Lapland 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 Lapland Longspur – the common name and sub-species reflect the nomenclature in use at the time the description was written.

In North America the range of the Lapland longspur extends from northwestern Mackenzie over the vast expanses of tundra of the arctic and subarctic regions northward to Ellesmere Island and eastward to the Atlantic Ocean in Labrador. The following remarks show the habitat this species prefers remains essentially the same over this enormous region. T. H. Manning, 0. Hohn, and A. 11. Macpherson (1956) found longspurs most numerous on Banks Island in “marshes and better vegetated country.” It occurs throughout Prince of Wales Island except on the barren disintegrated limestone uplands (Manning and Macpherson, 1961). J. Dewey Soper (1946) remarks that the species is scarce in the mountainous eastern half of Baflin Island and reaches the height of its abundance on the great tundras of the west where “it inhabits localized flats of a swampy nature, but its favorite breeding grounds are wide, moist tundras interspersed with ponds and streams and studded with grassy tussocks.”

V. C. Wynne-Edwards (1952) states that the most suitable habitats at Clyde Inlet on Baffin Island are wet tussocky meadows, and in the southern part of that island Sutton and Parmelee (1955) found the longspur inhabiting all wet grasslands of the vicinity. They remark:

Precisely why wet grasslands are so all-important to the longspur may be difficult to say. Availability of insect food for the nestlings, of dead grass for building material, of nest sites not readily accessible to certain predators: these probably have a part in determining the matter. To he attractive the terrain must be more than low and flat; if too well drained, too gravelly, too bare, it will not do. Dry, firm, thinly grassed areas just north of the Base were inhabited by Horned Larks and Semipalmated Plovers, but the longspurs lived elsewhere: a stone’s throw away, in the wet grassy places.

W. H. Drury (1961) describes the preferred and somewhat different habitat on Bylot Island as follows: “Longspurs occupied the thick, moss-floored vegetation of dry places in the uplands, primarily on the east- and south-facing slopes, not the bogs. Their distribution was similar to that of the Bell Heather [Ca.ssiope tetragona], but Bell Heather grew over large areas where we found no longspurs.”

On Southampton Island, Sutton (1932) found that nesting-sites were restricted to the grassy margins of streams, or to hummocks of grass in the little marshes.

I would add to these generally consistent remarks only my belief that many references to “grasses” actually refer to the numerous sedges (Carex) of the arctic, and that the species generally nests in those moist areas that are not excessively wet. The situation on Bylot Island Drury describes seems exceptional and somewhat at variance with the notes of others. However, Adam Watson (1957) states that longspurs attained peak densities on the Cumberland Peninsula of Baffin Island wherever the ground became slighly drier and more heathy with a good growth of willows (Salia~) and became uncommon on drier ground where the heath was closed. Drury (1961) suggests that the breeding sites on Bylot Island were selected “because the wet meadow sites were snow-covered too late in the species’ breeding cycle and because the species, displaced from ideal nest sites, selected a form resembling the overhang of a tussock.”

In the extreme northern part of the range on Ellesmere Island, habitat apparently suitable for longspurs is unoccupied. David F. Parmelee observed about 20 pairs breeding on the Fosheim Peninsula (80~ N.), but recorded none in roughly 400 square miles of seemingly suitable tundra between Slidre and Greely Fiords. J. S. Tener (1961) recorded an extreme northern nesting on Ellesmere Island when he saw a female feeding a flightless young a few miles north of Hazen Lake at approximately 850 N.

Spring: A sampling of the scattered records of spring arrival show a synchrony consistent with that observed in the ensuing nesting activities. At Bathurst Inlet, Mackenzie District, E. H. McEwen (1957) observed the first longspur to arrive, a female, on May 21, and 10 days later saw four males feeding in a sedge-covered flat. T. H. Manning et al. (1956) were present on Banks Island (De Sails Bay) in 1952 when the first longspurs arrived on May 29, when three were seen. The next day they counted 115 in 7)~ hours, mostly in flocks of 25. Males outnumbered females on that date and on May 31, but on June 1 the sexes were about equally represented. In 1953 the first longspur (a male) was seen at Sachs Harbor on May 19, and one was heard at Cape Kellett on May 21. Their numbers increased thereafter, and most of the birds seen up until May 26 were males. The migration was essentially over by June 3, at which time the birds were seen in pairs. Manning and Macpherson (1961) first observed longspurs on Prince of Wales Island on June 14, and remarked that the late date of arrival was likely due to the late thaw on the northern part of the mainland barrens. The males outnumbered the females by 4 to 1 for 3 or 4 days.

The severe climatic conditions arriving longspurs frequently meet throughout their range is well-described by Sutton (1932) from observations on Southampton Island:

On May 20, 1930, during a snow-storm, which eventually developed into one of the worst blizzards I ever experienced, I saw and collected a male Lapland Longspur at Itiujuak. The bird was associating with a large flock of Snow Buntings, practically all of which were males. It was f at. The gonads were much enlarged. The stomach and crop held particles of vegetable food and some bits of quartz crystal, but food had evidently been difficult to find. I noted the brightness of the yellow bill, the base and culmen of which had a slight greenish cast. I looked carefully for other longspurs in this flock of buntings, but saw none.

On May 27 an adult male was seen perched on the very edge of the ice, perhaps six miles from land, at the floe west of Native Point. We heard one (perhaps the same individual) from our tent on May 28. On the following day a male was seen at the Post, feeding with the Snow Buntings during a wind-storm. This bird was very weak, probably from starvation, and I nearly caught it in my hands. During the midst of the storm it flew upward, was caught in a blast of wind, was hurled southward, and did not return.

On May 30 I collected a specimen from a flock composed of five longspurs and a few Snow Buntings. AU these birds were males. On May 31 a male longapur was seen at the Roman Catholic Mission and another at the Factor’s dwelling.

On June 1 large, loose flocks, entirely composed of males, assembled on the bare ground about three miles north of the Post. There must have been two hundred birds in all. I collected seven. The gonads of all were considerably enlarged; and all were fat, but their gizzards were not full of food. The birds worked their way through the short grass diligently, walking for a time, then stopping to look round, only to continue their search. When they came to the edge of a bare patch, they usually flew over the snow to the next bit of open ground. In some places the tundra had many snowless patches, and here with almost mathematical precision each such patch of open ground harbored its single longspur, which searched carefully over all its chosen space, not neglecting the shadowy spots at the edge of the snow.

The retention of substantial deposits of subcutaneous fat by arriving migrants has been remarked upon by other writers dealing with arctic passerine birds, and this phenomenon can readily be seen to have considerable importance to the welfare of the birds settling upon a wintry landscape.

In southern Baffin Island Soper (1928) observed the first longspurs at Cape Dorset on June 3, and saw them sparingly the following few days. Soper (1946) states that they arrived at nearby Bowman Bay on June 4, and in three days both sexes were common. By June 10 they swarmed in thousands over the restricted strips of snow-free tundra. The numbers increased “up to the middle of the month, when the majority moved on to more northern resorts. Large numbers remained to breed, however, and by a wide margin were the most abundant summer residents of the tundra. On June 11, the males were first heard singing half-spirited snatches of song, but the first mad outbursts of their full and brilliant repertoire was not general until June 20.” In central Baffin Island, Wynne-Edwards (1952) observed the first longspur, a male, on May 29; three days later most of the males had arrived and females were seen.

D. B. 0. Savile (1951) states that two males and a female appeared at Chesterfield Inlet, Keewatin on May 19, 1950; on May 21 there were eight males and two females, and thereafter numbers increased; males were still somewhat predominant on June 7. The early arrivals frequented mainly the wind-swept beach ridges where there was little snow and seeds of the sea-beach sandwort, Arenaria peploides, provided abundant food.

Courtship and nesting: The courtship and nesting habits do not appear to differ from those described for the Alaskan race of this species. The same general and striking synchrony of activities in any single locality is apparent over the entire range of the species. Dates of the onset of courtship and nestbuilding may vary between years at the same or nearby localities, apparently because of weather, and consistent variation between populations may be due to latitude (indirectly to weather), or perhaps to local habits.

Sutton’s (1932) comments on the breeding schedule on Southampton Island are pertinent for the species as a whole: “On June 8 I collected another female and observed the first flight-songs. In the North Country, the program of the individual is virtually that of the species. When one male bird begins to sing, they all begin to sing; by the time one pair mates, all the pairs are mating; when the first egg is laid by one female, all the females of that species are laying the first egg, and so on throughout the length and breadth of the tundra.” This is certainly the first impression an observer of this species gets, and it can be carried further to include the molt of young and adults. Although the onset of particular activities among individual birds in the same population usually varies somewhat more than Sutton describes, his remarks point out fairly enough what is to me the most striking single adaptation birds have made to life at high latitudes: the general synchrony of events on the nesting ground and the compression of the time intervals between them.

On Banks Island, Manning et al. (1956) found pairs occupying nesting territories as early as June 1, although prior to June 2 the largest ovum found in a female was 1.5 millimeters. On June 5 a female was taken with a complete egg in the oviduct and three or four others had fully developed eggs in the next few days. Manning and Macpherson (1961) in their study of late-arriving longspurs on Prince of Wales Island (discussed previously) state that: “On June 26 a male was seen copulating with NMC 43094, a female with no empty follicles and rather small ova. When she was shot he turned his attention to NMC 43095, which had six empty follicles. The first nest was found on June 28. It contained six eggs. Another with six eggs was seen on June 30. Males continued to perform their nuptial flights regularly until the end of June, and some were singing until about July 10.”

Drury (1961) arrived at Bylot Island while the longspurs were still in flocks and their numbers were still increasing daily. Males took up territories between 15 and 25 June, copulation was observed on June 21, and eggs were laid as early as June 17 with clutches completed 22 June to 4 July. Wynne-Edwards (1952) found egglaying starting as early as June 6 on Biaffin Island.

More extensive variation in nest~lining materials than that noted for the Alaskan subspecies is interesting to note. Field notes of J. R. Cruttenden mention the use of caribou hair in a nest at Churchill, Manitoba. Wynne-Edwards (1952) noted ptarmigan and longspur feathers, plant down from willows, and hair from the arctic hare. Sutton and Parmelee (1955) observed linings of raven feathers and dog hair, while Watson (1957) found ptarmigan feathers, hare wool, lemming hair, and plant down from willows in nests on Baffin Island.

Eggs: Clutch size of the Lapland longspur remains generally consistent over the entire range. Drury (1961) observed an average clutch size of 4.6 (15 nests) on Bylot Island and found that the removal of eggs during and after laying had no effect on the number laid. Sutton and Parmelee (1955) found the average to be 4.4 in 22 nests on Baffin Island; all of these figures agree closely with the 4.7 average observed in northern Alaska. Wynne-Edwards (1952) based his larger figure of 5.4 on Baffin Island on only ten nests, Sutton (1932) obtained bonafide evidence of a larger average clutch size in an eastern population on Southampton Island, where 78 nests held an average of 5.8 eggs and 71 of these clutches, found between June 21 and July 21, contained six eggs. It is of interest that while no seven-egg clutches were observed in Alaska, Watson (1957) and Wynne-Edwards (1952) observed one and two of this size, respectively.

Wynne-Edwards (1952) presents evidence of a decline in clutch size as the season progresses: “five clutches begun between June 6 and 15 numbered 5, 6, 6, 7, 7, and five begun between June 19 and July 2 numbered 4, 4, 5, 5, 5. (The probability is about 30 to 1 against the apparent difference being due merely to chance.)” Drury (1961) states: “Early nests (hatching 3-9 July) contained consistently larger clutches (6, 6, 6, 6, 5, 4) than later nests hatching 10: 15 July (5, 5, 4, 4, 4, 4, 4, 3, 3).” Sutton and Parmelee’s (1955) data show 11 clutches hatching before July 3 averaged 4.7 eggs and ones hatching later averaged 4.3 eggs.

Egg color and pattcrn seems to be so generally variable over the range of the species as to merit no special discussion for this race. The measurements of 50 eggs average 20.9 by 15.3 millimeters; the eggs showing the four extremes measure 23.9 by 16.8, 18.8 by 15.8, and 19.4 by 142 millimeters.

Incubation: Considerable variation exists in the onset of incubation with respect to number of eggs laid, which may be a matter of adjustment by individual females to local climatic conditions. If so, it could be construed as an interesting adaptation to arctic conditions. Wynne-Edwards (1952) states that “Incubation begins with the first egg” because, as he elaborated in his discussion of the redpoll, “the temperature in June is below freezing for much of the 24 hours.” He found, however, that the actual hatching period in six cases ranged between 2 and 4 days. Watson (1957) found that the females at two nests definitely started to incubate from the laying of the first egg, and early incubation was suspectcd in most other nests, although in a few it definitely did not occur. At one nest the eggs were cold on many visits during the first two days of laying, but warm after the third day. At another nest the female did not sit for at least all of the two afternoons of the days she laid the first two eggs; here three young hatched the same day followed by the fourth early the next day. A much greater spread in hatching resulted from the early incubation at most nests. At the nest where all 5 eggs hatched, the hatching occupied at least 3 full days, and about 3 days at another nest where 5 eggs out of 7 hatched.

Drury (1961) notes similar variation in the onset of incubation and he discusses it for other populations throughout the range of the species.

The length of the incubation period appears to be generally the same throughout the range, 10 to 13 days (Drury, 1961), 12 days (WynneEdwards, 1952), 12 days (Sutton and Parmelee, 1955), 13 to 14 days (Sutton, 1932).

Young: Nearly every author has commented on the early departure of the longspur young from the nest: 9 days (Drury, 1961), 9 to 10 days (Wynne-Edwards, 1952), and 9 to 10 days (Sutton and Parmelee, 1955). Watson’s (1957) observations on Baffin Island are generally appropriate for the species. He states that nestlings showed a marked gradation in size, at some nests so great as to suggest periods of hatching in excess of 2 days. This gradation was evident in 8- to 10-day-old broods at the time of leaving the nest. In at least four nests the smallest individual was left behind in the nest and was apparently deserted and no longer fed for up to a day after the others had gone. Two such young died, one in the nest and one outside. Another that fluttered from the nest after most of a day alone was not more than 7 days old and had all retrices and primaries ensheathed. Most of the young left when 8 or 9 days of age, when the retrices and primaries were just beginning to appear. About the third week of July most of the young being fed by the adults were unable to flutter more than a few yards and their tails scarcely showed. Many seemed to have been deserted and dead young were often found.

Wynne-Edwards (1952) suggests that the habit of leaving the nest early may in part be directly correlated with increasing the clutch size without increasing the size of the nest.

Certainly towards the end the impression was given, both by the redpolls and the longspurs, that the nest was ready to burst and I was prepared at once to assume that dead chicks found in redpolls’ nests had been suffocated. There is moreover the growing danger of attracting the arctic weasel as the young become daily noisier and as the odor of feces mounts. (The latter probably also attracts the blowflies, whose larvae were found in all the examined nests of longspurs, snow buntings, redpolls, and wheatears.) It is not difficult therefore to see advantages in scattering the brood as early as possible, since it reduces the danger both from predators and from overcrowding in the nest. But the parents are probably presented with a more difficult task in providing enough food for each of the young after they are scattered, and the young are also deprived of the protection from cold and wet, and in general the uniform environment which they have previously enjoyed in the nest. It may be presumed that the advantages outweigh the dangers, and that once the young have acquired sufficient control of body-temperature their chances of survival are increased by dispersal.

Two additional important points must not be overlooked: that the frequent marked disparity in size of the nestlings can result in one or more remaining in the nest only to be abandoned as Watson noted, and that the chief predators of fledged longspurs are avian ones, mainly jaegers, to which dispersed young are certainly more viiinerable than those in nests.

Density: A number of authors have remarked on the generally high density of this species throughout the tundra regions. Manning et al. (1956) believed it to be by far the commonest and most generally distributed bird on Banks Island, and estimated the total adult summer population at 700,000. Likewise, Manning and Macpherson (1961) believed that the Lapland longspur was the commonest summer resident on Prince of Wales Island and from density figures of up to 56.8 birds per square mile estimated 250,000 present when nesting began. Sutton (1932) states that it was the most abundant bird on Southampton Island and comments on its general distribution. Drury (1961) refers to the longspur as the most numerous nesting species near his camp on Bylot Island (60 pairs in 7 square miles), and Wynne-Edwards (1952) states that at the head of Clyde Inlet on Baffin Island longspurs were the commonest birds, and in the most suitable habitats reached a density of one pair in 5 to 15 acres.

Much greater densities were recorded in Alaska, though this may have been due to the careful and intensive census methods used. The total population figures of young and adults can be exceedingly great in late summer, as mentioned in the account of the Alaskan subspecies.

Plumages: As mentioned in the account of the Alaska longspur, the two races are only doubtfully distinguishable. The eastern race is somewhat darker and, according to Manning et al. (1956) the eastern lap ponicus have wider and more deeply black centers in the dorsal feathers, and narrower more tawny and less buckthorn brown, but not markedly darker margins. The sequence of molts and plumages is the same in both races.

Winter: In Minnesota, T. S. Roberts (1932) writes:

During October and November immense flocks are passing southward across the state to winter in the southern Mississippi Valley beyond the winter range of the Snow Bunting. A very considerable number, however, remain in the southern half of Minnesota where they pass the winter months in the prairie and semiprairie regions, generally in great flocks, often mingling with the Snowflakes or fetding about the corn-and-weed-fields with the hardy Horned Larks. * * * The northward movement begins in Minnesota in late February and early March, and during the latter month and April countless myriads of these birds are traversing the state toward their Arctic homes, going in great, endless flocks, generally by night but often by day as well, passing steadily onward, sometimes high in the air, sometimes low over the earth, an incessant, twittering, hurrying stream of birds.

On February 25, 1878, Roberts “Found a flock in an old corn-field where, rising from the ground, they circled around and around for some time, occasionally crossing and recrossing the circle, parting above and around me and streaming by within a few feet. At length, they settled on a near-by fence and showed very little fear at my presence.” Again, he says on March 16, 1880:

When disturbed they would fly up two or three at a time and perch in the bushes, where, if not further molested, they would make an attempt at singing. On being startled into the air the whole flock circled about for a time, now and then leaving as if for good and as often returning to alight on the ground, or, in two or three instances, in the tops of a neighboring grove of small oaks. No sooner were they fairly settled in the trees than all burst out singing, and, although thus early in the season the individual effort was neither full nor continuous, the general chorus was quite loud and pleasing and kept up without breaks. The effect was very much like that produced by a flock of Blackbirds. These birds were not in very full plumage.

By April 23 the birds “were then in nearly full spring plumage and uttering an agreeable musical song.” E. T. Seton (1891) describes generally similar behavior of spring migrants in Manitoba.

In Maine, R. S. Palmer (1949) quotes A. H. Norton’s notes to the effect that: “the small flocks at Scarborough are found in sheltered places about the edges of grassy areas and out on the sand where vegetation is sparse. Here their food is mainly the seeds on the long spikes of marram grass (Ammophila sp.) and the seeds of orache (Afriplez sp.). Inland, they have been seen in fields where ragweed (Ambrosia sp.) and other weed stalks protrude above the snow. Associates, in decreasing order of frequency noted, are Horned Larks, Pipits, and Snow Buntings.”

C. W. Townsend (1905) says that in Massachusetts “Lapland Longspurs, although occasionally found by themselves, are more apt to be associated with Horned Larks and Snow Buntings. Flying and feeding with these birds, they generally keep together, however, in one part of the flock, although a few scattered birds are not uncommon. They frequent the dunes at Ipswich and the neighboring hills by the sea * * *

In a letter to Mr. Bent, Wendell Taber wrote: “On February 3, 1935, following a long spell of below-zero weather, I saw five of these birds on Ipswich beach in Massachusetts, accompanied by one snow bunting. My wife and I walked right up to the birds. A number of times I was within 4 to 6 feet of a bird. Ultimately I laid down and stretched out on my stomach near a bird. The bird kept right on feeding within a few feet of me, and came within 18 inches of my elbow. I watched it take a mouthful of snow and, seemingly, ‘chew’ it up and swallow it.”

Geoffrey Carleton writes in a letter from New York:

On March 28,1965 Mr. John L. Bull, Eugene Eisenmann and I were on a remote part of the Kennedy International Airport. It was a sunny afternoon with a sharp, cold sea breeze. About 25 Lapland longspurs Bhowed a preference for the tops of rocks piled along the shore as a breakwater, many of them evidently quarried for the purpose and 3 or 4 feet thick. Some of the longspurs were singing, particularly those most advanced into breeding plumage. Some were crouched so as to be more sheltered from the wind, and I believed they were taking advantage of the warmth of the rocks in the Burl. After short flights or periods of feeding on a nearby grassy bank, the birds returned to the rocks and resumed singing.

DISTRIBUTION
Range: Central Franklin, Greenland, northern Russia, and northern Siberia south to northeastern Texas, southern Louisiana, northwestern Mississippi, southwestern Ohio, eastern West Virginia, and Virginia.

Breeding range: The common Lapland longspur breeds from central Franklin (Banks, Prince Patrick, Melville, and Devon islands), Greenland (north to Thule and Scoresby Sound), Norway (lat. 710 N.), northern Russia (Arkhangelsk Government), Franz Josef Land, Novaya Zemlya, and the tundra of northern Siberia, including the New Siberian Islands and Wrangel Island, south to central western Keewatin (between Casba and Baker Lake: Hanbury River), northeastern Manitoba (Churchill), northern Ontario (Little Cape, Cape Henrietta Maria), northern Quebec (Cape Jones, Fort Chimo), northern Labrador (Okak), southern Greenland (Cape Farewell), southern Norway (Dovre), and central Sweden (lat. 630 N.); in eastern Siberia around the Gulf of Anadyr; recorded in summer north to Ellesmere Island (Slidre Fiord, Lake Hazen), and in Greenland to Germania Land.

Winter range: Winters from central northern Colorado (Barr), central northern Nebraska (Wood Lake), central Minnesota (Otter Tail County), central Wisconsin (Oconto County), central Michigan (Newaygo and Tuscola counties), southern Ontario (Kitchener), southern Quebec (AyLmer), Vermont (St. Johnsbury), Maine (Lubec), New Brunswick (St. John), and central Nova Scotia (Grand Pro) south to Oklahoma, northeastern Texas (Dallas), southern Louisiana (Jennings, New Orleans), northwestern Mississippi (Rosedale), Alabama (south to Marion and Montgomery), western Tennessee (Memphis), southwestern Ohio (Hamilton County), eastern West Virginia (Moorefield), and Virginia (Blacksburg, Shenandoah National Park, Falls Church, Back Bay); casually to Utah (Tooele County), Georgia (Augusta), Florida (Wilson), and Bermuda; and from England, northern Europe, and central eastern Siberia to France, northern Italy, Rumania, southern Russia, and Altai, casually to Iceland.

Casual records: Accidental in Baja California (Isla Cerralvo).

Migration: The data deal with the sp~cies as a whole. Early dates of spring arrival are: Pennsylvania: State College, March 22. New York: Nassau County, February 7. Massachusetts: Martha’s Vineyard, February 27. New Hampshire: New Hampton, March 15. Quebec: Seven Islands, May 25. Illinois: Urbana, February 12; Chicago, March 20 (average of 6 years, April 19). Michigan: Battle Creek, March 27. North Dakota: Red River Valley, February 19. British Columbia: Okanagan Landing, March 9; Mackenzie Delta, May 16. Yukon: Sheldon Lake, April 29; Old Crow,May4. Alaska: Yakutat, April 6; Unimak, April 16; Homer, April 19; Dyca, April21; Amchitka Id., April 25; Adak, April 25; Cape Thompson May 2; Attu, May 2; Cold Bay, May 5; Fort Yukon, May 4; St. Michael, May 1; Pt. Barrow May 20.

Late dates of spring departure are: Alabama: Courtland, March 27. Georgia: Qysterbed Island, March 24. South Carolina: Long Island Fill, April 20. North Carolina: Raleigh, February 20. Virginia: Blacksburg, May 1. Maryland: Baltimore County, February 10. Pennsylvania: State College, April 4; Erie, March 25. New Jersey: Cape May, March 24. New York: Long Island, May 9. New Brunswick: Grand Manan, March 28. Louisiana: Baton Rouge, February 4. Missouri: St. Louis, April 15 (median of 8 years, March 2). Illinois: Chicago, May 13 (average of 6 years, May 7); Urbana, April 21. Ohio: central Ohio, April 12 (median of 40 years, March 25). Michigan: Detroit, April 24 (mean of 10 years, April 19). Iowa: Sioux City, April 4. Minnesota: Lac Qui Pale County, May 30 (average of 16 years for southern Minnesota, April 8). Oklahoma: Payne County, March 31. Kansas: northeastern Kansas, April 12 (median of 10 years, March 12). Nebraska: Red Cloud, April 18. North Dakota: Cass County, May 21 (average, May 18); Jamestown, April 25. Manitoba: Treesbank, February 20. Wyoming: Laramie, April 3 (average of 5 years, March 17). Montana: Gallatin County, April 17. Washington: Destruction Island, April 28. British Columbia: Atlin, May 15.

Early dates of fall arrival are: British Columbia: Kispiox Valley, September 1. Washington: Westport, September 5. Oregon: Silt-coos Lake, October 15. Nevada: Mercury, October 10. California-San Diego County, October 2. Montana: Deer Lodge County, October 30. Idaho: Fort Sherman, November 13. Wyommg: Laramie, October 7 (average of 8 years, November 2). Anzona: Coconino County, November 15. Manitoba: Treesbank, August 24 (average of 22 years, September 6). North Dakota: Cass County, September 10 (average, September 18); Jamestown, September 18. Nebraska: Red Cloud, November 28. Kansas: northeastern Kansas, November 1. Texas: Austin, November 27. Mmnesota: Kittson County, September 8 (average of 5 years for northern Minnesota, October 21). Iowa: Sioux City, October 26. Michigan: Detroit area, September 25 (mean of 10 years, October 20). Ohio: central Ohio, November 1 (median of 40 years, November 15). Illinois: Chicago, September 17 (average of 11 years, October 4). Missouri: St. Louis, October 28 (median of 10 years, November 10). Tennessee: Nashville, December 31. Arkansas: Arkansas County, November 16. New Brunswick: Memramcook, October 1. New Hampshire: Pittsburg, September 30; New Hampton, October 22. Massachusetts: Martha’s Vineyard, October 18 (median of 5 years, November 1). New York: Long Island, October 3. New Jersey: Cape May, November 14. Pennsylvania: Erie, October 3; State College, October 26. Maryland: Laurel, December 3. Virginia: Shenandoah National Park, October 22. North Carolina: Swannanoa, November 14. South Carolina: Chester, January 1. Georgia: Augusta, January 1. Alabama: Gadsden, November 14; Marion, December 1. Florida: northwestern Florida, October 25.

Late dates of fall departure are: Alaska: Demarcation Point, September 2; Point Barrow, September 4; Nome, September 10; Attu, August 30; Glacier Bay, October 11; Kodiak, October 23; Cold Bay, October 25. Yukon: Macmillan Pass, September 4. British Columbia: Atlin, November 1. Manitoba: Treesbank, December31 (average of 21 years, November 16). Kansas: northeastern Kansas, November 26. Illinois: Chicago, December 14 (average of 11 years, November 21). New York: Nassau County, November 26.

Egg dates: Baflln Island: 52 records, June 12 to July 9; 25 records, June 22 to July 2.

Victoria Island: 61 records, June 11 to July 21; 31 records, June 15 to June 24.

Banks Island: 3 records, June 18 to July 10.

Bylot Island: 15 records, June 22 to July 4.

Manitoba: 15 records, June 12 to June 29.

Prince of Wales Island: 3 records, June 28 to July 1.

Southampton Island: 78 records, June21 to July 17.

ALASKA LONGSPUR
CALCARIUS LAPPONICUS ALASCENSIS Ridgway
Contributed by FRANCIS S. L. WILLIAMSON
HABITS
It is frequently said that every biologist should, at least once during his life, visit the tropics, particularly an equatorial rain forest with its amazing wealth of plant and animal species and its concomitant ecological complexity. More than elsewhere on earth the tropical environment affords unequalled opportunity to gain appreciation for evolutionary diversification. I think, however, it must also be said that every naturalist should at some time visit the arctic tundra, a relatively bleak and windswept environment characterized by a striking diminution of species of plants and animals and a concomitant ecological simplicity. These are factors which tend to make studies of adaptation and of ecological rclations especially rewarding.

For the vertebrate ecologist or zoologist there is no more suitable subject for studies of such relations than the Lapland longspur, in all likelihood the most characteristic species of vertebrate animal to be found throughout the arctic. The extensive circumpolar breeding range of this species is equaled by few other birds, and the generally consistent high density of individuals over such a vast area is likely not equaled at all. The Lapland longspur is found on the tundra of Kamchatka west through Siberia to Scandinavia, Franz Joseph Land, Greenland, northern Canada, and Alaska. In winter it can be found occupying large areas of southern Europe, Russia and China as well as most of the United States.

The great bulk of this imposing range is occupied by one subspecies, lapponicu.s, while two others, coloratus (Commander Islands, Kamchatka), and alascensis (Alaska, N. Yukon, N. W. Mackenzie) occupy more restricted areas. In Alaska, principal breeding area for the race alascensis, the distribution conforms essentially to the treeless coastal regions and islands. These include a relatively broad belt of tundra extending from the Arctic coast south and west along the Chukchi and Bering Sea to the base of the Alaska Peninsula. Westward in the Aleutian Islands the range extends to Attu (Murie, 1959). More peripheral range includes a small nesting group on Middleton Island in the Gulf of Alaska (Rausch, 1958), St. Lawrence Island (Pay and Cade, 1959), and the Pribiof Islands (Preble and McAtee, 1923). In the interior of Alaska longspurs occupy areas above timber in scattered localities including the Talkeetna Mountains (Schaller, MS) and the central Alaska Range (Dixon, 1938).

The habitat these small birds occupy within this geographic range includes nearly all that of open character. Generally exeluded are the extremely wet marshlands and meadows except as these contain more elevated areas such as hummocks or ridges where nests can be placed. The very xeric communities such as the Dryas and lichenclad rocky fellfields of the hills and mountains are also avoided. Otherwise, in such widespread arctic and sub-arctic vegetation as sedge tussocks (Eriophorum), the meadow-like tundra of Carez, low, ankle-high areas of willow (Salix), and in expanses of low dwarfbirch (Betula) intermixed with various heaths, the Lapland longapur is the most numerous bird of the north.

The subspecies of Lapland longspurs are differentiated primarily on the basis of color. For the most part the differences are slight, as pointed out by the following brief review. F. G. Salomonsen (1950) believes the longspurs of Greenland, on the basis of a larger bill and paler color, are separable from those of Scandinavia and for these he utilizes the name 0. 1. ,subcalearatus (Brelim). He includes all the birds of Canada and Alaska within this subspecies and states: “The N. American population comes nearest to 0. 1. subealcaralus and is best referred to that form; 0. 1. alascertsis Ridgway is hardly worth recognition.” TI. Johansen (1958) adopts this system and states “Three population groups can be separated: lap pomicus in Eurasia, &ubcalcaratus in North-America and Greenland, and colaratus on the Commander Islands and Kamchatka.” Portenko (1960) believes that the birds from Knmchatka are subspeciflcaliy distinct and has given them the name learnt sckaticus. Manning et al. (1956) indicate that the birds of Baffin Island are even paler, and thus further removed from the nominate race than those of Greenland, although they prefer to leave the nomenclature unebanged pending a direct comparison of good series of specimens. They suggest that the failure of Salomonsen to recognize alaseert.sis might be due to his lack of specimens in fresh autumn plumage for the comparison that demonstrates their distinctness.

In general, a weak dine of increasing paleness extends from Greenland to the Aleutian Islands. Over this vast area, there is no convenient or realistic point at which to make a subspeciflc distinction and, indeed, allocation to subspecies of specimens from Banks Island (Manning et al. 1953) is difficult under the present system. It seems reasonable that Salomonsen and Johansen have made the best interpretation of the situation.

With respect to differences in habits or behavior it is apparent that these vary little or none at all among the different races. Familiarity with the extensive literature dealing with this species combine with my own field experience over a period of 9 years, including 3 years when I conducted an intensive study of populational phenomena at Cape Thompson, Alaska, confirm this belief. The bulk of the account which follows derives from my own study at the latter locality.

Spring: The wintering ground of the Alaska longspur includes the southern part of British Columbia and the bulk of the western United States south to New Mexico and Texas. Migration from this area in spring must begin in March and early April as longspurs appear as early as April 6 in southeastern Alaska (Grinnell, 1910). I have observed migrants at Anchorage, Alaska, as early as April 24, and at Homer, Alaska, on April 19.

The general course of travel appears to be northwestward through British Columbia and Yukon Territory and northward through the western Mackenzie Valley (Irving, 1961). There is also a coastal movement of birds whose arrival times are somewhat different from those of longspurs moving through the interior. Irving has taken latitude 500 N. and longitude 1340 XV. as representing the northwestern sector of the wintering area and points out that from that locality migration extends 200 north and 140 west to the eastern border of the breeding range in Mackenzie, while to the west in the Aleutian Islands, migration extends 670, but represents only a 40 northward advance. The average arrival time for four years at Atlin, British Columbia was April 22 (Swarth, 1936). This station is located in the path of large migratory flights moving westward to Alaska and from whence the general sequence of northward movement may be traced (Irving, 1961).

Arrival in western and northern Alaska varies from the last of April until mid-May with an average arrival date of about May 10. The consistent arrival in April in southeastern and south-cent~ral Alaska and the Aleutian Islands indicates a coastal migration independent of the major movements of birds through the interior.

The first flocks are entirely or preponderantly comprised of males, and these are followed in a few days by flocks that are preponderantly females. Arrival of the longspurs coincides with the rapidly retreating snow cover and the increasing availability of nesting habitat. Late snowfall, especially in the arctic, is not uncommon and the weather is characteristically unsettled when the birds appear.

Courtship: Courtship and pair formation take place after the birds have arrived on the nesting ground. Scattered males take up residency on the tundra and show signs of increasing territoriality. At this time, usually mid-May, small groups of males continue to remain together quietly feeding, and intermittent stormy weather may cause others to leave the newly established or forming territories and regroup temporarily in sheltered areas such as willow-clad stream bottoms. Singing on the wing, or less commonly from more elevated parts of the territory such as sedge tussocks, commences rather abruptly after the males arrive. Territorial disputes, including spirited chasing of neighboring males at the borders of the territories, and alternate singing with adjacent males are commonplace. Some of the chases are long-lived with the pursuing male seemingly within inches of the fleeing adversary, and maneuvering adroitly to remain there as they are lost from view.

Actual contact between rival males is not uncommon and one intruder was seen to land on the back of a silently feeding male, thus initiating a prolonged and erratic chase during which he was routed. On another occasion two males approached to within a foot of one another on the ground; both crouched down in a threat posture for a moment, whereupon the intruder flew off a short distance and the remaining bird resumed his feeding between the tussocks. The intruder returned promptly and struck the feeding bird. This initiated new threat posturing with the birds crouching low and spreading their wings slightly.

As the females continue to arrive in increasing numbers and disperse over the tundra, the attention of the males is attracted more and more toward them. Courtship flights are common, and two or three males are frequently involved in chases of an individual female. The male pursues the female in frequently lengthy and circuitous flights and finally lands a few feet from her. Common behavior during these pauses is for the male to peck at the ground and move closer to the female until she flies up to the top of a tussock thus initiating a new flight. His actions seem to indicate that he is physiologically predisposed to copulation and is so motivated by the female’s presence. When her response is not one soliciting copulation, his actions may be carried over into the displacement activity of pecking. Later, as the female becomes motivated toward breeding, probably largely due to the persistence of the chasing male, she may react by reciprocal chasing.

Drury (1961) states that:

Pursuit flights were obviously mutual affairs and, I believe, closely associated with pair formation. They were longer, slower, and with less zigzagging tban territorial skirmishes. If the male caught up, there was a burst of rapid zigzagging but if he fell far behind, the female slowed down until he overtook her. We saw these flights before any displays on the ground, and throughout the period of ground displaying. Male No. 6, displaying to his female, ran across in front of her or up to her side, standing at about a 600 angle to the horizontal, with his breast feathers fluffed out, head held high and bill pointed slightly up, wings about half spread, drooped and quivering. While he ran in this way he was singing his regular flight Bong. The female ran slowly ahead of him, crouched in a horizontal position with her head partly lowered, wings partly spread, tail cocked just above horizontal, calling zeep, zeep. * * * She and the male often pecked stiffly at the ground. * * * The chestnut nape emphasized the stiff bow by the male. The female ran around and ahead of him; then flew; if he did not follow, she came back and repeated her actions until he did chase. During the chases she landed several times, ran along the ground, then flew again, fast and darting, or slowly and on quivering wings. During the period of pursuits, because their attention was on the females, males sang less often and chiefly on the ground.

In the presence of a female the advertising and/or territorial song flight was seen to be modilled into a courtship display. The male performed this flight in the usual manner, with the tail spread and wings near the horizontal in a gliding path. The entire flight is only a few feet in length; it is performed only a short distance above the ground a few feet from the female, and the male often carries nest material, apparently sedge or grass, in his bill. The chases, both aerial and on the ground, together with the modified display flights wherein the male may carry nesting material, ultimately result in the female becoming receptive to copulation.

Drury (1961) describes similar behavior as precopulatory:

Above nest No. 3 * * * I saw the male rise with fluttering flight straight up into the air, flying with difficulty because of a wad of dark material in his bill * * * . Before flying, he stood vertically, bill horizontal or pointed up, wings drooped at his sides, and tail spread and lowered. He sang with this material in his bill and succeeded in getting about 20 feet off the ground, then sang as he fluttered down again. He picked up even more material, and hopped onto a stone. He had so much in his bill now that he could not get off the ground, but he could sing. He stood a few moments and readjusted the material in his bill, put it down and picked it up again; then he pointed his bill up and fluttered his wings without getting off the ground. * * *

When approaching the female the male held his head high, neck extending and bill horizontal, with all the accumulation of dark material in his bill. He dropped the load and pointed his bill straight up, showing his black throat, while he dragged his wings as he walked up to her. Then he lowered his head, ruffled his scapular, back and rump feathers, and widely spread his tail * * * . She crouched with head low, tail cocked up, wings quivering and partly spread. He fluttered over and, without hovering, mounted on her back in the same position he had assumed in front of her, and copulated. After copulation he hopped off and walked in front of her in the same crouched position, while she stood up, raised her bill straight up and cocked her tail as high as possible, chattering.

Belligerence of a male toward a female was witnessed on one occasion when pairs from adjoining territories came into contact. The male of one of these pairs assumed a crouching threat posture with wings slightly spread before the female of the other pair. No other aggressive action was taken.

On another occasion the quiet side-by-side foraging of a male and female was interrupted by a violent struggle between them. They chattered loudly and actually seized one another with their bills. The male routed the female and came away with a bill full of feathers. Whether or not these birds constituted a pair was not determined, but in any event the hostile, defensive motivation of the male was apparently released by the female’s proximity.

Toward the end of May, as early as the 26th, the intensity of song and other territorial behavior of the mated males begins to subside. This becomes marked by the end of May, and as early as June 3 mated males were seen to tolerate the presence of other males singing around and over their territories. By June 12 song and territoriality virtually cease. In late May and early June it is commonplace to see two, three, or less frequently four males following a single female as she forages about the territory. The resident male often tries to drive off these intruders, but apparently a diminished agressiveness in the face of unfavorable odds nullifies his efforts. One explanation for the presence of these seemingly surplus males is that they are unmated, but this implies an unusually disproportionate sex ratio not at all consonant with what is known of passerine birds generally, and when all longspur nests on a given area are censused, the number of males in the area, whether counted in small groups or singly, generally agrees with the nest count.

Coincident with incubation and the general subsidence of territorial behavior, the males are attracted to any female who happens to be away from a nest. Undoubtedly a few males and females are unmated, and these together with birds that have lost nests, may commence flocking by mid-June. Males collected from obvious small flocks in generally uninhabited areas had testes smaller than those of the breeding component of the population.

Nestirtg: A striking feature of arctic bird life generally is the sudden onset of nesting after the arrival of the migratory birds. The Lapland longspur is an example of a species that has compressed this interval to a minimum. With the arrival of the females, which may be as much as two weeks after that of the males, the nesting cycle is set rapidly in motion. The females start nest-building as promptly as four days later, and many records show nest-building spans just three days.

Edward W. Nelson’s (1887) comments on general nest structure indicate that considerable size variation is commonplace:

The walls are thick and strong, composed of an abundance of material, or they may be a mere cup-shaped shell, barely sufficient to hold the eggs. The majority of nests are composed of rather coarse grass, sometimes with moss interwoven, forming a thick layer, which was frequently as thoroughly watersoaked as a wet sponge when the nest was collected. The amount of material used depends greatly upon the locality; in damp places a much greater amount is made use of, while in dry places the nests are much lighter. Though the outer part of the nest was frequently formed of old and often grimy or partly decayed vegetable matter, the interior was invariably composed of fine, soft, dry, yellow blades of last year’s grasses. These in many instances are unmixed with other material, and in others are combined with feathers of the Ptarmigan or other wild fowl. In a few cases the lining of the nest is a warm cup of feathers resting upon fine grass, and one has a thick lining of feathers and dogs’ hair. Some nests are so small that they may he inclosed in the hand, while others can scarcely be inclosed by both, and the smallest nest collected may be inserted entire into the cavity of the largest one.

I feel that much of this size vanation can be attributed to the use of a nest from a previous year. Nest depressions, and undoubtedly the old material they contain, are used repeatedly. This material may be carried to a new site and made into a more compact nest, or it may be simply left in place, slightly modified, and then lined with fresh grass. Such a nest would be considerably larger both in bulk and size of the opening than a new one. The virtual impossibility of finding old, unoccupied nests suggests that the premium placed on particular nest locations is great and their reuse routine. Nevertheless, my examination of many nests failed to reveal variation in size of the magnitude Nelson indicates.

Further variation in lining, apparently depending on availability of materials, is indicated by the remarks of J. W. Bee (1958) who found nests lined with snowy owl feathers and caribou hair, and A. M. Bailey (1943) who also found a nest lined with caribou hair. The slightly more than 100 nests I have found were all lined with nothing but willow or rock ptarmigan feathers.

Herbert Brandt (1943) gives the measurements in inches for 21 nests as follows: height 3 to 7, outside diameter 4 to 6.5, inside diameter 3 to 3.5, and depth of cup 2.5 to 3.5.

The nests are nearly always placed in depressions in the sides of small hummocks of moss or sedge (Carex) or in the sides of tussocks of grass (Calamagrostis) or sedge (Eriophorum). Invariably some vegetation hangs over the nest, usually grass or sedge, or it may be hidden by the leafy twigs of willow (Salix) or various ericaceous plants. Nests are also placed in more level terrain such as Garex meadows or alluvial flats grown to low willows, usually in the drier parts, and well hidden beneath the vegetation.

The nest nearly always has one conspicuous entrance, which compass readings showed to be generally on the south side. The prevailing wind in the study area was from the north, but longspurs at other locations tended to orient the nest opening similarly, probably to take advantage of the greater insolation from that direction. A. V. Mikheev (1939), working in Russia, observed that the bowls of the nests were inclined to the east, south, or southeast, a phenomenon he attributed to the rarity of southeast winds during the nesting season. At Cape Thompson south winds are also uncommon, but nonetheless they are the ones associated with summer storms and precipitation.

The general habitat utilized for nesting has been described earlier in this paper. In summary, it can be said that longspurs are adaptable in their choice of nesting locations, requiring primarily only open terrain with concealing vegetation, either hillside or flatland, and there preferring the more xeric sites with microrelief such as tussocks, hummocks, and low ridges. The use of the term zeric here is decidedly relative. Truly xeric areas such as rocky hillsides are assiduously avoided, and the remainder of the northern landscape, all mesic by general terminology, is inhabited. Here, however, a distinction must be made between areas with actual standing water and those only moist underfoot. The latter are most densely occupied, and the preference for elevated sites is conspicuous. Thus the comment by many authors that the Lapland longspur requires and selects moist areas for nesting is misleading.

Eggs: The Lapland longspur usually lays five eggs although in my experience the mean number is somewhat less. In 44 nests I studied at Cape Thompson in 1960 the average clutch size was 4.7. Although slightly more than 50 percent were 5-egg clutches the number of eggs laid ranged from 1 to 6. The clutch of one egg, and another of only two, were dutifully incubated and apparently were not the result of eggs having been lost. In 1961 the average clutch size was 4.9 eggs in 64 nests, and the range was from 2 to 6; slightly more than 60 percent were 5-egg clutches. Herbert Brandt (1943) studied 28 nests at looper Bay and found an average clutch size of 5.5 with 16 nests containing 6 eggs and 12 containing S eggs.

It seems likely that had Brandt observed a larger number of nests this mean number would not have been so much higher (nearly one egg) than the average per clutch at Cape Thompson. Clutches of three or fewer eggs generally represent renesting attempts and perhaps should not be included in these computations. They do not necessarily represent late nestings, and no evidence was obtained to indicate that clutch size declined as the season advanced. True second nestings are apparently nonexistent, renesting is uncommon, and such variation as exists in number of eggs relates only to the single clutch laid by each female at very nearly the same time during the nesting period.

The shape, color, and size of the eggs are extremely variable as the following descriptions will indicate. The shape is ovate tending to elongate, to ovate and the surface is slightly glossy. The ground color is pale greenish-white, but on most eggs it appears to be “buffy brown” owing to the confluence of spots, blotches, and cloudings of “Hay’s brown,” “Saccardo’s umber,” “bister,” or “mummy brown.” This is not to imply a blotchy appearance, as generally the background color tends toward uniformity. Most eggs have scattered spots or scrawls of black. Undermarkings when visible are “light mouse gray,” and occasionally this is the predominant color of the egg, with only a few spots or scrawls of dark brown or black. The markings are fairly evenly distributed over the entire surface.

Gabrielson and Lincoln (1959) describe the eggs as “light greenishgray, heavily marked with shades of brown.”

Joseph Grinnell (1900) describes a clutch of eggs as nearly oblongovate in shape with a ground color, disclosed only at the ends of the eggs, as very pale blue. He states further that “Otherwise the eggs are so completely covered with pigment as to be almost uniform isabella-color. Overlying this are scattered scrawls and dots of bistre.”

Nelson (1887) described the ground color of the eggs as light clay with a pale greenish tinge. He found eggs covered with a coarse blotching of reddish brown, principally at the large end, some with a fine reddish brown specking, and others with varying types and intensities of marking including spots, spots and blotches, and irregular zigzag markings of dark umber brown. The markings become heavier and darker colored (chocolate brown) until the ground color may be entirely concealed, or revealed as only a faint mottling of olive: brownish.

The measurements of 40 eggs average 21.1 by 15.1 millimeters; the eggs showing the four extremes measure 22.9 by 15.3, 22.8 by 16.3, 20.8 by 13.7, and 19.3 by 14.2 millimeters.

Incubation: A few records indicate a one-day pause between the completion of the nest and the laying of the first egg, but usually the first egg is laid the day the nest is finished, and one is then laid each successive day until the clutch is complete. Effective incubation apparently begins with the 3rd or 4th egg in the usual 5-egg clutch. In some cases it may start with the second egg, but apparently almost never with the laying of the first egg. The variation in the time of onset of incubation is evidenced by the commonly observed spread of hatching over a 1- to 3-day period. Incubation is by the female alone and is of 12 or 13 days duration, measured between the laying and hatching of the last egg in the clutch. Data from 14 complete nest-histories showed eight 1 2-day incubation periods and six of 13 days.

As both Nelson (1887) and Brandt (1943) point out, the male stays nearby during incubation and utters alarm notes when danger threatens. Early during the incubation period the female’s usual reaction is to leave the nest when an intruder is still some distance away, fly 10 to 20 or more yards, alight often near the male, and begin foraging. Both birds may utter alarm calls, and the female may fly from tussock to tussock or other vantage points while watching the intruder. Usually within 5 or 10 minutes after the intruder retires, she returns to the nest in a series of short flights interspersed with brief periods of foraging and watching. The last abrupt ffight takes her directly to the nest; only once have I seen a female walk to the nest.

As the time of hatching approaches the attentiveness of the incubating female increases noticeably. She shows a marked reluctance to leave the nest and frequently remains on it until approached very closely. Then she flies only a few feet, often chattering loudly and plaintively. Alighting with wings slightly spread and drooped and tail fanned against the ground, she runs fluttering in this position away from the nest. The responses of individual females when forced from the nest vary considerably. Some may omit either the alarm notes or the distraction display, or both.

Clutches of four or more eggs take between 24 and 48 hours to hatch, and may take as long as 72 hours. While I have no definite evidence of longer hatching periods, a few of the nest histories from Cape Thompson suggest that it could have run to a maximum of 96 hours. Only one nest with 4 eggs was observed to hatch completely in 24 hours.

Young: The most striking single feature in the development of young longspurs is their early departure from the nest, well before the plumage develops enough to permit flight.

My records from Cape Thompson show that most young leave the next after 8 to 10 days, with an average nest life of 9 days. A few histories show departure as early as the 7th day and as late as the 11th. The young left two nests in 6 days, probably because the nests were disturbed. W. J. Maher (1964) reported the nestling period at Barrow, Alaska, averaged 7.4 days with a range of 6 to 8 days. This relatively brief time was undoubtedly influenced somewhat by nest disturbance. Maher reported thatthe average weight of longspurs at hatching is 2.3 gin. and increases to 18.8 gin. 7 days later. When the longspur is able to fly on about the 12th day it averages about 21.9 gin. or 80.5 per cent of its adult weight. Maher noted that the gaping response is weak on the day of hatching and grasping with the feet appears on the 2nd day. The fear response of huddling down in the nest appeared on the 5th day and escape was attempted by the 7th day.

My records show that the young usually leave the nest over a twoday period, although in some instances all young left the nest within 24 hours. In one case the departure of six young apparently spanned 3 to 4 days. It is not uncommon for one young bird to remain in the nest a day or two longer than his nestinates, especially in the larger broods. This might be due to later hatching and/or to difficulty in getting a proportionate share of the food, for disparity in nestling size is commonplace.

The sexes share equally in the feeding of the young birds. After leaving the nest the brood is often split and each adult cares for a portion of the young. Maher (1964) inclosed nests with a fence and observed that the young begin to fly on the 12th day. At Cape Thompson fledging success was 1.68 young per pair of adults in 38 nests in 1960 and 1.9 in 53 nests in 1961.

Density: The following figures may bear out the belief expressed earlier that the Lapland longspur is very likely the most numerous bird in the north. At Cape Thompson the density per 100 acres was 14 pairs in low riparian willows, and up to 59 and 65 pairs in the widespread plant formations of Eriophorum tussocks and Carex meadow. Extrapolating these figures for larger areas of generally uniform habitat, such as Eriophorum tussock tundra, yields such total population figures as 11,800 breeding birds in 10,000 acres. With 1.9 young reared per pair, the total number of fledglings and adults present on this expanse in mid to late summer is about 22,200. Eriophorum tundra is the dominant plant communtiy not only of the Cape Thompson area, but of the entire foothill region of the arctic slope of Alaska (Spetzman, 1959). The lowland areas that are largely covered with Cares meadows may support even greater population densities.

Plumages: Tbe natal down is basally light tan in color, shading to yellow and gray for the distal third (Maher, 1964). Dwight (1900) states that the juvenal plumage is acquired by a complete post: natal molt. Maher shows that the nestlings appear feathered by the 7th day and can fly by the 12th day, although the primaries ~re still growing. The juvenal plumage is described by Gabrielson and Lincoln (1959) and by Dwight (1900) generally as follows: upper parts and sides of head rich buff, tawny buff, or clay-colored, streaked heavily with black; wings and tail deep clove brown; tertiaries and greater coverts edged with “Mars brown” and white-tipped; lesser coverts edged with white, primaries and tail with pale cinnamon; outer rectrices terminally bulfy white; below dull white, washed with buff across the throat; the chin, throat, chest, and sides streaked with black.

The postjuvenal molt begins about 3 weeks after fledging and involves mainly the body plumage and part of the wing coverts. It coincides generally with the timing of the complete molt of the adults. At its end the adults and immatures become, for all practical purposes, indistinguishable.

Dwight (1900) describes the first winter plumage of the male as follows:

Above, wood-brown and cinnamon streaked with clove-browo, the nape and sides of neck chestnut concealed by wood-brown edgings; lesser coverts edged with wood-brown. Median crown stripe, superciliary line, and anterior auriculars buff, posterior auriculars black. Below, white, the feathers everywhere dusky basally, the sides of chin and a crescentic area on the throat jet black veiled almost completely by long white edgings; the sides and flanks streaked with black.

Dwight further states that the first nuptial plumage is acquired by a partial prenuptial moult beginning in March in the United States which involves the anterior parts of the head, chin and throat. The black feathers of these areas and the creamy white ones of the sides of the head are acquired by moult contrasting with the chestnut collar which is assumed by loss of feather edgings. This moult does not usually extend to the posterior portion of the black throat patch where old black feathers with partly worn-off edgings are regularly found. Wear produces a distinctly black and white streaked appearance above with the collar clear chestnut as if unveiled.

The adult winter plumage is acquired by a complete post nuptial molt in July and August. Dwight notes it as “practically indistinguishable in many cases from the first winter plumage, but the black on the chin and throat is more extensive, and the colors richer and deeper, especially the wing edgings.” At Cape Thompson the complete molt begins during the last part of June and is essentially complete in all members of the population by the end of August. Timing of molt is similar for both sexes and takes about 50 days to complete. The earliest dates for inception of the annual molt of males and females were June 25 and July 2, respectively, while molt started as late as July ~ in males and July ii in females. Post: juvenal molt was observed to begin as early as July 10. The striking degree of synchrony manifest in the timing of molt accords with the brevity of the arctic summer and the short time available for this activity.

The adult nuptial plumage of the male is acquired by a partial prenuptial molt as in the young birds. Gabrielson and Lincoln (1959) describe this plumage as follows:

Head and chest deep black, relieved by a broad white or buffy stripe behind eye, continued downward (vertically) behind ear coverts and then backward along sides of chest; ground color of upperparts light grayish brown, with little if any rusty tinge, even on wings; sides narrowly streaked or striped with black; rest of underparts white; hindneck deep chestnut rufous; lesser wing coverts grayish, feathers black in center. Outer tail feathers white, dusky along midrib and on inner web toward base; the tail feathers narrowly edged with grayish white; lining of wing and axillaries grayish white. Bill dark, lighter at base; legs and feet black; iris brown.

The plumages and molts of the female according to Dwight (1900) correspond to those of the male, but the black throat patch is never so extensive and usually merely outlined with dull black streaks. The juvenal plumage is indistinguishable from that of the male. The first winter plumage is much veiled and streaked above with clove and cinnamon brown, the nape vinaceous; below it is white obscurely black on the sides of the chin and with a small throat patch, the sides and flanks black streaked. The first nuptial plumage is chiefly the result of wear, a few white feathers being acquired by moult on the chin.

Gabrielson and Lincoln (1959) describe the plumage of the adult breeding female as follows: “A dull-colored bird, striped above with dull black, rufous, and grayish white; an indistinct median light line on crown; collar on hindneck pale rufous, finely streaked with dusky; feathers of throat and breast black, concealed in part by grayish white and pale buffy margins of feathers, producing a mottled or streaked appearance; sides and flanks striped with dull black and pale rufous; rest of underparts white; tail and wings as in male.” The adult winter plumage of the female is like that of the first winter with perhaps more black on the throat.

H. S. Swarth (1934) records colors of the soft parts quite accurately, noting that in the young male the upper mandible is yellowish brown with pale yellow edges, the lower mandible a grayish flesh color, and the tarsus and toes brownish flesh color. The adults in summer have a lemon-yellow bill with a black tip, the iris is brown, and the tarsus and toes are dark brown.

Food: Gabrielson (1924) made an exhaustive examination of 656 stomachs, including 113 previously reported on by S. D. Judd, taken from birds collected in Alaska, five provinces in Canada, and 15 states of the United States, during every month of the year. He classifies 56 stomachs from Alaska and northern Canada, June to September inclusive, as representing “summer,” with the remainder being “winter.” Beetles, mostly chrysomelids and weevils, constituted 11.91 percent of the summer food. Fly remains, almost entirely the eggs and adults of crane-flies (Tipulidae) constituted 17.77 percent. Caterpillars, spiders, bugs, and fragments of other insects accounted for 17.75 percent. Total animal food was thus 47.43 percent of all food. Grass seeds formed 12.02 percent, seeds of sedges 4.16 percent, seeds of a variety of other plants 27.86 percent. Bits of grass and unidentified vegetable debris formed the remaining 8.53 percent.

These foods differ little from those observed through study of the contents of 90 stomachs taken at Cape Thompson during the summer of 1960. Animal and plant food were taken in approximately equal amounts with only a slight preference shown for the former. Important animal groups selected in order of occurrence in the stomachs were beetles (Coleoptera), flies (Diptera), wasps (Hymenoptera), bugs (Homoptera), and spiders (Araneae).

Gabrielson points out that over half of the 600 “winter” stomachs were taken in Kansas under very similar conditions. They contained largely millet and crabgrass seeds; animal food comprised only 3.97 percent, and four birds that had fed almost exclusively on carabid beetles of the genera Platynus and Amara accounted for half this total. Other animal food was chrysomelids, weevils, fly larvae, caterpillars, and spiders taken in varying quantities in every month except February. Goose-grass (Eleusine) was an important item, as were also sedge seeds. Seeds of pursiane (Port ulaca) were common but, owing to small size, made up only 0.82 percent of the bulk. Seeds of pigweed (Amaranthus) comprised 6.03 percent. Goose-foot (Chenopodium) and ragweed (Ambrosia) seeds were favored. Wheat formed 8.33 percent of the food and was eaten in every one of the eight months except October.

At Cape Thompson the young appeared to be fed animal food exclusively. This was also the observation of Rowell (1956) who noted that young longspurs were fed mainly adult Diptera, particularly culicids and chironomids, and were also given some beetles and caterpillars.

During 1960 the adults showed no dear pattern of food preference with the changing season. For approximately two weeks during late May and early June animal foods were taken to the near exclusion of plant foods, while prior to and following that period nearly equal amounts of both were eaten. The important point is that this arctic species is markedly adaptable in its diet and, depending on the day to day availability of certain items, readily switches to the most easily acquired foods.

G. W. Rawson (1954) comments on feeding behavior of this species in the Brooks Range of northern Alaska as follows:

Mosquitoes collected in large numbers on the outside of a white canvas tent two of us were using as living quarters. Several small flocks of immature Alaska Longspurs soon “caught on” to the fact that the tent acted as an ideal trap or concentration camp for mosquitoes and diligently fed on them both day and night. At first (before knowing what was causing the disturbance) the longspurs were quite perturbing because one or more would lose their footing and slide down the inclined roof of the tent causing a ripping sound suggestive of someone or something trying to rip open the canvas.

More commonly observed feeding behavior consists of constant walking about quietly concealed in such vegetation as sedge tussocks, often seemingly quite oblivious to the observer. Extremely strong winds seem to have no effect on the birds’ ability to find insects, and the young are fed just as constantly during gales that make flight difficult much above the ground.

Behavior: The general pattern of breeding behavior as detailed in the preceding sections, together with other facets of the annual cycle of the Lapland longspur, can be briefly reviewed as follows:

After a winter spent in flocks on the open country of the western and mid-western United States the birds begin the spring migration north to the breeding grounds of Alaska, Yukon Territory, and the Northwest Territories in March and April.

At Cape Thompson, in northwestern Alaska, the males arrived as early as May 5th, an event marked by the abrupt onset of song and display. Territorial disputes become commonplace, and defensive behavior involves chases, sometimes with actual physical contact, and threat posturing. Singing on the wing or from elevated points in the area seems virtually incessant. In less suitable habitat a male may defend an area roughly seven acres in size, while in favorable vegetation the territory is reduced to approximately two acres.

The females arrive as early as May 18. The attention of the males is turned toward them, courtship flights begin, and copulation may be observed as early as May 21. The end of May sees a marked subsidence of territorial behavior. Nest-building begins as early as May 23 and takes three days; only the female performs this task. The eggs are laid on successive days immediately following the completion of the nest. The average clutch consists of 4.? to 4.9 eggs and these are incubated for 12 days. The young birds remain in the nest for an average of 9 days and begin flying at the age of 12 days. Molt is initiated promptly in late June and early July, and the birds gather in small flocks by mid-July.

The longspur flocks feed quietly on the ground and may go unnoticed until approached closely. Gabrielson and Lincoln (1959) describe this flocking behavior very well:

After the breeding season, they gather in small groups, which gradually unite into flocks that sometimes consist of thousands of individuals by the time they reach their wintering grounds in the northern States. These flocks have the characteristic habit of taking flight when alarmed, flying in great sweeping circles, and then returning to alight on the same spot or very close to it. At other times, however, they may take off and go straight away, completely out of sight, before alighting again.

As the birds move morthward in spring the behavior differs somewhat as the compact flocks pause only to feed for a short time before moving on.

These aggregations take flight readily and seem unusually wary when contrasted with the tameness of the individuals scattered on their nesting territories. Their behavior is frequently like that Grinnell (1900) observed at Kotzebue Sound: “I would frequently meet with a male longspur standing motionless on some conspicuous hummock. If I approached too close he would attempt to get out of my way by stealthily running to one side, but if pressed he would take flight and mount upwards, circling high overhead and uttering his pleasing song.” I also have noted that territorial males pressed into flight often respond by initiating the flight song.

Voice: The varied vocalizations of this species include the conspicuous song of the breeding male, call notes, the alarm notes of both sexes, the alarm notes of the female when frightened from the nest, and flocking notes used during the non-breeding season.

Nelson (1887) writes vividly of the song of the male:

The males, as if conscious of their handsome plumage, choose the tops of the only breaks in the monotonous level, which are small rounded knolls and tussocks. The male utters its song as it flies upward from one of these knolls and when it reaches the height of 10 or iS yards it extends the points of its wings upwards, forming a large V-shaped figure, and floats gently to the ground, uttering, as it slowly sinks, its liquid tones, which fall in tinkling succession upon the ear, and are perhaps the sweetest notes that one hears during the entire spring-time in these regions. It is an exquisite jingling melody, having much less power than that of the Bobolink, but with the same general character, and, though shorter, it has even more melody that the song of that well-known bird.

This quotation is slightly misleading in that the bird seldom begins the song during the preliminary, upward flight.

The behavior of singing birds I witnessed at Cape Thompson did not differ from that Drury recorded (1961) on Bylot Island, which is briefly as follows: males occasionally sing from conspicuous perches on the ground, but the usual song is given in a flight during which the male rises to a height of 20 to 40 feet and then settles slowly to the ground singing several songs in rapid succession. During windy periods the birds often sing several songs while hovering overhead. Drury notes that “Songs started with a ringing and metallic ring, followed by a rolling and rapidly descending zizeleeaw; then a rolling, sustained zizelee-ee (ending with the highest note of the song ee), and closed with another rapid rolling and descending zizeleeaw.” Drury also uses the syllables see, serilee-aw, serilee-ee, .serileeaw to describe the rapidly descending flight song, and these seem to me excellent word representations of the sounds. He also reports a “whisper” song containing the same elements as the full song, but usually shorter, and given only by the male when in close company with the female before the start of nest-building. This I never heard at Cape Thompson where, over a two-year span, full song began as early as May 5 and continued as late as June 28.

The peak of song seems to occur during the period of greatest territorial activity, prior to the laying of eggs. Singing diminishes conspicuously after the female begins incubating. The last songs of the season, heard on July 21 and again on August 12, were weak and abbreviated.

The alarm note sounds to me like a plaintive peer and is given near the nest or elsewhere on the territory when the birds are disturbed. The same call is given by members of a flock when they are pressed into flight. Salomonsen’s (1950) descriptions of these and other notes for the birds on Greenland are applicable as well to the Alaskan members of the species and are generally more accurate representations than other published versions. He states:

The common call-note, used at the breeding-place, is a Boft melodious ee-yu or only yfi or chflp, uttered by both sexes on the ground and during flight, but more slurred and not so frequently by the female. Another note is a bunting-like dir-nt, uttered also on ground and in flight, used as a contact-note, during flight often as a lengthened, chirping dinninrinnirnit. A short and more harsh pinrh uttered by the male on the wing is sometimes heard, as well as an exited sparrowlike note during pursuing-flight. The call-note and the contact-note are often combined during flight, alternating like dit-ii-dit-ti. Birds moving about in flocks in the autumn use chiip when alighting, when foraging and at rest, dir-nit wben rising and in flight.

Almost as memorable as the full song of the male are the alarm notes sometimes given by the female when she is startled from the nesI~ late in incubation. These and the associated display are described by Nelson (1887): “In one instance a parent was driven from her eggs just as they were about to hatch, and she ran along the ground for a few yards, uttering a plaintive note, like ch~ ch~ ch~, in a fine, vibrating, metallic tone, at the same time dragging her outspread wings and tail upon the ground, and fluttering as though in mortal agony.~~ Field marks : P. A. Taverner (1934) writes:

The adult male has the throat black like the face, instead of white as in McCown’s, or buffy as in Smith’s and the Chestnut-collared Longapurs. Harris’s Sparrow has a similar black face and bib, but is otherwise an entirely differentappearing bird, with light grey ear-coverts, and no chestnut collar. Females and juveniles with the distinct or semi-obscured chestnut collar are easily separated from MeCown’s and Smith’s, but may be very similar to the Chestnut-collared. They are distinctly larger birds, however, wing 3.50 and over instead of 3.25 or under, and the collar is well developed instead of vaguely defined or absent. The whole bird is more sharply streaked. The black suffuses around ear-coverts and across the lower neck, and the underparts are solid white. Female and juvenile Chestnut-collared Longspurs may have a veiled black spot below a light throat, but the abdomen is a dusty buff y and usually shows more or less irregular intrusion of black. Autumn juveniles are still more confusing. They have a general appearance of a streaked buff y bird, with white, rarely cream, abdomen, with but traces of veiled black down sides of throat from corners of bill, and across upper breast. The tips of the ear-coverts are bordered by a conspicuous brown or black patch that is absent in the Chestnut-collared and MeCown’s Longspurs and much smaller or absent in Smith’s. The best test for the species, in this plumage, is the white or faintly cream abdomen. * * * In addition to details previously mentioned, other plumages are more streaky than other longspurs, and never as evenly buffy as Smith’s. * * * The subspecific distinction is slight.

R. T. Peterson (1941) notes that the spread tail is predominently black with white edges, whereas the amount of white in the tail of MeCown’s and the chestnut-collared longspurs exceeds the amount of black. The flight is undulating.

T. S. Roberts (1932) points out that female or autumn birds can be confused with vesper sparrows, but the longspurs have different tail patterns, customarily occur in flocks, and have different call notes. He also says they “resemble the House Sparrow more dosely than anything else, but habits and notes are very different.” Further, the “bird hops or runs instead of walks.”

Additional features brought out by H. F. Witherby et al. (1938) are the absence of white in the wing, the comparatively thick upper mandible, and the long, lark-like, hind claw.

Enemies: Observations at Cape Thompson indicated that the greatest losses of nests were caused by predation, which accounted for approximately 77 percent of the losses over a 2-year period. Arctic foxes (Alopex lago pus), ground squirrels (Citelius ‘undulatus), and weasels (Mustela erminsa, Musteta rixosa) were the predators believed mainly responsible, although the activities of the ground squirrels were restricted entirely to the gravel-covered stream bottoms where longspurs were relatively scarce.

After the young leave the nest they are particularly vulnerable to avian predation, and at Cape Thompson two species of jaegers, the parasitic and the long-tailed, feed on them regularly. James W. Bee (1958) reported that he observed short-eared owls and pigeon hawks preying regularly on longspurs in the vicinity of Point Barrow, Alaska. I watched a northern shrike capture and kill a young longspur in Mt. McKinley National Park.

Heavy precipitation at the time of hatching caused many nest losses at Cape Thompson. In 1961 rain associated with wind and low temperatures fell June 14 to June 21 following the peak of hatching on June 13. In the 66 nests under observation, 10 nest losses resulted from this storm, while only one before and one after that period were attributed to weather. Considering the relatively small number of nests I was able to observe, the loss must have been very great on a regional basis. At such times the females brood the young effectively for a while, but nevertheless must leave to feed, and frequently the resulting exposure is sufficient on a cumulative basis to cause losses of young. By contrast, in 1960 with 46 nests being watched, only one nest loss was attributed to the heavy rain that fell June 19 to 27, apparently because the young birds were by then sufficiently well feathered to endure the sporadic absence of the female.

Stormy weather is a well-known peril for migrating passerine birds, and the Lapland longspur is no exception. T. S. Roberts (1932) describes a spectacular mass kill in Minnesota during a March snowstorm in 1904:

On two small lakes, with an aggregate area of about two square miles, the ire was still intact and nearly bare from the melting snow. This exposed surface was thickly and evenly strewn with dead longspurs. By measuring a number of squares, counting the birds in each, and averaging these counts, it was possible to make a fair estimate of the number of bodies on the whole wee. A conservative calculation showed that there were at least 750,000 dead longspurs lying on these two lakes alone! The adjoining uplands, the streets of the town, and the roofs of the buildings were strewn with bodies in equal numbers. And this was only one locality in the extensive area throughout which the birds were killed.

Roberts adds that the total area involved was found to be approximately 1,500 square miles. This mass mortality had no noticeable effect on the numbers of migrating longspurs in succeeding years. With respect to such winter kills Lincoln (1939) writes “Almost every winter brings reports of storm destruction of thousands somewhere in the Middle ‘West until it seems amazing that the species is able to survive * * * catastrophes have been reported from eastern Colorado, Nebraska, and North Dakota.”

Winter: While much of the available literature on wintering Lapland longspurs cannot be referred to a specific subspecies, it is well known that eastern and western forms overlap broadly in the mid-western prairies where they occur in great numbers. Nearly every author mentions the great abundance of these birds in winter. A. L. Goodrich, Jr. (1945) remarks that in Kansas “They frequent the wind-swept plains and prairies, occurring in large flocks. Skimming or running over the ground, they range widely in search of seeds and berries of available grasses, weeds and low plants.” W. W. Cooke (1897) st.ates that in Colorado “When it first arrives it passes up into the lower mountain parks, but in severe weather it is confined to the plains extending to southern Colorado.” The species apparently moves rather extensively in winter, opportunistically seeking localities where the food supply can sustain the very large numbers present. This seems especially true in the northern perimeter of the winter range (latitude 500 N.) where they may occur in abundance in some years and only uncommonly in others.

DISTRIBUTION

Range: Alaska, northern Yukon, and northwestern Mackenzie south to Oregon, Nevada, Utah, Arizona, New Mexico, and northern Texas.

Breeding range: The Alaska lapland longspur breeds from southwestern, western, and northern Alaska (Aleutian, Shumagin, Pribiof, Nunivak, and St. Lawrence islands; St. Michael, Cape Lisburne, Barrow) to northern Yukon (Herschel Island) and northwestern Mackenzie (Mackenzie Delta, Rendezvous Lake); recorded in summer farther south in Alaska (Semidi Islands, Fort Kenai) and in Mackenzie (Fort Franklin).

Winter range: Winters from southern British Columbia (Lulu Island, Okanagan Landing), northwestern and central Montana (Fortine; Custer County), southwestern South Dakota (Rapid City), central northern Nebraska (Wood Lake), and northea.st.ern Kansas (Hamilton and Douglas counties) south to California (rarely, Litchfield, Death Valley), central Utah (Toocle; Duchesne County), Arizona (rarely, Imperial Dam, Meteor Crater), east-central New Mexico (Picacho), and northern Texas (Canyon); casually west to western California (Eureka, False Bay) and east to Tennessee (Memphis) and Ohio (Columbus).

Egg dates: Alaska: 159 records, May 23 to August 1; 90 records. June 5 to June 22; 112 records May 25 to July 7.

Mackenzie: 12 records, June 14 to July 1.