The Chestnut-collared Longspur nests and winters in native prairies, and loss of that type of habitat has affected the range and population size of the species. Territorial during the breeding season, Chestnut-collared Longspurs are gregarious during winter, when large flocks can sometimes be seen moving with an undulating flight pattern by each bird.
Highly variable spring and summer weather in the northern plains can cause mortality of nestlings by either cold or heat, though predation remains the largest source of nest failure. Grazed native pastures are preferred over planted domesticated pastures.
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Description of the Chestnut-collared Longspur
The Chestnut-collared Longspur varies in plumage in the course of a year, but has a white tail with a black triangle centered on the end and a pointed, seed-eating bill. Black belly, rufous nape, buffy throat. Length: 6 in. Wingspan: 11 in.
Female is drab brown with dark “ear patch, ” indistinct streaking on flanks.
Seasonal change in appearance
Winter males are substantially duller, resembling females, with just a trace of the reddish collar.
Juveniles resemble females.
Prairies and plains.
Seeds and insects.
Forages on the ground.
Breeds in the north-central plains of the U.S. and Canada and winters in the south-central plains of the U.S. and Mexico. Populations declining.
Bent Life History
Visit the Bent Life History for extensive additional information on the Chestnut-collared Longspur.
Chestnut-collared Longspurs prefer the short vegetation provided in grazed or mowed areas.
Males perform aerial song displays during the breeding season.
A fast warble, a two-note “kittle”, and rattles or buzzes are given
The black at the end of the McCown’s Longspur’s tail forms an inverted “T” rather than a triangle.
Lapland Longspurs have mostly black tails.
The nest is a grass cup on the ground near a plant or other object.
Color: White with darker markings.
Incubation and fledging:
– Young hatch at 10-13 days.
– Young fledge (leave the nest) in 10 days after hatching but remain with the adults for some time.
Bent Life History of the Chestnut-collared Longspur
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 Chestnut-collared Longspur – the common name and sub-species reflect the nomenclature in use at the time the description was written.
CALCARIUS ORNATUS (Townsend)
Contributed by GEORGE M. FAIRFIELD
When he investigated the birds of the Saskatchewan prairies in 1905, Mr. Bent (1908) wrote of the chestnut-collared longspur: “This and the following species [McCown’s longspurj were a constant source of enjoyment and interest; we never ceased to admire their beautiful plumage and their delightful little flight songs, during our long drives across the grassy plains.”
Indeed, the male chestnut-collar, with his black underparts, white winglinings and tail, and rich chestnut nape, adds a pleasant note of contrast to the rather monotonous aspect of the northern prairies. Where the grass is sparse and preferably less than 8 inches high, we see dotted here and there across the plain the black breasts of birds perched on low weeds guarding their territories. As we approach they take off and circle around with undulating flight, giving their short lively song and their high clear alarm call.
When the northern prairies became the great wheat-producing region of the continent, the amount of grassland available for the chestnut-collar was redu ced proportionately. The McCown’s longspur and the horned lark, adapted to sparser, more open ground than the chestnut-collar, were able to continue nesting in the summerfallow ploughed fields. The chestnut-collar, needing elevated song perches and a little cover for its nest, found the plowed lands too open. As the fields of growing wheat were too dense and confining for such an open-country bird, all that remained for its use were uncultivated pastures and wastelands. In heavily settled regions today the only suitable habitat for the chestnut-collar is on the outskirts of towns and cities where golf courses, airports, and idle lands provide the needed degree of cover.
T. S. Roberts (1932) thus describes the effect on the chestnut-collar of the agricultural altering of the virgin prairie in Minnesota: “Formerly an abundant summer resident throughout the prairie region of Minnesota * * ~’ï It has entirely disappeared in recent years from this range, except for a few small isolated colonies on the first sand ridge along the Red River Valley in western Pennington and eastern Polk counties * * * and in central Norman County.”
Frank Roy (1958) of Saskatoon, Saskatchewan, reports: “The Chestnut-collared longspur was an abundant bird in the Lucky Lake-Birsay region from the year 1937 to 1944. As many as 15 pairs used to nest in a fifty-acre pasture on our farm. By 1945, not more than seven or eight pairs nested in the same area. In 1946 five pairs remained. In 1947 not a single bird nested in the pasture (which, by the way, has been reduced to 20 acres in extent). Since that date, the Chestnut-collared Longspur has become progressively more scarce, even in the extensive tracts of pasture west and south of Lucky Lake and Beechy.”
Nonetheless, large areas of suitable grassland still exist on the northern prairies, and where conditions are favorable the breeding population of chestnut-collars can be quite heavy. Elmer T. Fox (in Belcher, 1961) found 11 nests on about 20 acres of pasture near Regina, and we found 19 territories (16 nests located) on 50 acres of waste land at Moose Jaw, Saskatchewan. As several of the early writers mention it as “abundant locally,” perhaps the species has always tended to nest in loose “colonies” in concentrated groups of territories. E. T. Seton (1886) notes that in Manitoba it was “Local in distribution, many pairs sometimes affecting a limited area of dry prairie, while again for miles no more of the species are to be seen.”
Much of the following information has been extracted from an unfinished study of the chestnut-collar in Saskatchewan. D. J. T. Russell cooperated with me on the breeding biology work, and several other friends have given help and advice.
Spring: Very little information is available on the northward migration of the chestnut-collared longspur. Comparatively few of the species have been banded (1,067 to 1962, with only 3 returns to the place of banding and no recoveries reported). The summer and winter ranges meet in northeastern Colorado and Nebraska, and much of their migration seems to consist merely of a northward drifting, in company with the flocks of other longspurs and of horned larks, from their winter home on the deserts and plateaus of northern Mexico and the plains of the southwestern United States to the northern prairies.
Several writers have noted their habit of stopping for several days at favorable feeding and watering places. W. H. Osgood (1903) says of their migration through Cochise County, Arizona: “In February and March the chestnut-collared longspur was exceedingly abundant. They were seen flying over at all times and at nightfall clouds of them would sweep over the house and on down to the grass at the edge of the alkali lake, whence they straggled out at daybreak.”
The birds pass northward through western Texas and Oklahoma, New Mexico, and eastern Arizona, across Colorado and Kansas, and arrive on their breeding grounds about the second week of April. R. D. Harris (1944) describes their arrival at Winnipeg, Manitoba: “On the first day of migration, no more than a single individual was usually noted, but afterward 10 or 12 were counted in a day. They arrived principally during the forenoon, either singly or in groups of two or three.”
Territory: We found that the chestnut-collared longspur usually defends a nesting territory of between 1 and 2 acres of prairie, though two of our pairs occupied territories of almost 10 acres of rather unsatisfactory habitat at the margin of the nesting area. As with other territorial species, the more the pairs are crowded together and the smaller the individual territories, the more defensive behavior is to be seen.
An important aspect of territorial defense consists of advertising by singing from conspicuous perches. Rival males spent many hours singing at each other from weed clumps, small shrubs, boulders, fences, or farm refuse. Our wire nest markers were favored perches, and one longspur even sang from the top of the car when it was parked near the nest. The birds rarely sang from the transmission line that crossed our study area; possibly its 20-foot poles were too high to suit their purposes. The birds stand out so strikingly against the pale prairie grass when singing that I believe the black breast is almost as important as the song in advertising territorial occupancy.
L. J. Moriarty writes in a letter of his observations of the species in eastern South Dakota:
The birds nearly always nest on the higher areas in our rolling short-grass lands. Seldom do I find them in hay land as they prefer open places with poor soils and short-spaced vegetation. Each male appears to have what I call his “singing rock” upon which he sits, flies up from to sing or to defend his territory and returns to constantly. These rocks are usually the largest ones in each pair’s territory, and are soon marked by a ring of droppings around the highest peak. Occasionally when no rock is available, a weed or a post will be substituted. The nest is usually within 25 feet of the singing rock. Each pair’s territory appears to be about 100 yards across. Of the many nests I have found, none have been closer together than 100 yards.
In Saskatchewan we observed chases of two types. The commoner occurs when a neighboring bird flies over an occupied territory. The male, and sometimes the female as well, take wing and follow him at a distance of a few feet. The chase usually continues over at least one more territory, from which another male will rise to join the chase. It is common to see three to five longspurs circling round and round after one another in undulating flight high above the fields, then dropping back to their own territories.
Occasionally a much more violent chase takes place, apparently when a male bird discovers a trespasser on the ground within his territory. He flies at the offending bird and chases it in a fast, erratic course low over the ground until he has driven it well away from his boundaries.
Fights occur frequently along territorial boundaries. Two males, each probably on his own territory, may be foraging only a few feet apart. Suddenly one of them, possibly considering the other to be trespassing, runs at him with lowered head. Facing each other, the two join battle with fluttering wings. They may rise together to a height of 4 to 15 feet before dropping separately to the ground, either to go their own ways or to start the fracas again. I have never seen a bird hurt in these exchanges.
We experimented with setting up a mounted male chestnut-collar about 50 feet from an active nest. The male first threatened the dummy. Facing it with his head held high, he pointed his bill up at a 45-degree angle and raised his breast to show as much of the black as possible. He then dived repeatedly at the interloper, delivering blows to the head and back. Such continued chastisement would be unlikely under natural conditions where the intruder would retreat or fight back.
Courtship: Courtship in the chestnut-collared longspur seems largely a matter of staying close together. Up to the time incubation starts, when the male isn’t singing we can usually find the pair foraging together quietly in the more open parts of their territory. We saw a number of sexual chases, in which the male chased the female low over the prairie in fast, erratic flight, apparently trying to force her down. After a minute or so the male was seen to fly back to the foraging area, the female following along behind. As Tinbergen (1939) found with snow buntings, these chases probably result from the male’s unsuccessful attempt to copulate when the female is not in the right state of physiological receptiveness. The only display we saw a male make under natural conditions was to lift the tail high over the back momentarily on two occasions.
That a much more intense display is possible was shown by a male’s reaction to a mounted female placed near the nest. Approaching the dummy female in a zigzag course, he erected his chestnut collar, stretched his head high, and looked ddwn at her over his lowered bill from directly in front. He then fluttered to her back and attempted to copulate. Dismounting, he ran away from her holding his body level with the ground, wings and wide-fanned tail almost dragging. He then circled back and tried once more to copulate, dismounted, and circled to the front. There he assumed an exaggerated “female precopulatory posture.” in which he raised his closed tail as high as possible until his chest almost touched the ground, bent his head back on his shoulders with bill pointing straight up, and the wings projecting back and down. In this remarkable posture he rocked slowly from side to side several times. He then pecked at the dummy’s head, rump, and cloacal area as if in annoyance, and repeated the display. Possibly this behavior is elicited only when resistance is met, for I saw one pair copulate a dozen times without performing any such preliminary display.
In the matings I witnessed the female took the initiative. Typically she flies over and lands beside the singing male, lifts her tail straight up and tilts her head backward with bill pointing up and wings held low and quivering. The male mounts for a few seconds, dismounts, and stands beside her with his chestnut collar erect and bulging out from the rest of his plumage. The female holds the copulatory position for two or three seconds after the male dismounts, then assuming normal posture, fluffs out her feathers momentarily and assumes normal posture again. I once saw a female hold a piece of grass in her bill during coition.
The female’s reaction to a mounted female placed near her nest was to attack it fiercely, pecking at the head and throat. This suggests that the female drives off rival or trespassing females. I have found no reports of polygamy in the literature, nor did we find any indication of it in the field.
Nesting: The chestnut-collared longspur nests typically in uncultivated grasslands. At Winnipeg, Manitoba, R. D. Harris (1944) describes its favored habitat as consisting of “~ * * prairie, its flatness relieved by occasional low ridges and shallow sloughs. The dominant vegetation was composed of grasses of the following species: Panicum virgatum, Poa arida, Agrostis hyemalis, and Agropyron tenerum. Wolfberry (Symphoricarpos occidentalis) in straggling patches, prairie sage ~Artemisia gnaphaiodes), goldenrod (Solidago canadensi,s and S. his pida), and gum-weed (Gr-indelia squarrosa) were also present.”
In Montana, A. Dawes DuBois (1935) writes: “The Chestnutcollared Longspurs prefer to nest in the low and slightly moist situations where the thicker and taller grasses afford adequate concealment. If the meadow is wet or flooded the nests are placed on higher ground but are often near the moist margins. * * * twelve nests out of eighteen were in low places. However, one nest was high and dry on a knoll of the rolling prairie, one was on sloping land near a coulee, one in a fence border of native sod between wheat fields, and one was in a patch of grass on a dead-furrow left by the breaking plows, in the midst of a field of winter wheat which was almost knee high at the time the nest was found.”
At Regina, Saskatchewan, Margaret Beicher (1961) found the birds preferred sparse to heavy grassland: ‘~* * * I noted x~ith interest that the longspurs were more numerous in a 160-acre pasture that had been over-grazed than in an adjacent 320-acre pasture with a good stand of grass. Fox had a similar impression when checking a small pasture near Regina View School, southeast of the city, for Chestnut-collared Longspurs’ nests. In 1960 when there were no cattle in the pasture and the grass was long, no nests were found when the area was checked with a rope; in 1961 when the pasture was patchy from grazing, it was noted that the longspurs had chosen the bare patches for their nests.”
We found a few nests at Moose Jaw, Saskatchewan, in very sparse pasture with little cover for the nests. Though one nest was in quite thick grass over a foot high, the preferred habitat seemed to be iangrazed fields of medium thickness up to eight inches tall which offered good concealment and yet allowed the birds easy walking. The nests were built on level or gently sloping prairie, and not in the low areas that would flood in heavy rains. The steeper slopes were not suitable for the cup-shaped hollows they dig for their nests.
The nest is built by the female alone. Three different females we watched gathered the material a short distance from the nest site, mostly within 100 feet, and usually walked with it to the nest. DuBois (1935) describes the nest-building process as follows:
On the afternoon of May 22 I discovered a nicely rounded hollow in the ground, amidst the grass. There was no loose dirt near it. When I returned the next morning the nest proper had been started at the rim. There were only two weed stems at the bottom of the bole. Two and a half days later, at 9:30 am., the entire excavation has been lined with dead grass blades and stems, but the material was as yet rather loose in the bottom, while the rim was apparently finished. The grass material of the rim had been interwoven with the basal stems of the standing grass which grew around the nest. It seems especially noteworthy that the weaving of the rim was the first work done upon the nest structure. By 8 a.m. on May 27 (about four days after construction had started) the nest was apparently finished, with some white hairs added to the lining. The first egg was probably laid early the next day, as there were two eggs on the evening of the 29th, three on the morning of the 30th, and four on the morning of the 3 1st.
The body of all nests examined was composed entirely of dried grasses. These grass materials consist of both blades and stems, varying in different nests as to age and texture. One nest was made of old, soft, and shredded grasses; others had very old grass around the rim but fresher grasses in the bottom. Old, weathered grass in the rim is clearly an aid to concealment.
The materials used for linings include grasses, rootlete and hair, in various combinations and proportions. * * * The rim of the nest is usually flush with the surface of the ground.
The main building material for the nests seems always to be dried grasses, with animal hair or feathers added to the lining when they are available. Of 33 nests I examined only 3 contained material other than grass: one had seven gray partridge feathers in the lining and the other two each contained one small passerine feather. All these nests were circular in shape except one that was slightly oblong. Seven nest excavations I measured varied from 75 to 100 millimeters (average 88) in diameter, and from 43 to 55 millimeters (average 51) in depth. Nine nests I measured varied from 45 to 70 millimeters (average 60) in inside diameter, and from 38 to 45 millimeters (average 43) in depth of cup.
The nest is usually well concealed under a clump or tuft of grass. Even when placed in an open spot the nests blend so well with the pale prairie earth they are very hard to see, and the general sameness of the surroundings makes them even more difficult to find. L. J. Moriarty writes (in litt.)
A good way of locating the nest if you don’t find it when the female flushes: and she usually walks away from it before flying: ia to sit quietly not too far away and watch with binoculars her return. She will not fly directly to the nest, but will alight some distance from it and walk around apparently unconcerned, nonchalantly picking up food. As soon as she is convinced that no danger threatens, she will go to the nest and disappear into it to resume incubating. The nest is usually placed in a tuft of longer grass beside a stone of perhaps baseball size. I believe such sites are selected partly for the shade offered by the grass or rock during the noonday heat, and also because cattle will not step or lie down on stones of this size.
The female sometimes builds the nest close enough to the boundary of the territory to involve the male in a continual battle with his neighbor. In one such case the two nests were only 130 feet apart.
Eggs: The chestnut-collared longspur lays from three to five and sometimes six ovate eggs that niay show considerable variability even within a single clutch. The ground is creamy white, spotted, blotched, clouded, and with small scrawls of dark purplish-browns such as “dusky drab,” “bone brown,” or “olive brown” and black. The undermarkings are often very prominent blotches of “pale neutral gray,” or “deep purplish gray.” Some eggs have the entire surface clouded with gray undermarkings and with only scattered spots of the purplish-browns and black; others will show considerable ground with the markings concentrated toward the large end where they often form a loose wreath. The measurements of 90 eggs average 18.7 by 14.2 mm; the eggs showing the four extremes measure 20.8 by 14.7, 18.3 by 15.9, and 16.2 by 12.7 mm.
Incubation: The eggs are laid early in the morning, one each day on successive days until the clutch is complete. At Moose Jaw, Saskatchewan, the laying time, as nearly as we could determine, was between 6:00 a.m. and 7:30 a.m., central standard time.
Incubation generally does not start until the clutch is completed, and is entirely by the female alone. Some confusion has resulted apparently from the occurrence of male plumage characteristics in an occasional female.
The male spends his time during the incubation period in foraging, singing, defending the territory against encroachment by neighboring pairs, and chasing off small mammals and birds that approach the nest. One male approached the incubating female with a mouthful of insects, but was deterred by the close proximity of the blind. This behavior was not seen again during many hours of watching and may have been anomalous, perhaps anticipatory of later nestling feeding.
As Harris (1944) notes, the male * * * selects one or two definite stations near the nest. He watches attentively for his mate, and when she flies to or away from the nest, he follows and alights beside her. Now and then throughout the day he launches himself into the air, rises to a height of 10 to 50 feet, sPreads wings stiffly, and, floating slowly to earth, delivers his short, clear melody. At the approach of a human intruder, the male retires to his favorite perch, from which he marks the intruder’s progress. With uneasiness growing stronger, he takes wing and flies back and forth over his territory, giving utterance to a warning wheer note and sometimes a song. Some males have a habit on these occasions of reaching the highest point of their flight directly over the nest.
As incubation progresses the female sits very closely and leaves the nest only when almost trodden upon. Sometimes she flutters away in a distraction display with half-spread wings as though crippled. She leaves the nest of her own accord, however, every half hour or so and joins the male feeding, or she may fly to a nearby pond for a drink. If the female has not returned after a few minutes, the male appears agitated and flies to the nest, or he chases the female until she returns.
Duflois (1935) found the incubation period at one nest to be l2~4 days. Harris (1944) reports the period “from the laying of the last egg to the hatching of the last” to be 10 days at three nests. Our observations at one nest support DuBois’s 123~-day period: the last two marked eggs in a four-egg clutch were 12 days 13 hours, and 11 days 13 hours old the evening before they hatched. This is confirmed also by Moriarty’s (1965) timing of a marked clutch: “(2:30 p.m.). Egg No. 1 was hatched, 14 days plus 6 hours after laying. This was 11 days and 6 hours after I was sure incubation started. Eggs No. 2 and No. 3 were pipped at this time and hatched, young still moist at 5:30 p.m. By this time No. 4 was pipped, to hatch and dry by 7 p.m. It took about 1 to 13~ hours to complete hatching after the egg was pipped.”
The young may hatch over a period of a day or two. As Harris (1944) describes the process:
“In one observed case, hatching of one egg required over half a day, but in the majority of cases it seemed to take a shorter time. An irregular series of perforations is made by the young bird around the circumference of the shell about mid-way down the main axis. When the cut is completed, and the young bird has finally extricated itself, the two halves of the shell are carried away by the parent; pieces of shell have been found as far as a hundred feet away from the nest.”
Young: Both parents feed the young and tend to nest sanitation. The fecal sacs may be either eaten or carried away and dropped at a distance. For the first few days the female broods the young up to 50 percent of the daylight hours, and the male makes most of the trips with food. Daytime brooding, which is exclusively by. the female, ends about the fifth day, except during cold and rainy periods and thereafter the female makes more trips with food than the male. At one nest the young were fed 7.2 times per hour (5-hour watch) when 3 days old, and 16 times per hour (9-hour watch) when 9 days old. Watches at two nests of 8- or 9-day-old young on June 13 and June 23 showed the daily feeding period to last 17 hours.
The parents feed the young whatever insect food is available. Harris (1944) found his birds feeding their young almost exclusively on grasshoppers, but this was at a time when they were very abundant. At our study area the most common food seemed to be small green caterpillars and small brown segmented worms. The parents subdued these by biting them and beating them on the ground before giving them to the nestlings.
The following account of development is based on daily observations at four nests of individually color-marked young:
First day: The newly hatched young are very weak. They have just strength enough to gape for food as they lie in the nest. They appear well covered with down on the head and back after the down dries. They weigh about 1.6 grams at hatching, and gain 0.6 to 0.7 grams the first day. (Moriarty, (1944) reports four young weighed a total of 5.48 grams, or 1.37 grams each at hatching.)
Second day: The young are much stronger and gape for food at any noise made near the nest. They now weigh between 2.0 and 2.5 grams and will gain about 1.4 grams during the day.
Third day: The young are growing rapidly and the down is now about half an inch long. They will gain about 1.8 grams daily from the 3rd through the 8th day.
Fourth day: The eyes of the more developed young are open a slit. The feather sheaths are beginning to protrude from the skin. The nestlings begin to call with a soft “peeping.”
Fifth day: The eyes are about half open, but the nestlings still gape at sound rather than sight. Daytime brooding has almost stopped. Barring strong territorial competition with neighboring males, the male will stop singing about this time.
Sixth day: The eyes are now open and the nestlings will gape at the wave of a hand. The body feathers are beginning to break from their sheaths, but there are bare areas between the feather tracts.
Seventh day: The young gape at any movement near the nest. The feathers of the back and side are out and the primary sheaths are 10 to 15 millimeters long. The nestlings call when the parents bring food.
Eighth day: The young now crouch in the nest when disturbed, and can scramble away if placed on the ground outside the nest. The head feathers are out and the primaries beginning to protrude from their sheaths. There are still tufts of down above the eyes and a few bits elsewhere on the body.
Ninth day: The young are very active, moving around in the nest and preening. They are noisy when the parents bring food. They weight about 14.0 grams, and daily weight gain reduces to about 0.7 grams.
Tenth day. This is the day the young are most likely to leave the nest. R. D. Harris (1944) found the young left in 9,10, or 11 days. L. J. Moriarty (1965) reports them remaining through the 14th day, when they each weighed 12.21 grams. The young we observed weighed from 15.3 to 16.1 when ready to leave the nest.
The above account refers to normally developing young. Frequently, however, one nestling is smaller than the rest (possibly the last to hatch), and less successful than his siblings in competing for food. He either dies shortly after hatching, or may fall further and further behind to die later in the nestling period. A. D. DuBois (1937) writes:
“On July 19 I witnessed the removal of the dead fledgling from nest no. 14. It was a surprising feat of strength, for the fledgling was very heavy, being eight or nine days old. Upon rapidly fluttering wings the parent rose with his burden straight upward from the nest; then it flew horizontally, and, while flying low, dropped the carcass at a spot about fifty feet away.”
I saw a similar occurrence at Moose Jaw, when a female weighing about 20 grams carried a dead 14-gram nestling for about five feet before dropping it. Another dead young she dragged away from the nest.
Harris (1944) describes a nest-leaving he witnessed thusly:
The movement, perhaps stimulated by my activity around the nest, began without warning. The birds suddenly became very restless, kicking violently, and soon were panting hard for breath. After a few minutes they stopped simultaneously, and were quiet for about ten minutes. Again they began, and this time one bird, curiously enough the smallest of them all, pushed itself over the rim and crawled and hopped away from the nest in a wildly erratic course, finally coming to rest beside me two feet from the nest. Meanwhile, another bird, which had projected itself over the opposite side of the nest, turned back, and, shoving itself across the backs of its fellows in the nest, went toward the first one. The birds began to utter the chi-eep note and were answered by their parents, which were flying about overhead. After a general period of rest, a third one managed to scramble out, and the second one, in amazingly strong hops, followed an aimless course around the nest.
Of the post-nestling period he writes:
On the day of nest leaving, the bird is quite incapable of flight, and, except for occasional attempts at hopping, it remains crouched in the grass, receiving food from its parents. It grows, however, with extraordinary rapidity. After another day it is able to fly, when alarmed, for 100 feet or more. The flight is direct and labored. After alighting, the bird crouches upon the ground: I did not determine the age at which it is able to stand upright and walk.
Fourteen days after hatching (four days after leaving the nest), the young bird begins to use the eu-lip call note characteristic of the species. Its flight has now become undulating.
On the fifteenth day the bird is still being fed regularly by the male parent and occasionally by the female. If another nest is to be started, the female stops caring for the young at a time varying from two to seven days after they have left the nest; thenceforth they are in the sole care of the male.
By the twenty-fourth day, the bird appears to be fully grown. It may still be attended by the male parent, but it has sometimes to assume a begging posture, with wings outspread and fluttering, before the parent will give it food.
It begins to wander at large on about the twenty sixth day. If the parents are finished nesting, young and old go off together, but otherwise the young bird joins roving bands of juveniles.
The first clutches are usually laid about the middle of May. DuBois’s (1935) earliest clutch in Montana was May 6th. In Saskatchewan we found two clutches apparently completed about May 11th. New nests are started until well into July, which allows ample time for a second brood, or for several relayings after unsuccessful nesting attempts. Whether more than two broods are ever raised in a season is not known. Harris (1944) watched two pairs complete at least two nesting in a season. He notes “a second nest built by one pair was placed 40 feet away from the first nest; another pair built their second nest 100 feet from the first.” Two second nests we found were 50 and 100 feet respectively from each pair’s first nest.
Plumages: Harris (1944) says newly-hatched young are covered with buffy gray down about one-fourth-inch long. On the capital tract two rows of down, beginning at the loral region, run posteriorly to the occipital region, where they join a transversely placed tuft. An isolated tuft stands above each eyelid. A wide patch occurs in the spinal region, narrowing as it enters the pelvic region. Down is abundant in the humeral tract. In the alar tract, it is distributed in two rows on the dorsal surface. A prominent patch is found in each femoral tract, and scattered tufts can he detected in the crural tract. Mandibles are flesh color, darkening at the tip. Tarsi, toes, and claws are pale flesh color.
L. J. Moriarty (1965) adds that the mouth lining is “yellowishorange.”
T. S. Roberts (1932) describes the juvenal plumage as:
“Feathers of upper parts dusky centrally, edged with buffy and grayish-white producing a scaled rather than striped appearance; throat, abdomen, and under tail-coverts plain white; pale buff on breast, more or less thickly streaked and spotted with dusky-black; wings, including greater coverts, buffy-gray, the latter tipped with pale buff; middle and lesser coverts dark, tipped and edged with buffy-white, producing two rather indistinct buffy wing-bars; tail as in adult.”
He notes that in the first fall and winter plumage of the male: “The black of crown and underparts and the chestnut collar are largely acquired at the postjuvenal molt but are almost entirely obscured by buffy-white edgings of feathers. *** The first nuptial plumage is assumed before spring by the wearing off of theï light feather-edgings and a partial molt on head and breast. The adult male, after the postnuptial molt, is similar, but the concealed black is more extensive, and the middle and lesser wing-coverts are purer black and white.”
He describes the male in breeding plumage as:
Top of head, a broad line back of eye, a spot below ear-coverts, and underparts from chin to belly black; a small spot on occiput, a broad line over eye, lores, and a collar across throat white; chin pale buff (the white collar separating the buff from the black of lower throat); a collar across hindneck deep clear chestnut; back striped with dull black and gray, indistinctly so on rump and upper tailcoverts; wings, including greater coverts, brownish-gray; middle and lesser coverts black, tipped with white, broadly on the lesser forming a white patch; tertiaries brown, tipped and edged with light; the closed tail appears dark above, white below; spread it is largely white except middle pairs of feathers; the two outer pairs of feathers are almost wholly white, the others white at base with terminal oblique dark areas, decreasing in extent outward, thus not producing the terminal barred appearance seen in McGowns longspur. The wing4ining, a patch on either side of the breast, belly, and under tail-coverts white. The black of the abdomen is sometimes streaked with chestnut. Bill dusky above, light on cutting-edges and at base below; legs and feet dusky-flesh color; iris brown.
The female he describes as:
“A dull-colored bird, smaller than the male, striped throughout above with dull black, pale rufous, and grayish; a slightly paler collar across hindneck; below dull buff, lighter on chin and upper throat; faintly streaked on sides and across breast with fine dark lines. Some females are mottled with concealed black on breast and upper abdomen, and obscurely streaked on sides. Wings and tail as in male, except two very faint wing bars. Bill, legs, and feet dusky-flesh color, lighter than in male.”
We found great variation in the breeding plumages of both male and female birds in Saskatchewan, so much so that we could recognize many individual birds. Some males had white areas of various sizes and shapes in the otherwise black underparts, and their throats varied from white to buffy yellow. Many females showed vestiges of such male plumage characteristics as black breast markings or dull chestnut collars, but we saw none that reached the extremes reported by DuBois in Montana and H. D. Harris in Montana. As Harris (1944) tells it:
Occasionally a male is encountered in summer with areas of chestnut on the black underparts. Another anomaly is the occurrence of females in male plumage. DuBois (1935: 69, and 1937: 107) observed at least three female of this type, one “with all the male markings”; the others in an intermediate plumage, with the black underparts, but lacking the chestnut collar. A female with this intermediate type of plumage was collected on June 14, 1933, by T. M. and A. H. Shortt on my study area (it was carefully sexed). * * * The whole plumage was like that of a male, except that all the browns were paler.
C. G. Sibley and 0. 5. Pettingill, Jr. describe a hybrid between the chestnut-collared and McCown’s longspur they collected in an area where the two species are sympatric near Regina, Saskatchewan. The specimen was a male and showed intermediate characteristics between the two species in each area where they normally differ. The bird acted like a McCown’s, but had a flight song like that of the chestnut-collared.
Food: Very little has been published on the food of the chestnutcollar. F. M. Bailey (1928) describes it as “Largely weed seeds and insects, including crickets and grasshoppers, leaf beetles and weevils.” Roberts (1932) lists: “Seeds of grasses and other prairie plants; goosefoot, pigweed, witchgrass and grains; grasshoppers, crickets, beetles, bugs, caterpillars, wireworms, ants, etc.”
Martin, Zim, and Nelson (1951) give the analysis of 43 stomachs from various times of year as showing the diet 100 percent plant food in winter, and 72 percent animal food in summer. The animal food consists of “many beetles, grasshoppers, and spiders, as well as numerous other kinds of invertebrates.” Leading the plant foods are dropseedgrass, wheat, sunflower, and needlegrass, with lesser amounts of panicgrass, three-awn, pigweed, bristlegrass, and gramagrass, goosefoot, and sedge.
Voice: Sibley and Pettingill (1955) give the following comparison of the flight songs of the chestnut-collared and McCown’s longspurs: “The flight songs of typical McCown’s and Chestnut-collared longspurs differ in movements and in song pattern. Both species fly gradually upward, their wings beating rapidly. From the peak of the ascent McCown’s proceeds to sail downward abruptly with wings held stiffly outstretched and raised high above the back. The Chestnut-collared, after reaching the peak of the ascent, prolongs the flight by circling and undulating, finally descending with the wings beating as rapidly as before. Both species sing after the ascent, but the song of the McCown’s is louder with the notes uttered more slowly.”
The chestnut-collar does not use the flight song as often as the McCown’s does, nor does his performance appear as formal. The McCown usually nests on flat, open places such as very sparse prairie or plowed, summerfallow fields where it must make frequent, short song flights to advertise its presence. The chestnut-collared’s habitat is usually well supplied with song perches from which the singing male is easily seen.
D. J. Borror (1961) found the pitch range in the chestnut-collared’s song to vary between 2,300 and 6,500 cycles per second. He gives the following comprehensive description of the song:
The song is loud, rather musical, somewhat similar in quality to the song of the western meadowlark, and is generally 2 to 2.5 secs in length. It consists of 5 to 10 (usually 7 or 8) phrases uttered 3 to 4 per sec; the phrases may contain from one to several notes. The 3 or 4 introductory phrases contain clear notes, and are of decreasing pitch; the final phrases are usually huzzy. The first phrase or two of the song are usually quite weak. The phrases are not repeated in a given song, hut some phrases contain repetitive elements. The songs of a given hird are essentially alike, differing only in the character of the final weak phrases; the songs of different hirds usually contain slightly different phrases.
I was pleased to see that Borror disagrees with the earlier published accounts that seem to follow Coues’ (1903) description of the song as “weak and twittering.” From the inside of a blind, the song of a bird singing on the roof can be deafening! I could distinguish no difference between the songs given in flight and those while perched.
The male chestnut-collared spends a great deal of his time singing. Throughout the nesting season song continues from one hour before sunrise until shortly after sunset with little regard to the weather. The only significant break occurs when the nestlings get so large that most of his time and energy is taken up with providing food for them. At one nest, however, the male was still singing 54 times per hour when the young died eight days after hatching.
Harris (1944) gives the following accurate account of other notes: “The common call note is a til-lup or til-lip (the accent on the first syllable), sometimes lengthened to til-lil-Lip. It is a general flocking and flight note, and in the breeding season it seems to express anxiety. The usual alarm note is a whistled wheer, used mostly by the male. A tzip and a rattling fri-ri-rip indicate extreme alarm and perhaps anger. On coming to the nest with food, the female sometimes utters a soft lu, and the young then stretch open their mouths. Low, conversational notes are exchanged between the parents at the nest.” To which it should be added that birds also utter continuous buzz-buzz-buzz notes during sexual and territorial chases.
Behavior: The different roles the male and female chestnut-collared longspurs play on the nesting grounds result in wide differences in their behavior. The male is well suited to his role of guarding the territory against encroachment by rival males. He makes himself conspicuous as possible by his flight songs and by perching above the level of the grass. The female is quiet and tame. She rarely perches above the ground level and she seldom flies.
The female is so protectively colored and so agile afoot that she does not have to be very timid. As she walks through the grass near the nest she can be extremely hard to follow. As you approach she will usually walk quietly away from you, occasionally flying 10 or 15 feet to avoid being stepped on if you get too close. Her close sitting during incubation and brooding also has survival value in foiling potential predators. In one extreme case a female allowed me to photograph her from a distance of 18 inches, and then catch her by hand on the nest for weighing and color marking.
The only time the female makes herself conspicuous is when trying to attract your attention away from the nest. I have five records of distraction displays by females in my notes, each when the young were hatching. In each case the bird fluttered two or three feet into the air in a series of six-foot jumps with head held low and tail flicked wide to show all its white. John Horton of Moose Jaw told me of a male doing a “broken-wing act” and trying to lead him away from a nest containing young about a week old.
Although the chestnut-collared lives peacefully with the other grassland birds and animals, it tries to drive away any that approach its nest too closely. Our study area supported a heavy population of Richardson’s ground squirrels, and whenever one came near a nest, one of the pair, usually the male, immediately attacked it. Fluttering and chattering over the “gopher’s” head, the bird drops down to peck its head and back. Sometimes the gopher rears up at the bird, and I saw one actually jump into the air at the attacker. Or there may be a pause while bird and squirrel stand facing each other a few feet apart. Eventually the longspur gets the animal running away from the nest, and keeps chivvying it from above its back until it is a safe 80 or 100 feet away from the nest. I saw one bird pause after attacking a ground squirrel flick his bill several times as though to clear it of hair.
I have seen the longspurs similarly attack gray partridges; meadowlarks, cowbirds, and Savannah sparrows. On the other hand I have seen chestnut-collareds being chased by meadowlarks and Baird’s sparrows whose nests they probably approached too closely.
Though a resident of the dry prairies, the chestnut-collar is strongly attracted to water and will visit regularly any that happens to be nearby to drink and bathe. I kept watch at a pool left by recent rains on our study area when the females were incubating their first clutches. Birds visited the pool periodically from a few minutes after sunrise until just before sunset, and their appearances seemed fairly evenly distributed. In 5% hours of watching I never saw two male or two female longspurs at the pool at the same time. One female bathed and preened herself twice over while the male watched impatiently. After five minutes he finally crowded up to her and chased her back toward the nest, as if urging her to return to the eggs.
Field marks: The male in spring with his chestnut collar, black underparts, and white tail can be mistaken fo~ no other species. In other plumages the birds are rather nondescript, and can best be told from the vesper sparrows, pipits, and other longspurs that frequent open ground by the tail pattern. The spread tail is mostly white, with a small dark triangle in tbe center with its base at the end of the tail.
Enemies: By reducing the amount of grassland habitat, man and his agricultural practices are undoubtedly the chestnut-collar’s worst enemy. Yet the birds must suffer heavily from the depredations of badgers, skunks, weasels, and snakes, as do all ground-nesting birds. Though as related, the birds will not tolerate Richardson’s ground squirrels near their nest, I have no real proof of any predation by these animals. Such nest-robbing birds as crows and magpies must also take their toll. One morning a crow crouched in the grass with just his head showing about 150 feet from a nest with four eggs. The parents investigated him, and then went about their affairs apparently unconcernedly. The crow watched quietly for almost an hour, and then flew off without discovering the nest. Of the 31 nests we had under observation for more than one day, eggs or young disappeared from 11.
Young birds found dead in the nest are usually covered with ants. Though this is no proof that they caused the deaths, the female may often be seen cleaning ants out of t.he nest and picking them off the nestlings.
Hawks, owls, and shrikes also take a few birds without doubt, though the only record in the literature is M. F. Gilman’s (1910) report of a Cooper’s hawk preying on a flock in Arizona.
Cases of cowbird parasitism are rare. None of the 36 chestnut: collared nests we found in Saskatchewan was parasitized, though three of the four lark-bunting nests we found in the same area contamed cowbird eggs. None of the 38 nest-record cards in the Regina Museum of Natural History mentions cowbird eggs. Of the 23 nests Harris (1944) studied in Manitoba, only one was parasitized. Herbert Friedmann (1963) concludes “This longspur is probably a not uncommon local victim of the northwestern race of the brown-headed cowbird. The paucity of records seems to be due to a scarcity of observers in the breeding range of the host. * * * North Dakota is the only area where this longspur has been observed repeatedly as a cowbird victim.”
Fall: Harris (1944) gives the following fine description of the start of autumn migration in Manitoba:
The birds collected into flocks before turning southward. Young birds were the first to gather, frequenting the outskirts of the nesting areas. With the termination of nesting about the middle of August, adults joined the flocks of juveniles. The species then entirely abandoned the grassy breeding grounds, and was found in adjacent ditches, dried-up sloughs, and similar low-lying, rough ground (though rarely stubble or plowed land). This rather remarkable change of habitat may be due to the availability of the autumn crop of weed seeds, combined with the reduced number of grasshoppers, which constitute the bulk of the species’ food in summer; but the cause may lie deeper than that, and involve the psychological and physiological changes bound up with migration. Young and old together spent the last half of August in the new habitat, in loose, restless flocks numbering up to 30 or more individuals. During September, southward migration began, and the latest date on which I noted the species at Winnipeg was September 28.
The birds wander southward in large flocks to their winter range, pausing at good feeding and watering point en route. Some remain on the southern plains of the United States, the rest continue on to northern Mexico.
Range: Southern portions of Prairie Provinces south to northern Sonora, central Chihuahua, southern Texas, and northern Louisiana.
Breeding range: The chestnut-collared longspur breeds from southern Alberta (Lundbreck, Beaverhill Lake), southern Saskatchewan (Conquest, Quill Lakes), and southern Manitoba (Brandon, Winnipeg) southeast to northeastern Colorado (Weld County), central northern Nebraska (Holt County), and southwestern Minnesota (Jackson County), formerly to western Kansas (Ellis County).
Winter range: Winters from northern Arizona (San Francisco Mountain, Springerville), central New Mexico (San Mateo Mountains), northeastern Colorado (Fort Collins), and central Kansas (Lamed, Manhattan) south to northern Sonora (Pozo de Luis; San Pedro River at boundary), central Chihuahua (Chihuahua), southern Texas (Rio Grande City, Brownsville), and northern Louisiana (Gilliam); occasionally south to Puebla and Veracrus (Orizaba).
Casual records: Casual west to California (Point Reyes; Darwin) and central British Columbia (Kispiox Valley), north to northern Alberta (Fort McI~viurray), east to the Atlantic coast in Nova Scotia (Cape Sable), New Brunswick (Nantucket Island near Grand Manan), Maine (Scarborough), Massachusetts (Magnolia), New York (four Long Island localities), and Maryland (Ocean City), and south to northern Florida (Tallahassee).
Migration: Early dates of spring arrival are: New York: Orient, April 21. Minnesota: Jackson County, April 12 (average of 5 years for southern Minnesota, April 30). North Dakota: Jamestown, March 26. Manitoba: Treesbank, March 24 (average of 23 years, April 11). Saskatchewan: Indian Head, March 30. Wyoming: Lararnie, March 16. I’.Iontana: Big Sandy, April 21; Terry, April 26.
Late dates of spring departure are: Texas: Austin, April 26. New Mexico: Gage, April 17. Arizona: Huachuca Mountains, May 3.
Early dates of fall arrival are: California: Inyo County, September 28. Arizona: San Francisco Mountain, September 18; Mount Graham, September 24. New Mexico: Willis, September 12. Oklahoma: Copan, November 11. Texas: Austin, November 12. Maine: Scarborough, August 13. Massachusetts: Magnolia, July 28. New York: Miller Place, September 14. Maryland-Ocean City, August 20.
Late dates of fall departure are: Montana: Miles City, September 27; Terry, September 23. Manitoba: Treesbank, October 12 (average of 24 years, October 3). North Dakota: Jamestown, October 7. Minnesota: Lac qui Parle County, October 12.
Egg dates: Alberta: 6 records, May 27 to June 11.
Minnesota: 23 records, May 19 to June 16.
Montana: 36 records, May 6 to July 19; 19 records, June 6 to July 7.
North Dakota: 27 records, May 23 to June 30; 19 records, May 26 to June 10.
Saskatchewan: 55 records, May 10 to July 9; 36 records, May 20 to June 10.
South Dakota: 19 records, May 20 to June 17; 13 records, May 22 to June 5.