logy, at Cambridge, Mass., which were obtained in the Boston markets, March 24, 1873, February, 1887, December 29, 1899, and January 24, 1893, respectively. So far as I know, these hybrids do not reproduce themselves, and in most cases this is probably due to the sterility of the individuals. Food: The prairie chicken, like other grouse, is adaptable in its food eating habits, varying its diet from season to season and sustaining its life on the food that is most abundant and easily obtained.
Dr. Sylvester D. Judd (1905a) reported on the examination of 71 stomachs of prairie chickens collected in the Middle West and representing all months of the year except July. The food consisted of 14.11 per cent animal matter, chiefly grasshoppers, and 85.89 per cent vegetable matter, made up of seeds, fruit, grain, leaves, flowers, and bud twigs.
According to Judd’s report the prairie hen is highly insectivorous from May to October inclusive, insects constituting one-third of the food of the specimens shot during this period. The species is particularly valuable as an enemy of the Rocky Mountain locust. During an invasion by this pest in Nebraska, 16 out of 20 grouse killed by Prof. Samuel Aughey (1878) from May to October inclusive, had eaten 866 locusts. Beetles and miscellaneous insects were eaten in smaller numbers.
From October to April, inclusive, according to the Biological Survey report (Judd, 1905a), the prairie hen takes little but vege table food, consisting of fruit, leaves, flowers, shoots, seeds, grain, and miscellaneous vegetable material. It is especially fond of rose hips, which comprised 11.01 per cent of the food. When the deep snow causes scarcity of other supplies, the sumac affords the prairie hen with abundant food. Seeds make up 14.87 per cent of the annual diet, of which grass seeds form 1.03 per cent, seeds of various polygonums 8.49 per cent, and miscellaneous weed seeds 5.35 per cent.
The prairie chicken eats more grain than any of the other native gallinaceous birds; the food examined by the Biological Survey was 31.06 per cent grain. The stomach of one bird shot in June in Nebraska contained 100 kernels of corn and 500 grains of wheat. Buckwheat, barley, oats, and millet are relished, but corn appears to be the favorite cereal, amounting to 19.45 per cent of the food. Wheat was next in order represented by 11.61 per cent. Like other gallinaceous birds, it is fond of mast such as hazelnuts and acorns, though it obtains much less than the rufl?ed grouse. A bird shot in Minnesota in March had bolted 28 scarlet-oak acorns. An analysis of organic material in the food of 17 prairie chickens collected in Wisconsin for the Wisconsin Conservation Commission, chiefly in fall, revealed that about 28 per cent of the food was animal and 72 per cent vegetable matter. Gravel constituted 6 per cent of the combined organic and inorganic material of the crop and stomach contents. The average weight of the crop contents was 25.7 grams; the maximum 83 grams. The average weight of the stomach contents was 14.3 grams, and the largest quantity found in any one stomach weighed 23.1 grams.
There were 84 kinds of vegetable matter and 82 kinds of animal matter represented in crops and stomachs of the prairie chickens collected in Wisconsin. Arranged in order of the percentages of the entire food eaten by the birds, the 25 more important foods follow: Short-horned grasshoppers, 26.7; ragweed, 11; oats, 10.8; clover, 7.7; black bindweed, 6.2; acorns, 4.5; greenbrier, 3.6; dogwood, 3.5; crickets, 3.3; buckwheat, 3.1; bramble, 3.1; blueberries, 2.4; rose, 1.7; hawkweed, 1.4; chokeberry, 1; galls, 0.94; ants and wasps, 0.88; poison ivy, 0.80; birch, 0.80; pin cherry, 0.64; woody d~bris, 0.64; bunchberry, 0.53; wild black cherry, 0.53; smartweed, 0.47; pigeon grass, 0.47.
It will be seen that the chief difference between the above list of foods and the results published by the Biological Survey in 1905 is the absence of corn in the recent list. In Doctor Judd’s list corn made up 19.45 per cent of the entire contents of all the birds examined. This may be accounted for by the change in methods of farming. In the past, corn was husked in the field and much grain was accidently left behind by the harvesters. To-day, in Wisconsin, a dairy State, all the corn that is raised is cut and made into ensilage for the cattle. Practically no corn is allowed to ripen in the field; hence it does not appear as a food for the prairie chicken. Both lists agree in the large percentage of grasshoppers comprising the food. The prairie chicken and sharp-tailed grouse are notable grasshopper consumers, which fact, together with their fondness for weed seeds, makes their presence a great asset to the farmers. Except in the northern part of their range, where very severe weather and deep snows prevail, there is sufficient natural food for the prairie chickens at all seasons of the year. The prairie chicken is there hard pressed for an existence, since it does not seem to be able to subsist on buds and other foods above the snow to the extent that it is done by the ruffed grouse. In Wisconsin, experiments conducted with winter feeding stations by the conservation commission have proved a great success. Plots of ground ranging from a half acre to two acres in extent are planted chiefly to buckwheat, with sorghum, sunflowers, broomcorn, and corn planted as accessory foods on most of the plots. Half of the crop is left standing and the other part is cut and placed in covered shocks, which are opened up after the deep snows arrive. According to the reports of the wardens in charge, as many as 200 to 300 birds visited a single station at one time, a strong testimonial for the practicability of such stations in game management.
Migration: The prairie chicken is a permanent resident in much of its range, but in the Northern States there is a regular annual movement of the birds southward at the approach of winter weather. There are counties in Wisconsin where prairie chickens do not breed, or are present in very small numbers during the summer season, whereas they are represented by large numbers of individuals during the winter months, especially when deep snows and extremely cold weather exist in the more northerly sections of the range from which the birds apparently come. Observers in Door County, Wis., have reported seeing flights of prairie chickens approaching the land from Green Bay. The birds supposedly came from the opposite shore, a distance of 12 to 15 miles, which, if true, means that they sometimes take flights exceedingly long and continuous for a bird of the type of the prairie chicken. A. E. Doolittle, superintendent of Peninsular Park, Door County, saw a flock of 300 prairie chickens headed northeast, up the shore of Green Bay, which he thought were en route for the Michigan side of the bay. William Fairchild, former keeper of Chambers Island (near the middle of Green Bay, a distance of about 7 miles from the mainland), saw two prairie chickens arrive from Marinette in April, 1927. They remained several weeks, then flew eastward to Door County proper.
R. M. Anderson (1907), writing of the prairie chicken in Iowa, states: “While a certain number remain throughout the winter, large flocks pass southward early in the winter, returning in March.”
The migration was even more marked in the past when the birds were abundant. J~ A. Spurrell (1917) states that there was a marked migration of birds away from Sac County, Iowa, until about 1875-. 1880. After that date, he says, corn became a common crop and birds wintered as well as nested abundantly in that section of the State. Prof. W. W. Cooke (1888), in writing of migrations of prairie chickens in Iowa, stated:
In November and December large flocks of prairie chickens come from northern Iowa and southern Minnesota, to settle In northern Missouri and southern Iowa. This migration varies In bulk with the severity of the winter. During an early cold snap immense flocks come from the northern prairies to southern Iowa, while in mild, open winters the migration Is much less pronounced. During a cold, wet spring the northward movement In March and April is largeW arrested on the arrival of the flocks In northern Iowa; but an early spring, with fair weather, finds them abundant In the southern tier of counties in Minnesota, and many flocks pass still farther north. The most remarkable feature of this movement Is found In the aez of the migrants. It Is the females that migrate, leaving the males to brave the winter’s cold. Mr. Miller, of Heron Lake, Mlnn., fairly states the case when he says: ‘~ The females in this latitude migrate south In the fall and come hack in the spring about one or two days after the first ducks, and they keep coming In flocks of from ten to thirty for about three days, all flying north. The grouse that stay all winter are males.”
In the spring of 1884, at Iowa City, Iowa, the first flocks passed over March 10, and the bulk March 22; at Newton, Iowa, the bulk was noted March 23.
Glenn Berner, of Jamestown, N. Dak., says in his notes: During the spring of 1924 I witnessed a decided northward movement of prairie chicken flocks numbering 10 to 100, some Quite high in the air: late in the afternoon: possibly 80 flocks being seen from one location In two hours and very few of them alighting.
0. A. Stevens, of Fargo, N. Dak., writes that he saw a flock of prairie chickens fly high overhead on October 25, 1930, a time that coincided with a marked migratory movement of other birds. Mr. Stevens has noted for several years this annual movement taking place during the latter part of October. He considers these flights as a distinct migration, as the birds were always moving in the same direction.
Information on the migration of the prairie chicken supplied by F. C. Lincoln, of the Biological Survey, is as follows:
Although not a true migrant in the strict sense of the word, the prairie chicken has been known to make more or less regular ifights north and south. Curiously, these movements appear to be confined chiefly to the females, the males remaining In the breeding areas during winter. The flights apparently were not infrequent a generation ago when the birds were much more abundant than at present. The extent of the movement and the number of individuals partici pating were dependent upon the severity of the winter In the northern part of the range. The exodus usually took place in November or December (Wisconsin, Madison, November 25; Iowa, Ogden, December 14, and Osage, December 30), while the return trip was made in February, March, or April (Iowa, Sioux City, February 4, Marshailtown, February 7, and Osage, February 13; Wisconsin, UnIty, February 15, Elkhorn, March 2, and Whitewater, April 1; and Minnesota, Fort Snelllng, March 21, and MinneapolIs, April 8).
Thus it is clear that the prairie chicken, at least in the Northern States, makes flights of considerable length, which we can consider of a migratory character. In addition to these movements there are shifts and concentrations of the birds that are very local and mainly concerned with the food supply. At the feeding stations established in Wisconsin it was not unusual for 200 to 300 birds to feed at a single station at one time, a total far in excess of the numbers breeding in the vicinity. Some of the birds may have migrated from the north, but it is probable that the mass of these flocks are merely aggregations from a limited region of a few counties.
Winter: In regions where deep snows prevail, the prairie chickens often dig themselves into the deep drifts to avoid the excessive cold. One observer at Green Bay, Wis., relates observing five prairie chickens alight on the surface of the snow, which was about 2½ feet deep. The birds walked up to some weed stalks that projected through the snow and then dug themselves in at places about 10 to 12 yards apart. A day later the same observer flushed the birds from the snow bank and found well-molded places on the ground among the weeds. There was an accumulation of droppings in each burrow, indicating that the birds had remained in the same spot during the night. The practice of digging into the snow has proved disastrous at times when it becomes covered over with a resistant layer of ice.
Gale W. Monson, of Argusville, N. Dak., says in his notes:
In winter the prairie chickens are our most conspicuous birds. They spend the nights in the tall grass of marshy meadows making small pockets for themselves In the snow. At sunrise they leave their beds and fly to the nearest cornfield, there to eat their fill of that grain. In the afternoon they return again to their sleeping quarters. Their chief enemy at this time is the snowy owl, which sometImes depletes their numbers to a noticeable extent.
Several observers in Wisconsin report that the tall marsh grass is frequented by the prairie chickens as soon as the water of the swamps is solidly frozen over.
John Worden, of Plainfield, Wis., states that during times of deep snows the prairie chickens are often in a semistarved condition. At such times the birds showed little fear of man and often allowed him to approach within a few yards before attempting to fly. A farmer living near Babcock, Wis., stated that in collecting shocks of corn following a heavy snowstorm, he had virtually to drive the birds away, and when flushed they flew but a few yards to the next shock of corn. Such behavior is probably very unusual except under very extraordinary circumstances when the birds are suffering with extreme hunger. According to Mr. Worden, the birds frequently alight in trees during winter, but he says they invariably roost on the ground at night.
F. Hall, of Babcock, Wis., states that during the winter of 1928: 29 a flock of about a dozen prairie chickens came regularly to the poplar trees of his back yard, but he was not certain to what extent they fed upon the buds.
Range: South-central Canada and the United States east of the Rocky Mountains, except the Southeastern States.
The full range of the prairie chicken extended north to southern Saskatchewan (Quill Lake and Indian Head); southern Manitoba (Oak Lake, Carberry, Westbourne, Ossono, and Shoal Lake); northern Minnesota (Crooked Lake); central Wisconsin (Unity, Wild Rose, and West Depere); Michigan, (Chathain, McMillan, Sault Ste. Marie, and Fourmile Lake); southern Ontario (Wallaceburg and Chatham); and Massachusetts (Springfield, Newton, and Cape Ann). East to Massachusetts (Cape Ann and Marthas Vineyard); Long Island, N. Y. (Miller Place and Hempstead); New Jersey (Barnegat); and southern Maryland (Marshall Hall). South to southern Maryland (Marshall Hall); District of Columbia (near Washington) ; probably Virginia and perhaps North Carolina; southwestern Pennsylvania (Blairsville); central Ohio (near Columbus); southern Indiana (Bloomington, Marco, and Bickwell); northwestern Kentucky (Henderson); southern Louisiana (Iowa Station and Calcasieu Pass); and Texas (Beaumont, Richmond, Edna, Port Lavaca, St. Charles Bay, Austin, and Tascosa). West to northwestern Texas (Tascosa); Colorado (Barton and Barr); southeastern Wyoming (Chugwater); northwestern Nebraska (Chadron); South Dakota (Pine Ridge Reservation, Kadoka, and Short Pine Hills); North Dakota (Bismarck, Charlson, and Crosby); and southern Saskatchewan (Johnston Lake and Quill Lake).
A specimen taken in the fall of 1917 near Huntley, Mont., is at present the only record for that State. Prairie chickens have been noted as rare in winter near Fayetteville, Ark., and are said to occur in that season at De Witt, Ark.
The prairie chicken and its eastern relative, the heath hen, have been extirpated over great areas in their former range. The heath hen is, in fact, extinct except for a single bird, which at the time of writing (November, 1930) was still living on Marthas Vineyard, Mass. Western Indiana marks the present eastern boundary of the species.
Many attempts have been made to transplant the prairie chicken into other parts of the country. These have all met with failure, except for an apparent introduction in northern Michigan. They are reported as thriving in the vicinity of Sault Ste. Marie and McMillan.
The range as described is for the entire species, which has, however, been separated into three subspecies. True americanus, the greater prairie chicken, formerly occurred from the southern parts of the prairie Provinces of Canada and eastern Colorado east to southwestern Ontario, northwestern Ohio, and western Pennsylvania. The range of the heath hen (T. c. cupido) included New England (to southern New Hampshire), New York, and other States of the Atlantic seaboard, probably south to and including Maryland. Attwater’s prairie chicken (1’. c. attwateri) is found in the coastal region of Texas and southwestern Louisiana.
Egg dates: Manitoba: 6 records, May 24 to June 20. Minnesota and Dakotas: 30 records, May 1 to June 18; 15 records, May 18 to 29. Wisconsin: 37 records, May 5 to July 10; 19 records, May 28 to June 20. Illinois and Iowa: 32 records, April 20 to June 6; 16 records, May 6 to 25. Marthas Vineyard: 3 records, June 2 and 5 and July 24. Texas: 4 records, April 3 to May 16.
TYMPANUCHUS CUPIDO ATTWATERI (Bendire)
ATTWATER’S PRAIRIE CHICKEN [Current A.O.U. = Greater Prairie Chicken]
The small dark race of the prairie chicken named attwaleri is confined to southwestern Louisiana and eastern Texas, mainly in low prairies in the coastal counties. It was described by Maj. Charles E. Bendire (1894) and named by him in honor of Prof. H. P. Attwater. He gives as the subspecific characters:
Smaller than T. emericanus, darker in color, more tawny above, usually with more pronounced chestnut on the neck; smaller and more tawny light colored spots on wing coverts, and much more scantily feathered tarsus, the latter never feathered down to base of toes, even in front; a broad posterior strip of bare skin being always exposed, even in winter, while in summer much the greater part of the tarsus is naked.
George Finlay Simmons (1925) describes its haunts as “rolling open, grassy, fertile upland prairies, where the grass is from 1 to 3 feet tall, old and thick and mixed with weeds; wheat and corn fields; takes to timber only during snow and sleety storms.”
Referring to its history, Simmons writes:
Formerly abundant on the open prairies, these wonderful game-fowl became extirpated in the Austin Region through two agencies; civilization and hunting. They disappeared rapidly as the country was settled up and as cultivated fields took the place of the extensive, wild, unfenced prairies; and hunters quickly killed the few remaining birds.
Its courtship performances, nesting habits, eggs, plumages, and molts are all similar to those of the common prairie chicken.
Eggs: The eggs are indistinguishable from those of the common prairie chicken. The measurements of 27 eggs average 42.3 by 31.5 millimeters; the eggs showing the four extremes measure 44.9 by 32, 42.4 by 33.5, 38.8 by 28.9, and 39.8 by 28,6 millimeters.
Food: Simmons (1925) says that it sometimes flies to treetops to inspect corn fields before allghting in them to feed; frequently feeds in the open in plain sight of observers several hundred yards away. During early breeding season, feeds largely on insects, such as grasshoppers, crickets, potato bugs, and other beetles; in fall and winter, tops and seeds of leguminous plants, tender buds and green leaves of late winter, fruits, berries, and waste grain of stubble and corn fields.
Behavior: Simmons says that it is observed singly and in pairs in spring; in fall and winter, roamed about in flocks of from 10 to 12 up to a 100 or more, moving about over the prairies and grain fields, generally keeping emong bushes and tall grass inland, the open prairies and grassy knolls along the coast. Stately of bearing, but otherwise very much like the domestic fowl in its actions. In spring, a “scratching ground” or smooth, open courtship ground is selected, where pairIng takes place.
Voice: The sanie observer describes the notes as “nondescrint calls; strange cackles, with a muffled booming love-call, uck-aA-umbboo-oo-oo-oa-oo-oo; and a loud beating iwom-ak-boom, perhaps produced by a beating of the wings; when alarmed, a rapidly repeated cluk-cluk-cluk-cluk; female, when flushed, utters a low kuk-A,uk-kukkuk-kuk-kuk-louk-kuk.”
TYMPANUCHUS CUPIDO CUPIDO (Linnaeus)
CONTRIBUTED BY Airnzo Orxu Gaoss
The heath hen and the prairie chicken are so closely related that they are now considered as geographical races and not as distinct species. In 1885 William Brewster (1885a) called our attention to differences between the pinnated grouse of Marthas Vineyard (heath hen) and the western pinnated grouse, or prairie chicken. He named the western form as the new species, because 7’etrao cupido of Linnaeus was the eastern form from the fact that its habitat is given as Virginia. The differences between the eastern and western birds are so slight and the variations of the individuals so great that ornithologists now concede that Brewster was not well justified in the establishment of a new species.
In prehistoric times the common ancestors of the heath hen and the prairie chicken probably ranged in an uninterrupted distribution from the Atlantic seaboard to the plains east of the Rocky Mountains. Later the birds of the East became separated from those of the West, and as a result of this isolation and differences in environment certain modifications arose that have resulted in the establishment of the two geographical races: Tympanuchus oupido cupido, the heath hen, and Tyrnpanuchu.s oupido americanus, the prairie chicken.
The prairie chicken is still flourishing and in recent years has been rapidly regaining its numbers in favorable sections of the Middle West, but the heath hen has been unable to cope with the changing conditions of its restricted environment, and to-day is represented, so far as we can ascertain, by a single male individual, which is living out its normal life on the scrub-oak plains of Marthas Vineyard Island, Mass.
The following account is made up primarily of modified excerpts of the contributor’s monograph on the heath hen (Gross, 1928) and from subsequent annual census reports:
Historical: The heath hen is among the first of the American birds to be mentioned in the writings of the early colonists who came to our shores. There is, however, such a dearth of material concerning the heath hen during these early times that we know but little concerning the conditions under which it existed, and the records are so incomplete that we are unable to determine with any degree of accuracy its relative abundance and distribution prior to the nineteenth century. Some of the earlier American writers designated the heath hen by the name ” heatheocke,” “pheysant,” or “grous,” but their notes and descriptions are such that they can be clearly referred to this species. William Wood (1635) in his New England Prospect writes as follows: “Heatheockes and Partridges be common: he that is husband, and will be stirring betime, may kill halfe dozen in a morning. The Partridges be bigger than they be in England, the flesh of the Heathecocks is red, and the flesh of the Partridge white, their price is four pence a piece.” Wood resided at what is now the city of Lynn in Massachusetts. His map included Cape Ann and the Merrimac River; hence it is evident that the heath hen existed in northeastern Massachusetts in his day. Thomas Morton (1637), writing concerning the heath hens, which he called “pheysants,” stated that these birds were like the pheasant hen of England in size but were rough footed and had “stareing” feathers about the neck. The birds, according to Morton, were so common that they seldom wasted a shot upon them. The writings of many others who followed indicated that the birds were distributed along the Atlantic seaboard from Maine and Massachusetts southward to Virginia and possibly the Carolinas. They were by no means evenly distributed over this region, but were restricted to certain areas whose features and productions were suitable for their existence. There were large heavily timbered areas that probably were never visited by the heath hen. In favorable localities, such as the brushy plains of eastern Massachusetts, they were abundant. Thomas Nuttall (1832) wrote as follows: “According to information I have received from Governor Winthrop, they were so common on the ancient brushy site of Boston, that laboring people or servants stipulated with their employers not to have Heath Hen brought to table oftener than a few times a week.” No published statement has ever been found that more impressively reveals to us the abundance of these birds in early colonial times. It was chiefly on the sandy scruboak plains of Massachusetts, Connecticut, Long Island, New York, New Jersey, and Pennsylvania that they existed in large numbers when the white man first came to America. The birds served as a valuable source of food, and because they were easily tricked and killed they were exterminated at an early date in the more accessible areas, and soon after 1840 were entirely gone from the mainland of Massachusetts and the State of Connecticut. The birds persisted for a longer time on Long Island, and a few continued to battle for existence on the plains of New Jersey and favorable places among the pines and scrub oaks of the Pocono Mountains in Northampton County, Pa. Since 1870 the surviving members of this interesting race have been restricted to Marthas Vineyard Island, off the southeastern coast of Massachusetts.
Because of conflicting reports and uncertain statements we can not be positive whether the heath hen was native to Marthas Vineyard or was introduced there from the mainland by man. In either case it is truly remarkable that the heath hen, after being so greatly depleted in numbers, has persisted for more than half a century in this very restricted area where excessive interbreeding has occurred and where the birds have been subjected to all the vicissitudes of diseases, enemies, and other adverse conditions.
In 1890 William Brewster (1890) made a careful census and at that time estimated that there were 200 birds on the entire island. Kentwood (1896) stated that there were less than 100 birds in 1896. By the beginning of the twentieth century the birds had reached a very low ebb in their existence. The year 1908 witnessed one of the most notable steps taken in the history of the heath hen in an effort to preserve it from extinction, in the establishment of a reservation in the midst of the breeding range where the birds could be protected from poachers and predators by competent wardens. Six hundred acres were purchased by private subscription, and an additional tract of 1,000 acres was leased by the Commonwealth for a reservation, which was systematically improved to make it attractive for the birds. There is no doubt that the prolongation of the life of the heath hen on Marthas Vineyard Island has been due to the interest taken in it by the State of Massachusetts, conservation organizations, bird clubs, sportsmen, and bird lovers. The State Department of Conservation expended $70,000, and thousands more were contributed by individuals, in the unprecedented efforts to prevent the bird from becoming extinct. Many attempts were made when the birds were abundant to transplant them to other favorable places on the mainland and to other islands such as Long Island, one of their former strongholds. Furthermore, the most experienced sportsmen and game breeders were unable to breed the birds in captivity, a fact indicating that the heath hen was very sensitive to radical changes in its enviromnent and that it would not yield to such methods of conservation. All the many experiments of introducing the western prairie chicken to the East have likewise proved unsuccessful.
When the reservation was established in 1908 there were only about 50 heath hens, but as a direct result of the efficient protection the birds increased very rapidly and by 1915: 16 they were to be found in all parts of Marthas Vineyard with the exception of Gay Head, the extreme western end of the island. It was then possible to flush a flock of 300 or more birds almost any day from the corn and clover plots planted on the reservation for the birds. An estimate made by William Day, then superintendent of the reservation, indicates there were probably 2,000 birds on the island. This was a great triumph for those who had encouraged and fostered the reservation, but unfortunately success was not long-lived.
In spite of the unusual precautions taken to prevent the spread of fire, a terrific conflagration broke out during a gale on May 12, 1916, which swept the greater part of the interior of the island, destroying brooding birds and their nests and eggs, as well as the food and cover of the birds on more than 20 square miles, right in the heart of the breeding area of the heath hen. This fire undid in a few hours the accomplishment of many years of work. A hard winter followed the fire, and in the midst of this came an unprecedented flight of goshawks, which further decimated the number of birds. The net result of this catastrophe was an amazing decrease m the number of heath hens, which according to official estimates was reduced to less than 150 birds, most of which were males.
There was a slight rally in numbers during the following few years, but the birds were too far gone to overcome the surmounting uncontrollable conditions of extensive interbreeding, declining sexual vigor, the condition of excess males, and, worst of all, disease. In 1920 many birds were found dead or in a weak and helpless condition, indicating that disease was then exacting its toll. The heath hen is very susceptible to poultry diseases, and when domestic turkeys were introduced to the island in large numbers the dreaded disease blackhead caine with them. The turkeys and heath hens fed on the same fields, and thus the disease was readily transmitted through droppings to the native birds.
The heath hen continued to decrease in numbers, and by 1925 it was apparent that they had reached their lowest ebb in history. The Federation of the Bird Clubs of New England then came to the front and offered to raise $2,000 annually to support additional warden service. In spite of this splendid cooperation the birds, after two years of effort on the part of all concerned, continued to decrease. The 1927 spring census revealed but 13 birds, only 2 of which were females. During the fall of 1928 only two birds were seen and after December 8 but one was reported. This bird was photographed from a blind on April 2, 1929, at the farm of James Green located on the State highway between Edgartown and West Tisbury. At that time it was the common expectation that the bird would step out of existence before the end of another year. (See p1. 1, frontispiece.) It was seen regularly until May 11, 1929, but after that date it disappeared among the scrub oaks to live a life of seclusion, as was customary for the heath hen to do in the past, during the summer months. After the molting season it again appeared at the Green farm to announce to the world that it was still alive. It was seen at irregular intervals during the winter, and after the first warm days of March it appeared daily at the traditional booming field at the Green farm. The bird was studied and photographs were taken again at the time of the annual census in March-April, 1930. The lone bird continued to appear at the Green farm during April and May, where it was observed by many ornithologists and bird lovers who journeyed to the island to get a glimpse of the famous last bird. The bird again disappeared during the summer, and no reports were received until it almost met a tragic death on September 15, when it was nearly rumi over by an automobile traveling one of the little-used roads leading across the scrub-oak plains. In October it resumed its daily visits to the open field on the Green farm and at the time of this writing (November 15, 1930) was still alive. It is the first time in the history of ornithology that a bird has been studied in its normal environment down to the very last individual. how long this bird will live no one can safely predict; its going is inevitable, and the death of this individual will mean the death of its race, and then another bird will have taken its place among the endless array of extinct forms. Ornithologists, bird lovers, and sportsmen the world over, however, will have the satisfaction of knowing that all that could be done has been done to save this bird from extinction. The State department has assured us that the last bird will be allowed to live, and when death comes, whether it is due to old age, disease, or violence, we shall know that the life of the last heath hen was not wilfully snuffed out by man. [During the fall of 1931, this lone survivor disappeared.]
Courtship: There was no part of the behavior of the heath hen more unique, more interesting, and more specialized than the extraordinary performances during the courtship season. I vividly remember the thrill of hearing and seeing the heath hen’s boom for the first time on Marthas Vineyard Island during April, 1923. At that time the birds came regularly to a definite part of the meadow west of the reservation house. A wooden blind, 4 by 6 by 6 feet, had been erected several years before for the convenience of the large numbers of ornithologists who journeyed to the island each year to get a glimpse of the heath hen. The blind had become a part of the environment of the drumming field around which the birds came to enact their fantastic dances without any fear of being harmed. At times one or two of the birds would even alight on the fiat top of the structure, offering unexcelled opportunities for study of the intimate details of their behavior.
The following observations, as recorded in my notebook for April 11, 1923, are typical of the many mornings spent inside the blind: “I left the reservation house at 3.30 a. m. It was very dark and only the faint light of the stars illuminated the way. The cold, enervating air quickened my step, and as I walked along, the frosted grass crunched under my feet with a metallic resonance. At this early hour all was quiet so far as voices of birds were concerned. I entered the blind, closed its door on creaky hinges, and prepared myself to wait patiently for the first note of the heath hen. A slight fog rolled in from the sea and for a time hid the stars. At 3.55 a. m. with the first dim light of dawn I heard the clear whistled notes of a bobwhite perched somewhere among the scrub oaks. At 4.05 the first robin was heard chirping, and five minutes later a vesper sparrow was singing its awakening song. In a short while a host of other birds were adding their notes to the morning chorus. At 4.21 the first “toot” of the heath hen was heard, a note that has often been mistaken for a muffled blast of a tug boat or a fog horn. Though I was well prepared for this deception, I must admit that I did not at first associate this curious note with the heath hen, for the light was yet dim and the fog obscured the view of the bird. At 4.27 a heath hen appeared from the scrub oaks on the south side of the meadow at a point relatively near the blind. The calls of this individual at once stimulated the birds on the western end of the field to greater activity in tooting. After 20 minutes a second and then a third bird came out of the scrub oaks on the south, and for a time all were busily engaged in feeding. One of the males flew to the roof of the blind, where he commanded a splendid view of the field and his companions. Later it was found that the resonant wooden roof proved to be an admirable place to conduct their stamping and courtship performances. At 4.45 the birds without any warning interrupted their feeding and began “booming or tooting” at a point only a few yards from the blind. The male on the roof joined his fellows on the ground. At this close range the call resembled wklioo: doo-.doooA. The note varied somewhat in Subsequent renditions and was variously interpreted as wkoo.-oodul: doo: o–o: o: o, whoodle: dook, or whoo: dook: dooh. The sound was accented on the second syllable or the first part of the second and then gradually diminished in intensity. It required from iy2 to 3 seconds to render the different versions given above. The number of calls a minute varicd greatly, according to weather conditions, temperature, time of day, and the season. The booming was interspersed with henlike calls resembling cac, cac, cac, or oc, oc, OC, CC, goc, goc, goc, god, occasionally ending with a queer call that sounded like auk: ae: e: e: e–e–ek. The males frequently leaped into the air to a height of 3 or 4 feet and so doing uttered a piercing iuliing wrrrrm~rb, followed by a curious indescribable laughterlike sound. In this wild demonstration the bird completely reversed its orientation in the air and landed on the ground, usually facing in the opposite direction. This leaping and screaming seemed to be augmented by similar performances of the birds on the other side of the field, and it was an evident challenge to their fellow antagonists. “At 5 a. m. the sun pierced the screen of fog and appeared like a giant fiery ball above the eastern horizon. The morning chorus of birds then rapidly diminished in volume, but the heath hen now prepared for real action. One male from each group ran rapidly toward each other in a defiant warlike attitude. When near together they hesitated, lowered and waved their heads, leaped at each other, and struck their wings vigorously as they lunged forward. A few feathers flew, but no real harm was done, and they settled back in a comfortable position and occasionally uttered a long-drawn-out but shrill cry, which fluctuated greatly in tone and intensity. One bird arose after a few minutes, circled, paced a few steps, and went through his repertoire of toots and calls without any interference from his antagonist. Later one ambitious male insisted on chasing one opponent after another, following after them rapidly on foot until they took wing. He flew after them for a distance of 30 to 50 yards, then returned to repeat the performance with another weaker member of the group of a dozen birds. It reminded one of boys at play after being pent up in school all day. These thrilling spectacles continued until 6.50 a. in., when with one common impulse all the birds left the field to the seclusion of the scrub oaks to remain quiet until the afternoon, when they again appeared on the drumming field during the few hours preceding sunset. But when the last glow of twilight faded into darkness the fantastic dance ceased to be resumed at the coming of dawn the next day.”
The first “tooting” calls of the year were usually heard the last week of February or the first week of March, the date varying from year to year and depending largely on the nature of the weather. In 1927 a series of warm days started the birds booming as early as February 12. Though an early beginning was sometimes made it was not until the latter part of April or the first week of May that the courtship reached its maximum intensity. It then gradually diminished, and by the end of May the performance was generally over~ but a few more persistent males often continued a few weeks longer. In 1923 the last “boom” for the year was recorded on June 11; and in 1920 a small group of males was still performing as late as June 20. After the month of June the birds ceased their nuptial displays until the mating season of the next year.
The following details of the courtship performance were obtained by repeated observations from blinds of the birds at close range and are supplemented by a study of captive birds and detailed laboratory dissections: The tooting is usually prefaced by a short run, followed by a very rapid stamping of the feet, a part of the performance that is not readily detected unless the observer is very near to the birds. The stamping is vigorous enough, however, to be distinctly heard at a distance of 25 or 30 feet, and certain males, which did their stamping on the resonant roof of the blind, produced a noise second only to the tooting that followed. In preparation for tooting the neck is outstretehed forward; the pinnae (neck tufts) are usually directed upward or forward; the primaries are spread and held firmly against the sides of the body and legs; and the tail is thrown upright at right angles to the axis of the body, thus displaying t.he white under tail coverts when viewed from the rear During this procedure the whole musculature of the body seems to be in a strained state of contraction, as if it required great effort on ïthe part of the bird. As the inflation of the orange-colored sacs (it is really one large sac with two lateral areas devoid of feathers) begins, the tooting sound is heard. Sometimes there is a slight inflation before any sound is given. The inflation seems uniform and does not fluctuate with the inflections and accents of the tooting call. At the end of the tooting the sacs collapse suddenly by the release of air through the nares or more rarely through the opened mandibles. The sacs do not produce the notes, as was thought by some of the earlier ornithologists, but have much to do with modifying the sounds produced by the syrinx (the vocal mechanism at the junction of the bronchial tubes). The sounds are produced by the air forced from the lungs, which vibrates specialized membranes of the syrinx under control of a complex set of muscles. The sound waves then issue through the trachea and glottis to the pharynx. In the production of such notes as the ordinary cackle the mandibles are opened and the air accompanied by the sound waves issues out of the mouth. In the tooting performance the mandibles are tightly closed, the throat patch is elevated, and the tongue is forced against the roof of the mouth (palate) by the mylohyoides muscles, which close off the exit through the internal nares. The tongue is bent in such a way that it causes the glottis at the base of the tongue to open directly in front of the esophagus. The air now coming from the respiratory system is forced to fill the modified anterior end of the esophagus, or gullet, which becomes distended like a balloon.1 While the air sac is filling, the sound waves produced by the syrinx beat against these tense drumlike membranes, which serve as resonators for the sounds and give them their great carrying power. The ordinary cackles and screams of these birds seem louder than the tooting or booming calls when one is near to the birds, but at a distance of 200 yards or more you can scarcely hear these calls, whereas the booming carries for long distances, often 2 miles or more under favorable conditions.
The female’s part in the courtship is a passive one. She minds her own business, and I have never heard her utter any calls or notes or show any concern in response to the ardent attentions of the males. When a male or pair of males came strutting and circling about a female she kept on with her feeding. If the males came too near she merely stepped to one side and continued with the serious business of procuring food.
‘A number of detailed experiments were performed with both dead and living birds, which clearly demonstrate tbe nature of the vocal mechanism aa deecribed abova See Gross (1928).
The females gave frequent calls and notes when attending their young. If the mother bird was suddenly surprised she gave a characteristic sharp call signal for her young to scatter and hide. If the members of the brood were very young and unable to fly, she feigned a wounded bird and cried out as if in great distress as she fluttered along the ground. If the young were older she usually sailed out over the scrub oaks and uttered a loud cackling call, which apparently was also for the purpose of attracting attention away from the young.
Nesting: The nest of the heath hen was built upon the ground and was usually composed of leaves, grasses, and twigs already in place, to which were added materials found near the nesting site. The nests were concealed by the low dense vegetation of the scruboak plains. Indeed they were so well hidden from view and the eggs so well covered when the bird was away that few of the nests were ever found, in spite of the great efforts various observers have expended to locate them.
William Brewster (1890) states: “Only one person of the many I have questioned on the subject has ever found a heath hen’s nest. It was in oak woods among sprouts at the base of a large stump and contained either 12 or 13 eggs.” There was a set of six eggs in the Brewster collection that were found in a nest in the woods near Gay Head on July 24, 1885. This set was described and one egg was figured by Capen (1886). One of the eggs given to the United States National Museum is figured by Bendire (1892, p1. 3, fig. 2). Bendire stated that the six eggs referred to above were the only eggs in any collection known to him.
In 1906, E. B. McCarta found a nest and nine eggs in a low but dense growth of scrub oak near the central part of Marthas Vineyard Island. Dr. George W. Field photographed the nest on June 2, and two days later the eggs were placed under a bantam hen. One of the eggs hatched on June 20, but unfortunately the chick was killed by the hen, and the other eggs failed to hatch. This set of eggs with the chick is now a part of a display group in the American Museum of Natural History, New York. On June 5, 1912, Deputy Warden Leonard, after a most prolonged and diligent search, found a nest and four eggs covered with leaves in a slight hollow surrounded by a dense mass of sweet ferns growing among the scrub oaks. The oaks in the vicinity were 2 or 3 feet in height. When the nest was visited on June 12 the bird was incubating. On June 21, Doctor Field was able to determine that there were eight eggs, and a week later he took an excellent series of photographs of the bird on the nest. The bird sat so closely on the eggs that it was dislodged only by active effort. Deputy Leonard had no difficulty in approach ing the bird, and she fought the approach of the hand in the same manner as would a sitting hen, ruffling her feathers, opening her beak, and striking viciously. The incubation period of the heath hen, according to Doctor Field, is 24 days.
Eggs: In addition to the eggs mentioned above in connection with the account of the nests, there are the following: An egg in the Brewer collection of the Museum of Comparative ZoSlogy marked Tympanucku8 cupido, Holmes Hole, Mass. (Holmes Hole is the old name for Vineyard Haven). There is an egg in the John E. Thayer Museum and another in the collection of the Boston Society of Natural History. The latter was picked up on the plains of Marthas Vineyard after the destructive fire of 1916. Bendire (1892) describes the eggs as “creamy-buff in color with a slight greenish tint, ovate in form and unspotted.” They are regularly oval in form, all specimens being quite uniform in this respect. The color is yellowish green of a peculiar shade. I have compared the colors of this set of eggs with Ridgway (1912) and find all of them to be a “deep olive buff.” One egg has a small spot of drab. All other heath-hen eggs that I have examined are of this same deep olivebuff color.
The measurement of the five eggs in the Brewster collection are as follows, 43.5 by 32.5, 43.5 by 32.1, 43.8 by 32.5, 43.9 by 32.4, and 46.2 by 32.9 millimeters, and the sixth egg, now in the United States National Museum, measures, according to Bendire, 44 by 33 millimeters. The egg in the Brewer collection collected at Holmes Hole measures 44.2 by 32.6, and the egg at the Boston Society of Natural History collected at Marthas Vineyard in 1916 measures only 40.3 by 30.4, the smallest egg of the species I have examined.
Young: The first young of the heath hen for the season usually made their appearance during June. The earliest record is of a brood of 8 or 10 young seen near Edgartown on June 14, 1913, by Dr. Charles W. Townsend. During the season of 1913, nine broods, with an average of four chicks to the brood, were seen. On July 15, 1914, a dead chick was found that was estimated to be about five days old. In 1915, 14 broods were reported. The first brood, seen on June 19, consisted of six chicks about five days old. Allan Keniston saw seven broods with an average of five chicks each during the summer of 1918, and in 1919 he reported broods on June 29, July 1, and July 11, the members of which were able to fly well at that time. For 1920 the following were noted: June 20, 10 young; June 24, 6 young; July 4, 2 small broods, the numbers were not recorded; July 9, 5 or 6 young. During the summer of 1921 a brood of sr7en was seen on July 3, and a few days later a brood of eight was recorded. On July 31 a brood of six, about two-third~’ g: own, was seen. In 1922, between June 15 and June 30, there were five broods reported containing four to eight chicks; the exact dates were not recorded. In 1923 the writer made a continuous search for nests and young throughout the summer, but the birds by that time were so greatly reduced in numbers that only one brood of two chicks was seen, and that on July 3 during a downpour of rain, as we suddenly surprised the brooding mother in the middle of one of the little-used crossroads. Since 1923 there have been no authentic records of broods of young birds, and I very much doubt if any young have been reared since 1925.
Plumages: The following description is based on a downy young about four days old: Underparts “cream-buff,” the throat and middle of the belly approaching “colonial buff.” Sides of the head “Marguerite yellow” with three small black spots back of the eye. Upper parts “tawny-olive” or “Isabella color,” the region of the rump “snuff brown” and “russet,” variously marked with black. There is a conspicuous mark on the forehead. The remiges and coverts marked with various tones of brownish drab and black, the feathers tipped with dingy white. The measurements of this specimen are as follows: Bill, 8; tarsus plus third tce, 39.8; wing, 42.5; length, 85; third toe, 23 millimeters. The natal down of a 2-day-old specimen hatched in captivity by Dr. John C. Phillips is similar to the above description but with the following differences: The underparts brighter yellow, the throat and sides of the head “amber yellow.” The bright yellows fade rapidly when exposed to air and light, and in chicks two weeks old the bright yellows of the underparts are faded to. a uniform “cream-buff.” The measurements of the 2day-old chick are: Bill, 7.5; tarsus plus third toe, 37; wing, 28; length, 79; third toe, 19 millimeters.
I have been unable to obtain a specimen of the heath hen in the completed juvenal plumage, but it is reasonable to infer that the sequence of the molts and plumages are similar to those described for the prairie chicken.
The first winter plumage of the heath hen is acquired by a complete postjuvenal molt except on the two distal primaries. The first winter plumage is similar to the adult plumage, but the younger birds are readily distinguished from the adults by their smaller size, by the more rufescent color of the upper parts, and by the coloration of the throat, which is “cinnamon-buff” in contrast to the “warm buff ~ or “cream color~~ of the adults and the white throat of the juvenals. The first plumages of the heath hen and the prairie chicken are so nearly alike that one can not readily distinguish them from each other. This ontogenetic resemblance indicates the close relationship of the two races.
The first nuptial plumage is acquired by a partial prenuptial molt, which does not involve the wings and tail and body plumage but is restricted to the head region. The nuptial plumage was usually completed by April. The second or adult winter plumage is acquired by a complete postnuptial molt during August and September. Specimens collected in October and November had the adult winter plumage completed. From the adult winter plumage onward the plumages are repetitions of the nuptial and winter phases.
Food: The heath hen in its feeding habits was similar to other gallinaceous birds, such as the ruffed grouse, in being very adaptable to the changing food supply throughout the seasons of the year. It was not dependent on any particular item but subsisted on what was most abundant and most easily procured.
During the spring months the heath hen congregated in the open fields and meadows of the farms to feed upon the tender shoots of grasses, sorrel, and other plants, but when these became hardened and less palatable with the approach of summer the birds changed their diet to fruits and insects. In fall, berries and insects, such as grasshoppers, were freely eaten, and in the winter months, acorns, seeds, and certain berries found throughout the range of the heath hen on Marthas Vineyard provided the birds with a livelihood. Comparatively little snow falls on the island, and hence it was an exceptional winter when the birds were not able to obtain sufficient food from native plants. Even at times when the ground was covered with snow the scrub oaks held out an exhaustless supply of food in the form of acorns.
Our chief knowledge of the food of the heath hen is based upon the meager notes of food included in the data with birds collected and preserved as skins in various museums, upon field notes, and upon studies of birds kept in captivity. The following comprise the principal foods:
The crops of three heath hens collected by C. E. Hoyle on January 10, 1891, and two on December 28, 1895, contained bayberries (Myrica carolinensis). A specimen of a heath hen killed by a snowy owl was reported by Allan Keniston to have had its crop gorged with bayberries. Birds kept in captivity at the reservation in 1915 ate very freely of bayberries. During the winter of 1923: 24 a flock of 15 birds frequented a large bayberry thicket near the south shore where they subsisted chiefly on these berries.
The bearberry (Arctoataphylos uva-ur&i), sometimes wrongly named mountain cranberry or cranberry, is extremely abundant throughout the central portion of the island and was freely eaten by the heath hen during the winter months when the trailing plants were not covered with snow.
Audubon (1840) stated that the barberry (Berberi8 vulgaris) was the chief food of the heath hen. This is also included in the lists of the food of the heath hen by other earlier writers, but there is no evidence that the birds in recent years depended very largely on the barberry for a source of their food.
The fleshy wild-rose berries, more frequently called rose apples or rose hips, were eaten by a bird collected March ‘1, 1896.
The birds fed very freely on wild strawberries, which were abundant in the meadows and open areas of the reservation. They were also fond of the cultivated varieties, as evidenced by their frequent depredations in gardens grown near the haunts of the heath hen.
The partridgeberry (Mitchella repen~) was so frequently eaten by the heath hen that the earlier settlers called this berry the heathhen plum. Not only the berries but the leaves of this plant were often eaten by the birds during the early fall and winter months.
The dryland blueberry ( Va in~iurn vacillaina), the low-bush blueberry (17. penneylvanicum), and the black huckleberry (Gaylueeacia baccata) were eaten during the berry season. On August 24, 1913, William Day saw a flock of 51 heath hens feasting on blueberries, and he also states that captive birds ate freely of blueberries provided for them.
The acorns of the scrub oak (Quercus ilicifolia) have been called the “bread” of the heath hen. No natural food is more abundant on Marthas Vineyard, and no food is more dependable during the winter months. The scrub-oak acorns are small and were swallowed whole by the birds. The scrub oaks provide one of the reasons why these birds have persisted in the scrubby plains.
Leaves were found in the crops of many specimens examined by Mr. Hoyle. In the spring months I found that the birds showed a great preference for the leaves of sheep sorrel (Rumero acetoedla) over other plants such as clover and tender blades of grasses that were equally abundant in the same field. The distribution of the birds on the field usually corresponded roughly with the distribution of this plant. One male specimen trapped on April 5, 1924, had its crop gorged with leaves of sorrel. There were in this specimen, 1,846 leaves and parts weighing 32.2 grams. Another bird trapped on May 20, 1924, had its crop completely filled with 38.6 grains of seeds of the sheep sorrel. Although other leaves, such as those of clover, alfalfa, and other herbaceous plants, were eaten, the birds exhibited a decided preference for the leaves of the sorrel.
Buds, including those of the scrub pine (Pin~us virginiana), were eaten by certain birds during the winter months.
The heath hen was especially fond of the grains of cultivated crops, such as corn, buckwheat, millet, and sunflowers, and in late years these were planted on the reservation, especially for the use of the heath hen.
The animal food of the heath hen consisted primarily of insects, chiefly grasshoppers, which were sometimes excessively abundant on the island late in summer and in autumn.
There are very few data concerning the food of the young, but, judged from the, food of the young prairie chickens, it probably consisted mainly of insects, especially in the case of the younger birds. The crop contents of one 5-day-old heath-hen chick accidentally killed on July 15, 1914, contained more than 80 insects representing 10 species. The vegetable matter was merely incidental and negligible in this specimen.
Garnet: The heath hen even when in its prime was never given a high place among game birds by the sportsmen. It was easily shot, because of its direct and laborious flight, and the habit of massing in flocks in the open fields made it too easy a victim for the pot hunter and the market gunner. The ease with which the heath hen was tricked and killed readily accounts for the rapidity of its early disappearance from the mainland, whereas the clever ruffed grouse of the woodlands still holds its own and is ever ready to challenge the wits of the most skilled sportsman.
The attitude of the sportsman in the past toward the pinnated grouse is well illustrated by the following excerpt from an article by Elisha J. Lewis in his book The American Sportsman, 1885.
So numerous were they a short time since in the barrens of Kentucky, and so contemptible were they as game birds, that few huntsmen would deign to waste powder and shot on them. In fact they were held in pretty much the same estimation, or, rather abhorrence, that the crows are now, as they perpetrated quite as much mischief upon the tender buds of the orchards, as well as the grain of the fields, and were so destructive to the crops, that It was absolutely necessary for the farmers to employ their young negroes to drive them away by shooting of gnus and springing loud rattles all around the plantation from morning till night. As for eating them, such a thing was hardly dreamed of, the negroes themselves preferring the coarsest food to this now much admired bird.
It is apparent that the heath hen was not considered an ideal bird from the point of view of the sportsman, and our efforts to save the heath hen were not made on the plea of its economic importance as & game bird. It is interesting to note, however, that it was the sportsman who took the initiative and who provided a large part of the funds to assist the State in the vain attempt to preserve this interesting race of birds.
Enemies: Man directly or indirectly is in part responsible for the disappearance of the heath hen from most of its former range. Comparatively soon after the coming of the white man, it was driven from one locality to the next, until it was forced to entrench on the scrub-oak plains of Marthas Vineyard Island. It is a striking example of a bird that has not been able to adapt itself to the changing conditions brought about by civilization.
Audubon (1840) wrote: “We frequently meet with the remains of such [heath hen] as has been destroyed by the domestic cat which prowls in the woods in a wild state.” What was true in Audubon’s day was true in the more recent years of the heath hen, when cats ranked high among the enemies of the bird. In addition to the cats reared on the island, large numbers were introduced by people who dumped them in the interior of the island when they left their summer homes in autumn. A large part of the effort of the State Department and the special wardens in the control of vermin was directed toward the semiwild house cat.
Large numbers of hawks are attracted to the island because of the abundance of mice and shrews, which live among the scrub oaks. Unfortunately, many of the hawks, for example the marsh hawk, which have a good reputation elsewhere, are frequently tempted, on Marthas Vineyard, to prey upon birds; and when the heath hen was common, these birds were also numbered among their victims. The goshawks, notorious for their killing of game birds, played their part in the history of the heath hen. The most notable instance of their wholesale depredations was in the winter of 1910: 17, following the destructive fire that swept over the island during the preceding spring. Other hawks, such as the red-shouldered, the rough-legged, the pigeon hawk, and others, as well as the different species of owls, were killed on sight by the wardens in charge of heath-hen protection. It is probable that the wholesale killing of hawks and owls so upset the balance of nature that it acted as a boomerang to the heath hen.
Disease was one of the most important factors in the recent decline of the heath hen and was one that man was unable to control. Blackhead is a disease common to poultry, but, so far as we know at present, it is unusual in birds living in a free, wild state. The heath hen, however, had the peculiar habit of congregating in the open fields near farmhouses where poultry was kept. In most instances, chickens and turkeys had access to the fields visited by the heath hen, and thus the dreaded disease was readily transmitted to the native birds. Blackhead was found in the adult heath hen, and this is presumptive evidence that the disease was very destructive to the young.
Internal and external parasites were found on the few heath hens examined, but these were all of minor importance as compared to the disease blackhead.
In the recent history of the heath hen it was well known that there was a great excess of male birds. This abnormal ratio may have been brought about in part by some hereditary influence, but it is certain that this condition was aggravated by the fires that ravaged the island during the breeding season. At such times the females were destroyed on the nests, whereas the males escaped the conflagration. Furthermore, a female with young was subject to more danger of being killed than the male, which never cared for the young.
Other factors that played their part were the excessive interbreeding, which was destined to occur after the heath hen was restricted in range and to exceedingly small numbers of individuals. It was also found upon examination of dissected specimens that many of the birds were sterile.