The Horned Lark is a bird of open country with short vegetation, and it occurs over a very large range worldwide. Like many songbirds in this habitat it has flight songs which are used to delineate territories and attract mates. Winter flocks of Horned Larks can number in the hundreds.
Female Horned Larks dig a nest cavity from the soil, or find natural depressions in which to place their nests. She also lays a sidewalk of sorts leading up to the nest, though she does not always use it when approaching the nest.
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Description of the Horned Lark
The Horned Lark has sandy brown upperparts, a white belly, a yellow throat and black bib, and a black patch below each eye. It also has two small black tufts, or “horns” that are not always visible. Different subspecies vary somewhat in the coloration of the throat and upperparts. In flight, Horned Larks show mostly black tails with narrow white edgings. Length: 7 in. Wingspan: 12 in.
Males are brighter than females.
Females are similar to males, but duller in color.
Seasonal change in appearance
Juveniles lack the yellow and black head pattern, and have dark upperparts heavily marked with pale mottling.
Horned Larks inhabit fields, prairies and airports with short vegetation, shorelines, and tundra.
Horned Larks primarily eat seeds and insects.
Horned Larks forage by running or walking, and gleaning food from the ground.
Horned Larks have a holarctic distribution, including breeding across nearly all of North America, and wintering across most of the U.S. and Mexico. The population has declined in recent decades, after a large increase following deforestation in the late 1880s.
Bent Life History
Visit the Bent Life History for extensive additional information on the Horned Lark.
The shape of a bird’s wing is often an indication of its habits and behavior. Fast flying birds have long, pointed wings. Soaring birds have long, broad wings. Different songbirds will have a slightly different wing shape. Some species look so much alike (Empidonax flycatchers) that scientists sometimes use the length of specific feathers to confirm a species’ identification.
- Male, arctica, Washington, Mar.
- From below
- Male, merrilli, Washington, Mar.
- From below
- Female, merrilli, Washington, May
- From below
- Female, juvenile, merrilli, Washington, July
- From below
Wing images from the University of Puget Sound, Slater Museum of Natural History
There are about 21 described subspecies of Horned Larks in North America.
Horned Larks are sometimes seen in mixed species flocks with longspurs.
The typical song consists of a few chirps followed by rapid tinkling calls.
- Sparrows and longspurs lack yellow throats.Dickcissel
Dickcissels do not have “horns” or the black facial marking of the Horned Lark.
The Horned Lark’s nest is depression in the ground lined with grass and other fine materials.
Number: Usually lay 2-5 eggs.
Color: Greenish-white with darker markings.
Incubation and fledging:
The young hatch at about 11 days and fledge at about 10 days, though remaining dependent on the adults for another week.
Bent Life History of the Horned Lark
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 Horned Lark – the common name and sub-species reflect the nomenclature in use at the time the description was written.
OTOCORIS ALPESTRIS ARCTICOLA Oberholser
This large, pale race of the horned larks is about the size of alpestris aZpe8tms, but is decidedly paler. Dr. Oberholser (1902) says of it: “This form is one of the best marked of all of the races of Otocori~ alpe.stris, differing from the typical subspecies in its very much paler upper surface, more pinkish nape, upper tail-coverts and bend of wing, as well as in the pure white of throat and eyebrow. * * This is the race to which, through misapprehension of the identity of Dr. Coues’ type, the name leucolaema has, by common consent, been applied. Examination of the rediscovered type, however, proves it to belong to another race, as fully explained under its proper heading, and leaves the present subspecies without a name.”
At the time Dr. Oberholser described it, its breeding range was supposed to be confined to Alaska, “(chiefly the interior), with the valley of the Upper Yukon River.” Since then it has been found, apparently breeding, above timberline on several of the interior mountain ranges of Alaska, and in Alpine-Arctic regions in the mountains as far south as Washington.
In the Stikine River region of southeastern Alaska and northern British Columbia, Harry S. Swarth (1922) reports it as “seen in small numbers on the mountain tops above Doch-da-on Creek. There, on July 11 and again on July 23, they were found on the open, mosscovered slopes above timber line, associated with rosy finches and pipits.”
It remained for Dr. Walter P. Taylor (1925) to extend the known breeding range of this lark to a point much farther south than any of the localities in British Columbia from which it had previously been reported. He writes:
During field work in the State of Washington in 1919 we found the subspecies breeding well south of the international boundary line. On August 5 of that year a small Horned Lark, as yet unable to fly, was captured at an altitude of 7,300 feet on Panhandle Gap, Mount Rainier. The locality is well above timberline, in the AlpIne-Arctic Zone, and a favorite resort for Ptnrmlgan and mountain goat. The so-called Gap is in reality a broad ridge, to the north dropping off abruptly to the Sarvent Glaciers, on the south sloping gently to Ohanapecosh River. Although the date of capture of the young bird (August 5) seems a Uttle late, at least for localities at lesser altitudes, the season on Panhandle Gap was at its heIght, and the ground, only recently uncovered by the snow, was blanketed with grass and flowers. On being pIcked up the young Horned Lark disgorged three locust-like insects and a small green worm.
The mother remained close at hand while we watched the young bird, uttering a solicitous call-note resembling chivew, cl&ipew.
Specimens taken in this and neighboring localities proved to be typical of areticola. He sums up the status of the pallid homed lark in Washington as follows: “The subspecies occurs as a common migrant and breeding bird at least from July to September in the Alpine-Arctic Zone of the Cascade Mountains south at least to Mount Rainier (Taylor); east to Chopaka Mountain (Taylor); and west to Mount -Baker (J. M. Edson); in winter, as early as November and probably to March, it is found in the lowlands of eastern Washington, north and east to Cheney, south to Walla Walla (Lyman); and west to Moses Lake (Cantwell) and Benton County (Decker); it is of accidental occurrence, during migration, in western Washington (A. K. Fisher).”
I cannot find any information on the nesting of this subspecies except the following brief statement by Maj. Allan Brooks (1909): “Mr. [C. de B.] Green this year took the eggs of the Pallid Horned Lark on the high mountains above timber line, between the Okanagan and Similkameen valleys and collected the female, which is now in my collection. This is the breeding form on all the high mountains of the Province, Otocoris a. merrilli being restricted to the arid lower levels; nowhere do their breeding ranges impinge on each other.”
I do not know what became of the eggs collected by Mr. Green, who is not now living.
Probably the nesting habits of this race and its eggs are not materially different from those of alpestris or hoyti.
The measurements of three eggs from the Pearson Mountains, British Columbia, are 23.7 by 16.5, 23.2 by 16.1, and 23.3 by 15.5 millimeters; three eggs from the Ashuola Mountains, Wash., measure 24.1 by 16.5, 22.8 by 16.5, and 22.8 by 16.0 millimeters; these are well within the limiting measurements of eggs of the northern horned lark and, in fact, average about the same in size.
The plumage changes of the pallid horned lark are apparently similar to those of the other races of the species. Mr. Swarth (1922) writes: “Two adult males, taken July 23, are beginning the annual molt, shown mostly in the wing coverts. The young bird, taken July 23, is in juvenal plumage throughout. Compared with the young of various southwestern subspecies of Otocoris alpestris, it is extremely dark colored. Ground color of the upper parts is blackish, throat and lower belly are white, and there is hardly a trace of rufous or vinaceous anywhere.”
The food and general habits of this race are probably similar to those of the other northern subspecies, with due allowance for the difference in its alpine habitat.
HOYT’S HORNED LARK
OTOCORIS ALPESTRIS HOYTI Bishop
Dr. Louis B. Bishop (1896) named this bird in honor of his friend, William H. Hoyt, of Stamford, Conn., and described it as “similar to Otoco~d.s alpestris but with the upper parts generally paler and~ more gray, the posterior auriculars gray rather than brown, and the yellow of the head and neck replaced by white, excepting the forehead, which is dirty yellowish-white, and the throat, which is distinctly yellow, most pronounced toward the center.”
The breeding range is generally understood to extend from the west shore of Hudson Bay westward to the valley of the Mackenzie River, northward to the Arctic coast, and southward to at least Lake Athabaska. Just where it intergrades with arcti~ola on the west, or with leucolaema and praticola on the south, does not seem to be definitely known. It is the central one of the three northern races, occupying most of central-northern Canada. This is just where we might expect to find an intermediate form. Reference to the comparative descriptions of the three races by Oberholser (1902) and Ridgway (1907) will convince the reader that it is strictly intermediate in all of its characters between the dark alpe8tris and the pale arcticola. The wisdom of describing and naming an intermediate race seems open to question, as it immediately produces two more sets of intermediates. Such intermediate forms seem to occur in regions north of Hudson Bay, such as Baffin Island and Southampton Island. J. D. Soper (1928) collected 36 specimens ~n Baffin Island, of which he says:
The great majority are typical 0. a. iwyti, are as large as alpestris, but have white eyebrows and much white on face and sides of neck. Five specimens from Nettihing lake represent birds found associating with the typical 0. a. liovti. These five, if not typical breeding alpestris, are much nearer to that race than to l&opt~. Amongst the specimens are several that are Intermediate between these extremes. A male specimen taken at Nettilling lake, June 25, has a pure white Instead of well-marked yellow, throat and seems indistinguishable from typical 0. a. arcticola. A few other birds have white feathers In mosaic pattern, over the yellow throat, suggesting a mixture of bloods rather than a fortuitous development of white feathers.
Referring to the above remarks, Dr. George M. Sutton (1932) says: “I think Mr. Soper is wrong in calling ‘the great majority’ of this series hoiyti, or in referring to them as having white eyebrows and white faces. In looking at these birds in an off-hand way, their faces do appear to be rather pale; but when compared with the Southampton breeding birds, the decidedly yellow face and heavy, 8tUb bed bill show up immediately; and furthermore, when compared with a series of seven breeding birds from the Labrador coast (supposedly alpestri8) they do not appear to be in any way greatly dissimilar from them.” Referring to Southampton Island, he says:
Hoyt’s Horned Lark is a common summer resident in the region of South Bay. It is not so common farther west, at Capes Low and Kendall, and I do not know whether it occurs at all in the extreme eastern, higher part, where its place may be altogether taken by alpestri8. Unfortunately no summer collecting was carried on about East Bay and Seahorse Point, so I do not know which race breeds there. Alpestrie, apparently, Is the only form which occurs there as a migrant.
Hoyt’s Horned Lark arrIves in the spring a little later than the Snow Bunting and Lapland Longspur, and departs somewhat earlier and more definitely in the fall. It has never, to the best of my knowledge, been recorded in winter.
Frank L. Farley tells me that both alpestris and hoajti are common summer residents at Churchill, Manitoba.
Spring: Referring to the spring passage of the pallid and Hoyt’s horned larks at Aweme, Manitoba, Stuart and Norman Griddle (1917) write:
They usually arrive within a few days of each other and with the Lapland Longspurs in large flocks about April 6. Soon the ploughed fields are swarming with them and their value as destroyers of noxious weed seeds must be considerable. * * It is an interesting sight to see these birds, in company with thousands of Longspurs, circling for miles around some large hawk, though their object in doing so is a mystery and seems to be almost ignored by the hawk. Their music, as they fly around in milllons, fills the air, producing an effect which is long remembered. Both Horned Larks and Lapland Longspurs may also be seen to rise some 30 feet, uttering as they drop a short song. It is evident, however, that this is oiily a prelude to what is to come when the birds reach their true homes.
Taverner a.nd Sutton (1934) found these larks very numerous when they arrived at Churchill, Manitoba, on May 28. “They were everywhere, feeding confidingly with Snow Buntings even about the doorsteps of the offices and workshops of the townsite. They sang more persistently and finely than we had ever heard them before. The male of a pair nesting close to our Churchill camp habitually perched on the ridge-pole of the tent and sang continuously for many minutes, deserting his post only for momentary feedings or when he flew to the adjoining tennis-court, where he continued to sing. About June 10 the species became less noticeable about the door-yards, but continued abundant all over the tundra.” Both races, alpestri.s and /ioyti, were present on their arrival, but “most of the yellow-faced birds left with the transients.”
Nesting: Dr. Sutton (1932) writes: “The female alone builds the nest, and performs all the duties of incubation. While the male occasionally brings her food, there are regular periods of the day when she leaves the nest to preen, bathe, and feed. The nest is usually built in the open quite a way from water, often on a sloping plain or plateau, and not often on the highest part of a ridge. It is placed in a cup-like depression in the tundra. It is made of stalks of weeds, grasses, and small leaves, lined with soft vegetable material, especially the tassels of ‘bog cotton,’ and the phunous pappi of some of the flowering plants. I did not note any ‘pavements’ near nests.”
A nest found by Mr. Soper (1928) “was located in low, tundra-like ground, though fairly dry; was built into a small depression on the side of a grassy hummock; fashioned with a thin layer of dead grasses for the walls, and lined on the bottom with white down from the dwarf Arctic willow.” This was discovered on June 16 in a small upland valley near Nettilling Lake, Baffin Island; it held five eggs.
Frank L. Farley tells me that at Churchill, Manitoba, the “nests are usually set deep into the tundra and well protected with the last season’s growth of grass. They are made of grasses and liberally lined with ptarmigan feathers.”
Eggs: Hoyt’s horned lark lays ordinarily four or five eggs. The eggs described by Bendire (1895), as quoted under my account of the northern horned lark, are, of course, referable to this race, as they were taken by MacFarlane near the Anderson River, which is supposed to be within the range of Hoyt’s horned lark. The reader is referred to this account, which will apply equally well to most of the races of Of ocoris alpestris. The measurements of 33 eggs average 23.2 by 16.5 millimeters; the eggs showing the four extremes measure 26.7 by 18.8, 31.1 by 19.0, 21.6 by 16.3, and 23.6 by 15.5 millimeters.
Young: Dr. Sutton (1932) says: “The first newly batched young were found on July 3. The period of incubation therefore is probably about thirteen or fourteen days. The young are fed by both parents on insect-food, which is abundant at that season. The fully fledged young go about with their parents during the rest of the season. Unless the spring is unusually early, but one brood of young is raised. The young at the time of leaving the nest are in a much spotted and very pretty juvenal plumage, which is completely moulted in late August, apparently at about the same time the adults perform the post-nuptial moult.”
The plumage changes, food, and general behavior of Hoyt’s horned lark are apparently similar to those of the other northern races of the species.
Dr. Sutton (1932) says that, on Southampton Island, “its principal enemies are the Parasitic Jaeger, which eats the eggs and captures both young and adults; the weasel, which searches the ground carefully for the nests of small birds; the Arctic Fox; the Snowy Owl, which chiefly captures the young birds at the time its own young have to be fed; and the Duck Hawk. The Herring Gull does not greatly disturb this species, for it hunts chiefly along the lake-shores and coast and not in the high country of the interior.
“Hoyt’s Homed Lark leaves Southampton for the south among the earlier fall migrants. It has entirely disappeared before the last of the Snow Buntings, redpolls, pipits, and Lapland Longspurs have gone.”
Winter: According to the 1931 Check-list, Hoyt’s homed lark wanders southward in fall and winter to Nevada, Utah, Kansas, Michigan, Ohio, New York, and Connecticut, thus spreading out over a wide winter range and apparently thinly distributed and mixed with some other subspecies. Walker and Trautman (1936) say that in central Ohio, for example:
Hoyt’s Horned Lark (Otocoris atpestris Aoyti) Is by far the rarest of the three races that occur in this region. Most of our records are of one or two individuals associated with large flocks of 0. a. alpestris. These birds,. with the white superciliary line and pale dorsal coloration of praticola, but fully as large as the alpesfris with which they associate, are not difficult to identify in the field. The greatest number recorded, on December 29, 1928, at Buckeye Lake, was five in a flock estimated to contain 100 indivIduals of alpeslrte. Many large winter flocks of larks which we have carefully examined contained no hoyti nor have we found any flocks composed entirely of hopti. * * ~ The available central Ohio records for this race range from November 26 to March 17. Upon a few occasions we have heard a short song from Individuals of this race, and twice our attention was first attracted to the birds by a peculiar quality of the voice which seemed distinctly different from that of alpestris.
NORTHERN HORNED LARK
OTOCORIS ALPESTRIS ALPESTRIS (Linnaeus)
The horned larks form a most plastic species that has been split into a large number of subspecies, more or less recognizable, scattered over much of the northern portions of North America, Europe, and Asia. Our northern horned lark, 0. a. alpe8tris, stands as the type of the widely distributed species, because the name given by Linnaeus was based on Catesby’s bird that was supposed to have come from somewhere in the Carolinas. But the European race, 0. a. flava, is closely related to it and was once supposed to be identical with it. The northern horned lark is one of the largest and one of the darker-colored races of the North American subspecies. It might well have been called the northeastern horned lark, for Hoyt’s horned lark ranges fully as far north and the pallid horned lark ranges much farther north than alpestris.
Most of us know the northern horned lark only as a winter visitor, for few of us have enjoyed the privilege of seeing this hardy bird in its summer home on the northern barrens. To my late friend and companion, Dr. Charles W. Townsend (1923), belongs the credit for the discovery of the southernmost breeding station now known of this lark on the barren summits of the Shickshock Mountains, near the northern coAst of the Gasp~ Peninsula, Quebec, and about 200 miles south of Canadian Labrador. On the tableland above tree limit, on the summit of Mount Albert, 3,640 feet, he found a breeding colony of northern horned larks and pipits. He secured two specimens of the bird for identification, and says:
It was breeding In considerable numbers, for, at a very moderate estimate, I concluded there were twenty pairs. I saw several full-fledged young, and the old birds flew about with insects in their bills, scolding me anxiously. Occasionally I heard the flight song and saw the bird high In the air. * * * The summit of Mount Albert consists of a table-land some fifteen miles In extent, rising a little at the edges to plunge down In chasms and precipices. Protected by the northern rim of hard schists Is a straggling forest of black spruce and fir, rising to a height of five or six feet, with tops blasted by the arctic gales, and, on Its southern edge, a little lake imbedded in the mossy and grassy tundra. Beyond are great plains of brown serpentine rock masses, riven’ and heaved about by the frost, and beyond are other plains that appear almost as green and smooth as a lawn.
The flora is arctic in character, and comprises many species common to Labrador, such as curlew-berry, Labrador tea, pale-leafed laurel, moss camplon and creeping birch and willows.
Late in the spring and early in the summer of 1909 Dr. Townsend and I cruised along the south coast of the Labrador Peninsula, the north shore of the Gulf of St. Lawrence, traveling from Quebec to Esquimaux Point by steamer and from there to Natashquan, about 85 miles, in a small sailboat. On our arrival at Esquimaux Point, on May 24, we saw small flocks of northern horned larks; these were evidently migrating birds, for on our return there on June 2 these birds had all left. At Natashquan, on June 1, we collected a pair on the open, dry tundra near a small pond a short distance inland; they were evidently breeding there, for the female showed the wellknown signs of incubation. This is probably the western limit of the regular breeding range of the northern horned lark on this coast; from this point eastward the coast becomes more progressively Arctic m character. Audubon’s breeding record was much farther east, near Bras d’Or.
In 1912 I spent the month of June in Newfoundland, where I found the northern horned lark living and probably breeding on the treeless and tundra-like plains about Gaiftopsail near the center of the island. During that same season I cruised down the northeast coast of Newfoundland Labrador with Capt. Donald B. Mac
Millan as far north as Okak. We found a few pairs of these larks scattered all along the coast, but more commonly from Battle Harbor to Nain, wherever they could find the open and exposed situations that they like on the treeless coastal strip or on the rocky, mosscovered tops of the numerous islands. They seem to prefer the barren hilltops, where large beds of reindeer moss or other lichens of various colors partially cover the rocks and tundra and where there are no trees except the diminutive dwarf willows and birches, which grow only a few inches high and spread out over the ground, prostrated by the Arctic gales.
Spring: Most of the wintering flocks of horned larks leave New England before the end of April, though a few may linger well into May. We found a few small migrating flocks at Esquimaux Point, Quebec, on May 28, but by June 2 they had left. Lucien M. Turner says in his unpublished notes that in the vicinity of Fort Chimo, Ungava, “these birds are common in the spring migration only, arriving just after the middle of May”; but he found them breeding later on near the mouth of the Koksoak River. Taverner and Sutton (1934) found alpeatris migrating in company with koyti on May 28 at Churchill on the west coast of Hudson Bay. Dr. Sutton (1932) collected two from a flock of five males on Southampton Island on May 19, which were evidently migrating; he thought that alpestris occurs on this island only as a migrant, koyti being the breeding form there. Some of J. D. Soper’s birds, collected on Baffin Island, seemed to him to be referable to alpestris, though Mr. Soper (1928) says that the “great majority” of the breeding birds are typical of hoyti. Evidently these two races mingle on migration and probably intergrade in the regions north of Hudson Bay.
Courtship: I saw the courtship flight song of the male when I was in Labrador, but, as I could not hear it very well, I prefer to quote Dr. Townsend’s (Townsend and Allen, 1907) excellent account of it:
The bird suddenly mounts high Into the air, going up silently in irregular circles, at times climbing nearly vertically, to such a height that he appears but a little speck In the sky, several hundred feet up. Arrived at this eminence he spreads his wings and soars, emitting meanwhile his song, such as It Is: one or two preliminary notes and then a series of squeaks and high notes with a bit of a fine trill. The whole has a jingling metallic sound like distant sleigh bells, although the squeaks remind one strongly of an old gate. The whole effect, however, Is not unpleasant,: even melodious. Having finished one bar of his song, he flaps his wings a few times, closes them and sails again, repeating the song. One bird repeated his song twenty-four times and remained In the air one and a half minutes; another remained In the air three minutes, during which he repeated his song thirty-two times. During all this time the bird Is flying in curves or Irregular circles, sometimes In straight lines, or If the wind be strong, he beads up Into It and remains In the same place. The performance ended, he plunges head foremost down to earth, reaching It in a marvelously short space of time. The descent Is as silent as the ascent.
Nesting: Audubon (1841) gives the first account of the nesting of this species, as follows:
The Shore Lark breeds on the high and desolate tracts of Labrador, in the vicinity of the sea. The face of the country appears as If formed of one Ufl; dulating expanse of dark granite, covered with mosses and lichens, varying In size and colour, some green, others as white as snow, and others again of every tint, and disposed In large patches or tufts. It Is on the latter that the Lark places her nest, which Is disposed with so much care, while the moss so resembles the bird In hue, that unless you almost tread upon her as she sits, she seems to feel secure, and remains unmoved. * S The nest Is Imbedded In the moss to its edges, which are composed of fine grasses, circularly disposed, and forming a bed about two inches thick, with a lining of Grouse’s feathers, and those of other birds.
Townsend and Allen (1907) say: “At Frenchman’s Isle on July 16th, we found the nest of a Horned Lark composed of dry grass and a few large feathers, deeply sunk into the reindeer lichen and moss in a level piece of ground. There was no shelter or covering of any sort. It contained three dark-skinned young, clothed sparingly in sulphur-yellow down. Their eyes were not yet open.” Bendire (1895) quotes, from some notes sent to him by E. A. Mcllhenny, an account of a nest found by him on an island near Cape Charles Harbor on July 18, 1894, as follows: “The nest was embedded in a slightly inclining bank of moss and entirely below the surface of the moss; it contained five richly marked eggs, slightly incubated. When I found the nest it gave me the impression of being very small for the bird; but this was due to the fact that the entrance was small, and the hollow was enlarged under the moss. The nest was deeply cupped, having a thickness of about 1 inch of fine dry grass; it was lined with the down from reindeer moss and the white feathers of Ptarmigans. C. W. G. Eifrig (1905) reports a nest found near Cape Chidley: “The nest, placed on the ground, partly sunk in the moss, is made of moss, plant stems, grasses, finer toward the cup; this is lined with feathers and caribou hair. The outside diameter is 5 in., of cup 2 in., depth of cup, 1.75-2 in., outside depth, 2-2.50 in.”
Eggs: The northern horned lark lays three to five eggs, probably oftener four than any other number. Major Bendire (1895) gives a very good description of the eggs of the “pallid” horned lark (= lzoyti), which would apply equally well to those of this and the other races. He says that they: are mostly ovate in shape, less often elongate ovate. The shell Is close gralned, rather strong, and shows little or no gloss. The ground color Is mostly drab gray, sometimes grayish white; in an occasional specimen a faint greenish tint is perceptible, which fades out In time. The entire surface of the egg 1. profusely blotched and sprinkled with different shades of pale brown. In some specimens the markings are bold and well defined; In others they are minute, giving the egg a pepper-and-salt appearance; and again they are almost confluent, causing a uniform neutral brownish appearance. In some specimens the markings are heavier and become confluent about the larger axis of the egg, forming a wreath and leaving the ground color on the smaller end of the egg plainly visible; in fact, there appears to be an endless variation in color and markings as well as in size among these eggs and scarcely any two sets are exactly alike.
The measurements of 29 eggs of the northern horned lark average 22.6 by 16.5 millimeters; the eggs showing the four extremes measure 24.1 by 16.5, 24.0 by 17.4, 21.3 by 16.8, and 23.5 by 15.5 millimeters.
Young: Audubon (1840) writes:
The young leave the nest before they are able to fly, and follow their parents over the moss, where they are fed about a week. They run nimbly, emit a soft peep, and squat closely at the first appearance of danger. If observed and pursued, they open their wings to aid them In their escape, and separating, make off with great celerity. On such occasions It is difficult to secure more than one of them, unless several persons be present, when each can pursue a bird. * * * By the first of August many of the young are fully fledged, and the different broods are seen associating together, to the number of forty, fifty, or more. They now gradually remove to the islands of the coast, where they remain until their departure, which takes place in the beginning of September. They start at the dawn of day, proceed on their way south at a small elevation above the water, and fly in so straggling a manner, that they can scarcely be said to move in flocks.
Plumages: The nest found by Townsend and Allen (1907) “contained three dark-skinned young, clothed sparingly in sulphur-yellow down.” The juvenal plumage, which is acquired before the young bird leaves the nest and is alike in both sexes, is a fine example of concealing coloration, for it blends in so well with the surrounding lichens and mosses as to make the bird almost invisible in its open and unprotected nest. The crown is dark brown, almost black, and spotted with brownish white; the back is slightly lighter brown, mixed with dusky, and each feather is tipped with a spot of yellowish white, giving the whole upper surface a conspicuously spotted effect; there is a subterminal black bar on each of the scapulars with a broad terminal margin of yellowish white; the lesser and median wing coverts have large terminal spots of yellowish white; the greater wing coverts and remiges are margined with brownish buff; a superciliary stripe and a spot below the eye are pale yellow, as are the chin and throat, this color extending up the sides of the neck almost to the superciliary stripe; the chest is pale brownish buff, spatted with dusky; the rest of the under parts are very pale yellow or yellowish white.
A complete postjuvenal molt late in summer, mainly in August, produces a first winter plumage, in which the sexes are different. Dr. Dwight (1900) describes the first winter male as “above vinaceous buff, brightest on nape, vinaceous cinnamon on rump flanks and wing coverts streaked on head and back with sepia. Forehead, lateral ‘horns’, lores, auriculars and triangular breast patch black, veiled by overlapping pale buff or pinkish feather tips. Wings deep sepia, primaries much darker, edged with whitish, the rest of the wing feathers edged with vinaceous cinnamon. Tail brownish black, the outer rectrices edged with white, the middle pair paler, broadly edged with pinkish Isabella color. Below, dull white, the chin, sides of the head and forehead strongly suffused with lemon or canary yellow, a buffy band across breast below the black patch, flecked with dusky spots.”
The first winter female is similar but lacks the black forehead, which is streaked instead, the breast patch is smaller, the back is more streaked, and the colors are duller. The first nuptial plumage ls acquired by wear, with possibly some slight evidences of a prenuptial molt; the wearing away of the light-colored feather tips brings the black areas into clear prominence; and old and young birds look much alike, though young birds usually show more dusky streaking on the chest and flanks than adults.
Adults have one complete postnuptial molt on their breeding grounds, about August, and the nuptial plumage is acquired by wear, as in the young birds. Adults in fall are quite similar to the young at that season, the black areas being obscured by brownish tips, the yellow areas deeper yellow, and the white of the under parts more or less streaked with grayish brown.
Food: W. L. McAtee (1905), in his paper on the relation of horned larks to agriculture, publishes a long list of the vegetable food, mainly seeds, and the animal food, mainly insects, eaten by these birds, most of which does not apply to the northern horned lark. He has much to say about the injurious effect of weeds on agriculture and the cost to farmers in their control. Horned larks feed largely on seeds, perhaps mainly weed seeds, and so do many other birds, but I have always felt that the good that birds do in destroying weed seeds is a myth. Nature is so prolific in the production and so effective in the distribution of the seeds of plants, that only an infinitesimal percentage of those distributed can possibly find room to germinate; and no matter how many the birds pick up, there are always many times more than enough to cover the ground with verdure in a remarkably short time. Has any one ever known of a case where birds have kept even one square yard of ground free from weeds by eating the seeds? I certainly have not. Therefore, it seems to me that the eating of weed seeds is a neutral rather than a beneficial factor in the economic status of birds.
Cottam and Hanson (1938) have published a paper on the food of some Arctic birds, in which they give the results of the examination of three stomachs of this lark, taken at Indian Harbor, Labrador, in July. This shows what a large percentage of the summer food consisted of insects. They say:
Adult and larval Lepidoptera were consumed In numbers by each bird and averaged more than a fourth (27.33 per cent) of the entire amount consumed. These included the genus Agrotia sp. and undetermined Geometridae and Noctuldae. Large ants (Camponotus sp.) were next in the order of importance of the animal foods, averaging 7 per cent of the total. Other hymenopterous material, including ichneumon wasps, added another 5.67 per cent. A number of dipterous forms were next in order with 4.33 per cent, followed by spiders with 3.33 per cent. Leaf-hoppers, aphids, and other true bugs supplied 2.67 per cent, while mollusks, mostly a small M~tUua edulis, made up the remaining 1.67 per cent animal food, making an aggregate of 52 per cent.
Of the 48 per cent vegetable material, 31.67 per cent consisted of fruits and seeds of the bog hilberry (Vacdnium uliginosum), while the remaining consisted of cyperaceous seeds and undetermined vegetable debris.
While with us in winter the food must consist almost wholly of seeds, such as waste grain in the stubblefields, the seeds of forage plants in the fields, the seeds of eel grass, sedges, wild oats and mallows along our coasts, various grass seeds, and the seeds of ragweed and a host of other weeds. Probably some dried fruits or’ berries are eaten, and perhaps some insects in their dormant winter stages.
Most of its food is apparently picked up from the ground, where it spends most of its time walking nimbly along among the stubble, in the short grass or over the salt marshes. Dr. Townsend (1905) says: “It picks at the grass-stalks from the ground, never alighting on them as do the snow buntings and longspurs. It sometimes flies up from the ground, seizing the seeds on the tall grass or weedstalks, at the same time shaking many off onto the ground, which it picks up before flying up to repeat the process. Horned larks are frequently found in roads picking at the horse-droppings, especially when much snow has covered the grasses and weeds. They also come into the farm-yards for scraps of food.”
Behavior: As we see them in winter northern horned larks are decidedly gregarious, occurring in flocks that range in size from half a dozen to a hundred or more birds; they are seldom seen singly or in pairs as in their summer haunts. As we walk across some flat salt marsh near the shore, or some bare stubblefield farther inland, we may be surprised to see a flock of these birds arise from the ground, where their quiet movements and concealing coloration had rendered them almost invisible. They rise all together, and we hear their faint sibilant twittering as they circle about, now high in the air in scattered formation, now close to the ground in more compact order, showing a bright glimmer of white breasts as they wheel away from us, then suddenly disappearing from our view against the dark background as they turn their backs toward us, and finally vanishing entirely as they all alight on the ground not far from where they started. Their flight is light and easy, with a somewhat undulating motion; and the flocks are rather loose and irregular, yet they are apparently all in touch with each other and guided by a common impulse. As they alight on the ground they scatter out and walk about rapidly on their short legs, taking rather long steps, as shown by the marks of the long hind claw in the soft mud or sand. Horned larks are essentially ground birds; I have never seen one alight in a tree, and, so far as I know, no one else has. The top of a rock, stone wall, or low stump, not over 3 or 4 feet above ground, is about as high as they care to perch, and that not very often. They prefer open ground, especially bare ground or where the grass is short, and they are almost never seen where the vegetation grows rank and high. Among the stubble or tufts of short grass, they walk or run in a crouching attitude, reminding one more of mice than of birds; often they squat and hide until too closely approached. They are not particularly shy, if carefully approached, and seem to feel aware of their ability to conceal themselves in scanty cover. If we remain motionless while the bird is hiding, it will soon lift its head and look about, but at the slightest movement on our part it squats again or runs or flies away.
Dr. Townsend (1905) says: “It is a persistent fighter or extremely playful, whichever you will, and is constantly engaged in chasing its fellows. I have seen two face each other for a moment, with heads down like fighting cocks, the next instant twisting and turning in the air, one in hot pursuit of the other. When in flocks with the other winter birds, they more frequently chase them, especially the smaller Longspurs. I have also seen them chase Snow Buntings, and often Ipswich Sparrows that were feeding with them, end once, what appeared to be a Prairie Horned Lark.”
Voice: Aside from the courtship flight song, described above, the vocal performances of the horned lark do not amount to much. Townsend and Allen (1907) say of other notes, heard in Labrador: “The familiar sibilant squeaking call note was commonly used, and also a note which we do not remember to have heard during the migration in Massachusetts. This sounded like zzurrit and was often preceded by another note thus, whit-zzumit. These notes were occasionally so soft and sweet that they recalled the trilling whistle of the Least Sandpiper.”
Ralph Hollmann (1927) writes: “The common note of the Horned Lark is a shrill tsee, or ts~e-de-ree, and a still sharper double-syllabled ti-sick. The song is thin and unmusical, suggesting the syllables, taip, tsip, ts4e-di-di.” Dr. Townsend (1905) writes it “tss8wee it, tS8wt, the sibilant being marked.” This is the note commonly heard in winter, usually uttered in flight, but often given from the ground or from some slight eminence. 0. 5. Murie writes to me that one of these birds appeared to answer his imitation of its notes.
Field marks: If the bird is facing the observer, the conspicuous head markings are unmistakable, though in young birds and females, especially in fall and winter, these markings are much obscured. While walking away from the observer on the ground, no conspicuous field marks appear, but a horned lark can generally be recognized by its thick-set appearance, by its habit of walking instead of hopping, and by its mouselike movements. It is larger than the pipit or the Lapland longspur or any of the sparrows with which it is likely to he associated. The pipit has more white in the tail than the lark; and the lark does not wag its tail as the pipit does. When the lark is flying overhead its black tail shows in sharp contrast with the white under parts.
Winter: The old name, shore lark, seems very appropriate for this bird, for while with us in winter in New England it is far more abundant along the coast than elsewhere. Here it is often associated with the snow buntings and the Ipswich sparrows in the sand dunes and on the beaches, or with the pipits and Lapland longspurs in the salt marshes and meadows. Flat, open, brackish meadows along our tidal rivers are favorite resorts, and the birds are often seen about the shores of lakes and even in stubble fields and plowed lands farther inland.
In Massachusetts the northern horned lark is more abundant as a migrant than as a winter resident, though it is here in some numbers all winter. Dr. Townsend (1905) says of its seasons in Essex County: “During the first half of October, Horned Larks are found in small numbers, but they become abundant in the latter half of the month, increase through November, and reach their height in December. During most of January they are common but in the latter part of that month and in February and early March comparatively few are to be found, while in the latter half of March they again increase in numbers but are never as common as in the fall, and a few may occasionally be found early in April.”
Walker and Trautman (1936), referring to its status in central Ohio, write:
The Northern Horned Lark (0. a. alpestris) is unquestionably the dominant race during the winter months. * * Flocks of from twenty to one hundred individuals are usually present by early November. The peak of abundance occurs during December, January, and February when flocks of 200 or more are frequently encountered. The largest flock noted by us was estimated to contain 600 individuals and was seen in the cornfields of the Scioto mver bottom-lands a few miles south of Columbus on February 18, 1928. The largest number recorded In a single day was that of an estimated 2000 Individuals, the combined number of several flocks which were encountered along a three mile stretch of road ImmedIately south of Buckeye Lake on February 14, 1929. During the month of March there is a rapid decline in numbers.
Severe winter weather or cold storms, especially snowstorms, sometimes drive these northern larks as far south as North Carolina or even South Carolina, where they seek their food in the shelter of bare furrows or in the lee of tufts of grass in the fields. When the ground is covered with snow they manage to find food by scratching little hollows in the snow, or they resort to the barnyards to pick up hayseed and waste grain. Probably most of the horned larks seen in the Southern States in winter are of the prairie subspecies.
Range: Circumpolar; from the Arctic coast of both hemispheres south to northern Africa and South America.
Breeding range: The North American breeding range extends north to Alaska (St. Michael and Fort Yukon); Yukon (Herschel Island); and the Northwest Territories (Liverpool Bay, Horton River, Kent Peninsula, Cape Fullerton, Bowman Bay, and Resolution Island). East to the eastern part of the Northwest Territories (Resolution Island); extreme northeastern Quebec (Button Islands and Cape Chidley); Labrador (Okak, Davis Inlet, and Rigolet); eastern Quebec (Battle Harbor, Cape Charles, and Loup Bay); Newfoundland (Canada Bay and Cape St. Mary); New Brunswick (Scotch Lake); Maine (Eustis and Waterville) ; rarely eastern Massachusetts (Essex County, Plymouth, Barnstable, and Nantucket Island); rarely western New Jersey (Mount Holly and probably Gloucester and Salem Counties); rarely the District of Columbia (Washington); central Virginia (Lynchburg and Naruna); western Tennessee (Nashville); eastern Arkansas (Helena); eastern Texas (Galveston, Corpus Christi, and Brownsville); Tamaulipas (Miquihuana) ; Hidalgo (Real del Monte) ; Veracruz (Mirador and Perote); and eastern Oaxaca (San Mateo). South to southern Oaxaca (San Mateo, Oaxaca, and Mitla); Mexico (Valley of Mexico); Guanajuato (Silao); Durango (Durango); and central Baja California (Santa Rosalia Bay and San Ignacio Lagoon). West to Baja California (San Ignacio Lagoon and San Quintin); California (San Clemente Island, San Miguel Island, Santa Cruz, and Red Bluff) ; Oregon (Fort Kiamath and Wapinitia) ; western Washington (Seattle and Tacoma); British Columbia (Spence Bridge, Chilcotin, 150: mile House, and Wilson Creek); and Alaska (Kenai Mountains and St. Michael).
The range as outlined is for the entire species, which has been separated into no less than 16 currently recognized subspecies or geographic races in North America. The typical form, the northern horned lark (Otocori8 alpe8trzs alpestris), breeds in the Ungava Peninsula, Labrador, and Newfoundland; Hoyt’s horned lark (0. a. hoyti) occupies northern Canada from Hudson Bay west to the mouth of the Mackenzie River and south to the northern parts of the Prairie Provinces; the pallid horned lark (0. a. arcticola) breeds in Alaska (except the Pacific coastal region) and south in the mountains through British Columbia to Washington; the desert horned lark (0. a. teucotasma) occupies the region from southern Alberta to New Mexico and Texas, east on the Great Plains to South Dakota and Kansas, and west to Nevada; the prairie horned lark (0. a. praticola) is found from southern Manitoba and Quebec south to eastern Kansas, Arkansas, Kentucky, and Virginia; the Texas horned lark (0. a. giraudi) occupies the coast of Texas and northeastern Tamaulipas; the streaked horned lark (0. a. strzgata) inhabits the Pacific coast region of Washington, Oregon, and northern California; the dusky horned lark (0. a. mer’rilli) is found in southeastern British Columbia, eastern Washington and Oregon, northeastern California, and northwestern Idaho; the island horned lark (0. a. insu~aris) is confined during the breeding season to the Santa Barbara Islands of California; the California horned lark (0. a. actia) is found in California south of San Francisco Bay, east to the San Joaquin Valley and south to northern Baja California; the Magdalena horned lark (0. a. enertera) is confined to the central part of Baja California between Santa Rosalia and Magdalena Bays; the ruddy horned lark (0. a. rubea) is the race of the Sacramento Valley, California; the Montezuma horned lark (0. a. oceidentalis) breeds in central Arizona and New Mexico; the scorched horned lark (0. a. adusta) appears to be confined to a relatively small area in southeastern Arizona; the Mohave horned lark (0. a. ammophila) nests in the Mohave Desert and Owens Valley, Calif.; and the Sonora horned lark (0. a. leucan8~iptila) occupies a region extending along the Colorado River from southern Nevada and western Arizona south to northeastern Baja California. Additional races of this species are found in Europe, Asia, and Mexico, as well as one (0. a. peregrina) that appears to be localized in the vicinity of Bogota, Colombia.
Winter range: The species is found throughout the year over most of the breeding range, although it withdraws during the winter from the northern regions. At this season it is found north to southern British Columbia (Arrow Lake); southern Alberta (Warner and Sullivan Lake) ; southern Saskatchewan (Eastend and Skull Creek); northern North Dakota (Charlson and Grafton); northern Minnesota (Fosston and Iron Range); southern Wisconsin (Westfield and Greenbush) ; southern Ontario (Guelph, Toronto, and Ottawa); central Maine (Bangor and Calais) ; and Nova Scotia (Grand Manan and Kings County). Occasionally they are recorded in winter from points farther north in Alberta (Battle River) and Saskatchewan (Dinsmore); as well as from southern Manitoba (Winnipeg), southern Quebec (Montreal and Quebec), and southern New Brunswick (Scotch Lake). In the eastern part of the country the southern limits of the winter range extend south of the breeding range to Florida (St. Augustine and Apalachicola, rarely Daytona Beach and Miami); and northern Alabama (Decatur and Leighton).
Spring migration: Because of the fact that in the East the horned lark winters regularly to the northern parts of the United States and southern Canada, and in the West is resident south to and beyond the Mexican border, dates of arrival and departure are not numerous for the seasonally unoccupied regions.
In the southeastern part of the country late dates of spring departure are: Florida: Hastings, March 18. Georgia: Athens, February 25. North Carolina: Pinehurst, February 22.
Early dates of spring arrival at points north of the winter range are: Quebec: Montreal, February 23. Labrador: Gready, April 15. Newfoundland: Raleigh, April 24. Mackenzie: Fort Simpson, April 28. Yukon: Forty-mile, May 10. Alaska, Demarcation Point, May 6.
Fall migration: Late dates of fall departure from the North are: Alaska: Swan Lake, August 19. Yukon: Russell Mountains, September 4. Mackenzie: Fort Simpson, October 3. Labrador: Ticoralak, October 12. Newfoundland: October 14. Quebec: Montreal, November 15.
Early dates of fall arrival south of the breeding grounds in the eastern part of the country are: North Carolina: Raleigh, November 21. South Carolina: Chester, December 3. Georgia: Clayton County, November 16. Florida: Daytona Beach, November 20.
Egg dates: Arctic Canada: 7 records, June 14 to July 9.
Arizona: 15 records, April 8 to July 28; 8 records, April 29 to June 10, indicating the height of the season.
California: 106 records, March 20 to June 23; 52 records, April 17 to May 20.
Colorado: 26 records, April 9 to July 16; 14 records, May 10 to June 9.
Illinois: 32 records, March 3 to July 6; 16 records, April 25 to June 17.
Iowa: 19 records, March 27 to July 16; 10 records, May 14 to June 19.
Labrador: 7 records, June 1 to 30.
Lower California: 2 records, April 6 and 23.
Montana: 46 records, April 10 to July 7; 23 records, May 7 to June 4.
New Mexico: 11 records, May 20 to July 6.
New York: 14 records, March 19 to May 30; 8 records, April 1 to 22.
Northwest Territories: 5 records, June 1 to July 18.
Santa Barbara Islands: 5 records, April 4 to May 14.
Saskatchewan: 4 records, May 15 to June 9.
Texas: 22 records, February 20 to June 20; 12 records, April 24 to May 23.
Washington: 43 records, May 20 to June 25; 21 records, April 25 to May 30.
Wisconsin: 11 records, March 21 to June 25; 7 records, May 15 to June 18.
DESERT HORNED LARK
OTOCORIS ALPESTRIS LEUCOLAEMA (Coues)
On the more barren plains of the far West, we find this horned lark replacing the familiar prairie horned lark of the more fertile prairie regions of the Middle West. It is about the same size as and only slightly paler than praticola. In comparing it with neighboring races, Dr. Oberholser (1902) says: “This form may be distinguished from praticola by the markedly more cinnamomeous tint of cervix, upper tail-coverts and bend of wing, as well as by the paler color of the back, where the blackish of praticola is replaced by sandy brown. From arcticola it differs in reduced size, usually yellow throat, nape more tinged with cinnamomeous, lighter and brownish instead of blackish back; from giraudi in larger size, generally paler throat, together with paler, much more brownish upper surface; from merril2i in larger size and lighter, more brownish coloration.”
In southwestern Saskatchewan, in 1905 and 1906, we found horned larks very common on the prairies, on the barren hills north of Maple Creek, and on the alkaline plains. As we drove along the narrow wagon trails over the rolling plains, the monotony of the scenery was often relieved by seeing one of these little brown-backed birds running along in the wagon ruts ahead of the horses, perched on a clod of earth beside the road, or springing into the air to pour out its quaint little ditty, not quite equal to the rapturous flight songs of the chestnut-collared longspurs but, nevertheless, pleasing.
We collected a series of horned larks here and in Alberta, most of which, particularly those taken on the prairies in the more eastern portions of the region, were referable to the new form described by Dr. Oberholser (1902), 0. a. er&thyraia, while those collected on the alkaline plains and sagebrush plains farther west were more clearly referable to leucolaema.
Courtship: Although the habits of the various races of the horned lark are all very much alike, different observers have described them somewhat differently, illustrating certain phases of behavior more clearly than has been done by others. For this reason, I shall, at the risk of some duplication, quote freely from two excellent papers by A. Dawes DuBois (1935 and 1936) on the habits of the desert horned lark. He noted that the male, in his upward nuptial flight, usually ascends at an oblique angle, but that against a strong wind he rises almost vertically. He then goes on to say (1936): “After remaining aloft for a time, singing his best song, which comes to the human ear but faintly from so great a height, the bird suddenly folds his wings and drops like a bullet. With ever increasing velocity he descends until one might fear for his life; but he spreads his wings just in time to avert a violent end, skillfully turning his course into a glide which carries him horizontally, near the ground, until his momentum has been spent. He then alights quite easily, as though nothing important had happened. * * * I doubt that the bird world holds a more awe-inspiring event than this headlong drop from the sky.”
He thinks that this spectacular dive “far surpasses the performance of the nighthawk” but does not compare it with the thrilling dives performed by some of the humrningbirds, which seem equally inspiring. He says:
Another ceremony of the season is the fighting exhibition, which takes place in the air a few feet above the ground. The two males engaging in it begin their advance and attack while on the ground but immediately rise together in a whirl and flutter of gallantry. * * There seems never to be an injury, nor even a victory. I have never seen a drop of blood drawn or a feather lost in the encounters. * S The third sort of maneuver is an exciting chase. Two, three, or four birds usually take part in it. They fly in close formation with great swiftness and remarkable skill. It looks like a game of follow the leader, with instant response to every change of the leader’s course: a course of rapidly changing, meandering curves.
Nesting: Mr. DuBois (1935) has given us a very full account of the nesting habits of this lark in Teton County, Mont., based on four years of study of 58 nests. His data are given in far too much detail to admit of more than a few extracts from them, as follows:
Two peaks of nesting activity occur, one about the end of April, the other early in June, indicating two broods in a year. * * The Desert Horned Larks avoid the more luxuriant growths which are to be found in moist situations
They prefer the dry bench lands. There Is no special preference as to surface contour so long as the situation is a dry one. Nests occur on knolls or slopes, or In the dry depressions of the benches. * * The number of nests which one finds near old dried droppings of horses, and sometimes of cattle, seems much greater than the laws of chance would account for. * * * Only one nest was found in a cultivated field. ï It was in a field of young spring wheat which stood in drills, about two Inches tall, the ground being otherwise bare.
The nest is Invariably built in a rounded hollow In the ground, which Is evidently scratched out by the birds, the excavated dirt In the form of fine scratchings being thrown out to one side of the nest. The dirt is almost always on the east side, which Is also the side least protected by vegetation. Usually the top of the nest structure Is flush with the ground surface. ï The materials used for the body of the nest are dead grasses, including both stems and blades, usually without any other materials.
The nest built on cultivated land contained rootlets and old dead grass. The linings were more varied, including a bit of rag, some tiny bits of rabbit fur, soft, silky, white plant down, and seed pods, heads, tips, or leaves of yarrow, which when dried are gray or white and of soft, velvety texture.
“All nests examined, with only one exception, were provided with pellets of dried mud at the entrance or elsewhere around the nest. Them are little cakes or broken pieces of the cracked crust which forms on the surface of mud when baked by the sun. * * * The pellets are used chiefly to cover the loose dirt thrown out in excavating the hole for the nest.”
The inside diameter of nine nests averaged 2.49 inches, and their inside depth averaged 1.92 inches. The ground hollow for one nest was 4 inches in diameter and 2 inches deep. The time required to build the nest, after the hollow was dug, varied from 2 to 10 days. As to concealment of the nests, he says: “The prevailing short grasses of the bench lands do not afford much cover. The concealment of nests in general, so far as the surrounding grass is concerned, is very incomplete, sometimes quite meager. Nevertheless, the nests are not easy to see. In most cases there is some protection from grass on the west side; sometimes it slants over the nest, owing to the prevailing winds.”
Eggs: The desert horned larks usually lay three or four eggs, perhaps very rarely five, though none of Mr. DuBois’s 58 nests contained five. These are practically indistinguishable from those of the other races of similar size. Some sets are somewhat paler and less heavily marked than those of the darker races. The measurements of 50 eggs average 22.1 by 15.6 millimeters; the eggs showing the four extremes measure 23.9 by 15.5, 23.4 by 17.0, 20.6 by 15.2, and 19.6 by 15.2 millimeters.
Young: In his second paper (1936) Mr. DuBois gives a full account of the developments of young desert honed larks, to which the reader is referred for details, as only portions can be quoted here. He writes: “Apparently the male never assists in the duty of incubation, but both parents take part in feeding the young as soon as the eggs have hatched. The food for the young is evidently of a solid nature from the beginning. A parent was observed carrying a smooth green caterpillar in the afternoon of the day of hatching. Large larval insects are fed to the young birds after they are strong of flight. Parent birds were seen carrying excreta away from the nest when the nestlings were two and three days old.” He tells of the hatching of a chick from an egg held in his hand:
When It was ten hours old, nearly all of its natal down was dry, fully three-eighths of an Inch long, and very fluffy: a marvelous transformatlon! * * * At the age of seven or eight days the nestlings are fairly well feathered and the natal down is confined to the feather tips. * * ï When ten days old the young have left or are, In most cases, leaving the nest. They are not able to fly but can run very well. It appears that they usually leave in the fore part of the day. * * * It is easy to Identify the horned lark nestlings, at any stage of their development, by looking Into their mouths. The mouth lining Is orange, and there are distinct black marks in the mouth and on the tongue. This distinguishes them at once from the nestllngs of Mecown and Chestnut-collared longspurs, which have plain pink mouths and throat linings. When the young larks have grown up, the orange color fades and the black marks disappear.
Leon Kelso (1931) also gives a detailed account of the development of the young in four nests of this lark, to which the reader is referred. lie states: “It is evident that the rate of growth and length of time spent in the nest by nestling Desert Horned Larks varies according to the time of the year. The young of nests 1 and 4,in the months of April and May, respectively, remained in the nest at least ten days, while those of 2 and 3, in July, stayed in the nest not more than 7 days. The size attained appeared to be comparable in all instances. * * * The first nest has a lining of thistle-down, contained 1505 pieces of material and weighed 16.75 grams; the second had no lining, was built of 805 pieces, and weighed 7.7 grams.”
Enemies: Mr. DuBois (1936) writes:
On their nesting grounds the Desert Horned Larks have to contend with their share of enemies and sources of accident. Among the natural enemies, weasels, skunks and ground squirrels came to my attention, not to mention man, whose poisoned baits set out for ground squirrels apparently kill more birds than spermophiles. One day, by quick action, I Intercepted a weasel on his way to a nest to get the last nestling. The birds, of course, are powerless to defend their young against weasels and skunks. It Is believed that the abundant ground squirrels often destroy eggs, and possibly sometimes take a nestling; but the adult larks are not afraid of them. It is common to see the larks driving a trespassing squirrel away from their premises. They go after him from the air, in a series of dashes; and quite often the two birds attack together.
The barbed wire fence, new in part of the region when these notes were made, was a source of unexpected danger. Several carcasses of horned larks were found at different times beneath wire fences * ï * Storms cause the greatest destruction of nestlings. Eggs and young are kept dry during all ordinary rains. But In some years the destruction of nests by severe and protracted storms Is doubtless, over an extensive region, practically total. A continuous rainstorm of three days duration, coming first from the east, then from the north and finally from the northwest, killed all the nestllngs that were known to me.
William G. Smith wrote to Major Bendire (1895): “While I lived in the Platte River Canyon, 40 miles west of Denver, Colorado, a terrible snowstorm set in suddenly in April, and with it came thousands of these birds, which tried to shelter themselves under projecting banks. The majority were soon so chilled by the intensely cold wind which was blowing at the same time, that they could not move, and were quickly smothered by the drifting snow; and after this melted bushels of their dead bodies could be picked up everywhere.”
Probably this horned lark is imposed upon occasionally by the cowbird, but Dr. Fried~inann (1929) cites only one authentic record; it would seem as if horned larks might be frequently victimized where cowbirds are common.
Winter: After the second brood of young is strong on the wing these horned larks gather into immense flocks and roam about the country preparing to move southward. Many extend their winter range into the more arid regions of our southern border States and even into Mexico. But through a large portion of its range this race is largely resident and is found less commonly as far north as Montana in winter. It is common in winter at least as far north as Wyoming. In Colorado, it is very abundant in winter, traveling about in enormous flocks in company with some of the other subspecies. When the ground is bare the flocks spread out over the plains and fields, but when the snow covers their feeding grounds they congregate about the ranches and farmyards to feed on the waste grain or come into the towns and cities to be fed by the residents; sometimes in severe weather thousands of the birds come into the towns, where people feed them regularly on millet and other seeds, scattered on bare spaces, where the birds often gather so thickly as to almost cover the ground.
Claude T. Barnes writes to me from Utah: “Though the day be cold and drear, and all vegetation well nigh covered with snow, in the fields west of Salt Lake City, flocks of pretty horned larks are daily seen feeding on the seeds of the pigweed, saitbush, ragweed, amaranth, and other noxious weeds, which here and there protrude through the snow. If the snow becomes too deep for them, they even venture into the city.”
PRAIRIE HORNED LARK
OTOCORIS ALPESTRIS PRATICOLA Henshaw
Contributed By GAYLE PICKWELL
Out at the bleak end of the ecological series of bird habitats, which begins with the heavy forests and ends with the barrens, lives America’s only true lark, Otocorie alpestri8 (Linnaeus). In that region extending from Missouri to the Atlantic and from Kansas to Ontario the particular form of this lark is Otocoris alpestris praticola Henshaw, the prairie horned lark. Far from the treeless Arctics, far from the deserts, this lark finds as its barrens the plowed fields of the Midwest, the tree-denuded, wind-swept hilltops of the Northeastern States, and those peculiarly unnatural and artificial barrens, the hazards of these modern-day golf courses.
If for no other reason than that here is a bird nesting where no bird has a right to nest, a bird in a niche that demands not vegetation but lack of it, a bird alone and unique in its nesting site without a competitor and far out at the end of the series: if for no other reason than this purely ecological one: the prairie horned lark invites close study. But if we add to this the fact that it is a lark, a representative of our only lark, with the song of a lark, the ways of a lark, and many a habit and idiosyncrasy peculiarly its own, and that it is an intriguing bird of the open field, then the bird becomes even more interesting.
The prairie horned lark, because of its tendency to occupy the most barren regions as its home, interested me very early, for desultory observations of this bird were begun while still a boy in eastern Nebraska. The lark nests were found on the ridges of listed corn and an observation of a song still remains clear and trenchant. We were shocking wheat, hence it was mid-July, when a lark was seen climbing the air for his song. We watched him against the vivid sky during his long minutes aloft; were amaz~d by that final headlong drop to earth.
Study of the prairie horned lark was initiated in eastern Nebraska, continued intensively in northern Illinois (Evanston) for two years, then transferred to Ithaca, N. Y., and concluded. Since that time I have lived in California, where the prairie horned lark is replaced by the California horned lark, a closely related subspecies.
‘Derived largely from PIckwell, ‘The Prairie Horned Lark” (iOn).
Henshaw (1884) erected the subspecies Otocoris alpeatris praticola, splitting it from 0. a. alpestris. Prior to this records of a new form of lark and new lark breeding records were published from lower Ontario and New York. These were variously interpreted as a “paler form” or as a southward extension of 0. a. alpestris. Following 1884 a consistent and progressive series of records demonstrated that the prairie horned lark, coming up probably from Michigan through Ontario, invaded successively New York, Vermont, Massachusetts, New Hampshire, Maine, and Connecticut. From New York or Vermont it seems to have invaded Quebec much later; and lastly on the north, probably from the New England States, New Brunswick, Prince Edward Island, and Nova Scotia. Shortly after its entry into New York the lark appeared in western Pennsylvania, then farther east in that State, and south into Maryland and West Virginia. Recently the prairie horned lark has been recorded as breeding in the District of Columbia and Virginia.
Less complete evidence seems to show that Indiana, Michigan, Ohio, probably northern Kentucky, and southern Missouri have been occupied by this lark since the white man has entered and altered those regions. The regular advance of the bird, always consistent with geographic conditions, is suggested as an irrefutable evidence that such an extension is bona fide. It is suggested that this extension of range has resulted from changes that civilized man has made by deforestation and cultivation; thus creating permanent or seasonal semibarren conditions that the prairie honed lark requires.
The drier portions of the prairies of Illinois have probably long been occupied by this lark. The studies of Forbes and Gross (1922) seem to indicate that the lark, though it probably breeds in Lower Austral, Upper Austral, and Transition Zones, seems to prefer the Transition in that State. It is suggested that the prairies of northern Missouri, Iowa, Wisconsin, Minnesota, eastern portions of Kansas, Nebraska, North and South Dakota, and southern Manitoba probably formed the ancestral home of the subspecies. Nearly all this vast region would have been suitable for two broods in March, April, and early May, though the bird would have been forced to the more barren regions as the grasses became vigorous in late May, June, and July. That this lark species is ‘versatile in the matter of occupation of new territory seems to be further demonstrated by the observation of Glitke and Saunders in Europe with regard to 0. a. fla’va.
Courtship: Prior to the establishment of well-defined territories, fighting between males is promiscuous; after that fighting takes place only on territory boundaries, where two lark areas juxtapose. The males, at the boundary line, frequently strut before each other and often peck the ground furiously, like barnyard cocks, but all fighting is in the air. On a boundary this fighting often results in a curious game of “tit for tat,” as the male larks chase one another back and forth. Every adventitious lark, wandering into established territories, is promptly evicted by the male. Such a bird will leave without protest. So far as noted, the female is never the direct cause of fighting; in fact fighting is most frequently noted when the female is brooding and the male is no longer attending her. Only once was a female noted driving out another lark, a male. She was defending a recent nestling.
The female has no courting maneuvers and was never observed to sing. Only once was she seen to importune sexual attention and then by a crouch and flutter similar to the actions of the English sparrow. The male struts frequently before the female with wings dropped, tail spread, and horns up. He will assume this attitude before another male at the territory boundary.
Nesting: The literature shows a surprisingly large range of habitats in which the prairie horned lark has been known to nest. These habitats, resulting for the most part from agricultural activity or other human agencies, are those that most nearly result in barren conditions. It does not matter that these barrens may be seasonal or otherwise very temporary, if they are suitable for the initiation of nesting. That bare ground is the determinant is shown by the fact that variations of moisture, soil, elevations, and temperature will all be tolerated in the selection of nest sites. The prairie horned lark thus, it seems, does not differ greatly in the ecological condition of breeding habitats from other horned lark subspecies.
Some typical Chicago marsh in the Evanston region was drained for a golf course. The course was later cut up into real-estate subdivisions; sewers were laid exposing a wide area of bare soil in the streetways; and old sand hazards remained here and there. This series of activities provided nesting sites for many larks. More than a score of nests were located on this area (about 90 acres) in 1926. A plot of vegetable gardens bordering this region on the west, where larks had probably nested for some years, was also subdivided and the vegetation subsequently neglected. Here several larks also nested.
The advent of vegetation in both areas and the demand of the lark for bare ground forced a seasonal succession of horned-lark breeding sites first from lot surface, to streetway, to sand hazard, to vegetable garden, in the order that each was successively occupied by verdure.
At Ithaca, N. Y., one nest was located on the overturned sod of a former hay meadow. Most of the observations there made, however, were on a tract of ground that was largely fall wheat, partly fall rye, and the remainder devoted to experimental vegetable gardens. The growth of the wheat forced the larks from its surface by late May. The gardens and portions of the fall rye area that were turned under as green manure remained suitable throughout. Clean vegetable gardens will always present a considerable amount of bare soil, and the prairie homed lark is usually able to occupy such gardens until late in June.
A breeding territory was delimited by a male lark on February 7, 1926, at Evanston, Ill. From his selected territory he could not be driven. This territory was about 100 yards square. Late March snows disrupted all territories, and it was not learned whether the original sites were ultimately resumed or whether the same territory was maintained through more than one nesting. The pressure of vegetation in late May and June greatly modified the territories at Evanston and caused, eventually, the abandonment of most of those on the erstwhile golf course.
At Ithaca, N. Y., a male lark was forced to mark territory for the first time on March 13, 1927, though it had undoubtedly been established some time before this. Territories voluntarily marked were somewhat larger than those indicated when the birds were forcibly driven about. The regions of a breeding territory most frequently occupied were those boundaries that joined the territories of a neighboring lark.
The territores at Ithaca were much larger than those at Evanston, possibly because fewer larks attempted to occupy them. At Evanston they were seldom over 100 yards square, whereas at Ithaca they ran out to lengths of 300 yards and widths of 200 yards, in March and April. In general all suitable territory was occupied at Ithaca and most boundaries were established by the margins of unsuitable areas, though a large amount of suitable territory, extending beyond, was used only in part by the bird. Boundaries between males were often definitely established on ground that had no natural marker whatsoever.
The territory history of three pairs of larks was followed from March to June at Ithaca. One influence only modified the territories, namely, the growth of vegetation. One territory, completely on fall wheat, was abandoned by the close of tbe second nesting in May. Another territory, in part on fall wheat and in part on the gardens, was gradually reduced to the gardens, from an area once 300 by 200 yards to an ultimate area about 100 by 50 yards. A third territory, almost entirely on the gardens, suffered no major reduction. But the owner of this third territory, which abutted that of the second, gave no ground to the latter.
Though most of the feeding was done on the nesting territories, a neutral feeding territory was discovered, and others were indicated because, now and then, the larks would go off on purposeful flights entirely out of their areas.
The female would mark the same territory as that marked by the male, and if anything she was more closely restricted to it than the male. She selected the nest site with little or no regard to the center of the area.
The literature contains four February records of nests and many records of March nests in many States, and two or three records of nests in July. I have records of nests from about March 21 to July 12, in 1926, at Evanston, Ill.; from about March 11 to June 28, in 1927, at Ithaca, N. Y.
It is suggested that such a strange phenomenon as that of a passerine bird nesting in March in eastern United States cannot be easily explained. The bird has too long a nesting season to explain it on the conditions that might exist in early spring alone; and then, in the range where the prairie horned lark was studied, nests are frequently destroyed by inclement weather and many young die of starvation at this season. Since this bird demands barren conditions, and not verdure, for a nest site, the conditions are suitable very early, and it is suggested that an early-nesting physiological cycle may have been acquired in a more propitious climate and subsequently carried north and east. It is further noted that 0. a. actia of California nests in March where conditions are quite ideal.
With one exception all of the 14 observed nests of March and April were not begun until the mean temperature rose above 4O~ F. for two or more days in succession. The exception was the initiation of a nest on the fi~rst day that the temperature rose above a mean of 400 F. Once weather conditions suitable for the initiation of nesting activities prevailed, no subsequent weather, no matter how severe, except deep snow only, would inhibit these activities. Even birds that had nested in March and whose nests were destroyed by late March and early April snows, would not renest until weather conditions were as given, though this necessitated a delay of nearly three weeks in two cases at Ithaca, N. Y. That this was a delay caused by the weather is easily demonstrated by the fact that an exceptional case, as noted above, began renesting on a single suitable day, but two other larks waited two weeks longer for renesting or until weather again was suitable and for a longer period. It is known that two of these birds, and probably all, had former nests.
On the basis of this known weather control it was possible to calculate the frequency, over a period of years, with which nestings would occur in March, by a study of weather summaries for the month. The results showed one year when nesting was impossible and 16 years of possible nestings, at Evanston, Ill., for the years 1910 to 1927, inclusive, except 1924. During 10 of the 16 years nestings could have been successful; during 2 they would have been destroyed by snow; and during 4 weather and snow would have made success problematical. At Ithaca, for the years 1916 to 1927, inclusive, the summaries showed one year when nests were impossible and 11 years of possible March nests. During 5 of the 11 years, nestings could have been successful; during 4 they would have been problematical. Summaries could not be obtained for years previous to the earliest here noted. On the basis of those obtained it is shown that northern Illinois has more favorable weather in March than southern New York. New York, it will be noted, is a State recently occupied by the lark. It is concluded that S or more inches of snow, lasting two or more days, would destroy a nest.
It is suggested that the discovery of nests during nest building is possible by locating first the calling or singing male. At this period the male will be attending the female closely and she will be discovered shortly. The status of nesting can always be determined by the actions of the female. During nest building she is very restless, runs here and there, flies up and away, but shortly returns. Eventually she may disclose the site of the nest excavation. These reactions are instinctive responses to the desire for nest concealment. All nest building seems to be done by the female.
During egg laying the discovery of a nest is at best accidental. Neither male nor female has been noted to approach a nest during this period. They express no solicitude beyond that of nest concealment, thus displaying a remarkable nonchalance, especially on the part of the female. This reaction is so marked that an observer can nearly always be assured of the status of nesting whenever it is noted.
When incubation has begun the behavior is very different, as is also the behavior after the eggs have hatched. These reactions will be noted later. During these periods nests may be located by a systematic search that involves driving the male about until the female is noted. She will flush from the nest and the male will go to her. Then a patient watch of the female will, after a variable length of time, disclose the nest. When young are being fed the male will, at times, disclose the nest much more quickly than the female, for he assists in feeding and has nest-concealing instincts that are very poorly developed. Though the nest of the prairie horned lark is never concealed from above, it fits its semibarren environment so closely that a promiscuous search over a breeding territory is nearly always tiresome and unavailing. An incubating or brooding lark, as will be discussed later, often remains close to her nest on a chilly day or very early in the morning or toward evening. Nests can be found under these circumstances by a systematic search of Likely habitats and so flushing the bird from the nest.
No evidence of the use of a natural depression was noted either at Evanston or at Ithaca; all were dug by the female. According to Sutton (1927) and my own observations of 0. a. 8trngata in western Oregon, this excavation is dug with both beak and feet. The nest is constructed usually at the edge or partially under a grass tuft or clod, which, in the case of the prairie horned lark, lies most frequently on the west, northwest, or north, possibly because the cold and violent winds of the early nesting season come from this direction. The body of the nest consists of coarse stems and leaves with a finer lining within. The time spent in nest construction varies from two to four days.
The majority of the nests of the prairie horned lark showed a variable amount of clods, pebbles, or similar items laid about the margin usually on the side away from the protective tuft or clod. These so-called “pavings” were always composed of the material most easily obtained regardless of its permanency. It is suggested that the purpose of “pavings,” if there is a purpose, arises from the method of nest construction and from the desire of the larks to have a bare-ground nest approach.
Eggs: The egg has a background of gray with an occasional greenish tinge, which background is almost completely concealed with a fine speckling of cinnamon-brown. The cinnamon-brown often forms a denser ring about the larger end. The average size was found to be 2.25 cm. by 1.55 cm. The eggs of natural second sets seemed to be a trifle larger than first sets of the same individual. The number of eggs per set varied from two to five; the average was about four, the sets of fewer numbers occurred early, those of larger number, later. [ArmoR’s Non: The measurements of 50 eggs average 21.6 by 15.7 millimeters; the eggs showing the four extremes measure 24.6 by 15.5, 23.1 by 17.3, 18.3 by 15.0, and 21.6 by 14.5 millimeters.]
Incubation: The incubation period was determined to be 11 days. Only the female incubates.
The male shows little or no solicitude during the incubation period. The female has a highly developed series of automatic instincts of solicitude, which are modified by time of day, condition of weather, and frequency of disturbance. The most highly developed and probably the most recently acquired of these has been given the name ‘nest concealment by abandonment, or casual abandonment. The female leaves the nest, in this reaction, when an intruder is at a long distance, and flies quietly away, low against the ground, and does not show other solicitude for a very considerable period. The distances of the intruder from the nest during this reaction vary from 25 to 100 yards or often farther, a greater distance, it will be noted, than would disturb even a timid lark under other circumstances. A reaction that in many ways is the reverse of this, but still a marked exhibit of solicitude, is that called distress simulation. This consists of a precipitate flushing and rapid flutter over the ground after the nest has been approached closely. This reaction would be given most frequently on very cold da5’s; in the dusk of very early morning or evening, and when the bird was flushed very shortly after she had returned to the nest. It is certainly more primitive than the first reaction here described and is probably a culmination of the more frequent distraction display that most birds present when their nests are disturbed. Between concealment by abandonment and distress simulation there occurs a complete gradation, which, since the reactions are exact opposites-in expression, involves a curve that drops from the first to the zero point and then rapidly ascends to the expression of the latter. Thus, between the two, lessened expressions of either reaction would result, with a curious hiatus midway in which the incubating bird would allow an intruder to approach closely and then leave without an expression of either type of solicitude. Experimental flushing of an incubating bird from a blind showed that the bird, in one case, would give distress simulation if flushed in an interval that was less than two minutes from the time of her return; but would give casual abandonment if flushed after an interval of five minutes. A female lark, shortly after being forced from a nest, would express her agitation by aimless ground pecking, and, to be sure, would eventually be driven by the incubation urge to return to tlie nest even though an intruder might be much nearer than he had been when the nest was originally abandoned. This complex of instincts involved both the urge to incubate and the urge to protect. The instinct to protect, by whatever method, would all be overshadowed in time by the instinct to incubate.
Young: With the exception of those nests of early April, in which incubation began before the set was complete, all young hatched within an hour or two of each other. The young are fed within an hour or two following hatching. In most cases the male assisted the female in feeding the young. In carefully observed cases he visited the nest less often but brought greater burdens and fed more young at a visit than did the female. A total number of observed feedings during one day (April 30, 1926) was 108. The male fed 39 times and the female fed 69 times.
Observations of the adults and dissection of a few nestlings showed that some vegetable matter (weed seeds) is fed early in spring, but that even in March most of the food is animal matter. Later in the season grasshoppers become conspicuous in the diet. The adults dig up both cutworms and earthworms.
The male shows solicitude for the nest and its contents for the first time after the hatching of the eggs. His solicitude is restricted to calls. The female will leave her brooding in typical concealment by abandonment when conditions are appropriate as when incubating; likewise she will go from the young in distress simulation under conditions as noted previously. Proportionately the number of concealments by abandonment decreases and distress simulations increase slightly with young in the nest. Other reactions, which are various primitive expressions of solicitude, or intermediates of the two just mentioned, increase proportionately. Perhaps a return of more primitive instincts indicates a sum total of greater solicitude. Since the female is frequently absent from the nest in food foraging, she will come in, as an intruder approaches, with calls and cries. One or two references in the literature show that the reactions to dogs is the same as to man, but that liens are driven off by entirely different methods.
The larks removed all excreta throughout the full extent of nest occupancy. Early in the season much of the exoreta was eaten by the adults; later it was dropped to the ground 50 or more feet away. This seasonal change of habit may have been related to the available food supply. The instinct compelling excreta removal proved itself very powerful, at times overcoming strong solicitude for nestlings and even fear.
The young showe4 a psychic development closely related to their rate of growth and not to their age. Young of the same age or but one day younger than their nest mates often presented a psychic development two to four days behind them. This was due to uneven feeding, which occurred frequently early in spring because of uneven hatching or an inadequate food supply.
Normal nestlings give a food response indiscriminately up to the fifth or sixth day. Just prior to this time their eyes open. Following this they respond not at all or momentarily only. They withdraw at a touch from the hand on the sixth day and sink back quietly into the nest in crouch-concealment between the seventh and ninth days. Upon being removed from the nest at this age they sit quietly upon any object upon which they are placed; prior to this time they wriggle about when taken from the nest. They leave the nest on the tenth day and then express fear by hopping and calling wildly when disturbed. An expression of this type of fear, prior to the tenth day, would take them from the nest prematurely.
Weight-growth curves show a gradual increase over the first three days, a very precipitate rise (except for April nestlings) for the next three or four days, a marked leveling during the seventh and eighth (in one case the sixth), and a gradual rise during the ninth and tenth. Nestlings in May grew slightly more in the same period than a nestling in June and much more than a nestling in April. This discrepancy of growth seems closely (though perhaps indirectly) correlated with the temperatures of these seasons.
A lessening in weight growth occurs, normally, between the seventh and eighth days. This is brought about by the simultaneous unsheathing and drying of most of the feathers. On the other hand, growth in length shows, if anything, an acceleration at this period due to the extension of the tail. Marked variations in growth occurred in the various broods measured and in the different young of the same brood. This was brought about by two things: The fact that a slight difference in age gave the older larks a great advantage in securing food from the parents; and the fact that food was more plentiful later in the season than at the beginning.
Length-growth curves show a precipitate rise during the first three days, a slight leveling during the next three days, and a precipitate rise during the sixth, seventh, eighth, ninth, and tenth days. The cause for the intermediate leveling is not understood, but the rise toward the end of the nesting period is brought about by the growth of the tall.
In the early breeding season the enemies of the young are weather and a scanty food supply. The weather may result in snows that bury them from sight. The scanty food supply may result in the starvation of one or more of the nestlings. Starvation results from the automatic feeding reaction of the adults wherein the nestling nearest that part of the nest habitually approached by the adults will receive the first feeding; if the food is scanty this bird will receive all or nearly all the food. Only when food is so abundant that the first nestlings fed do not swallow promptly will the remainder of the brood be fed. In this case food is withdrawn from the mouth and put in the next and so on. The female lark rarely brings more than will go into one mouth; the male may feed two or more, but never four or five, at a time. Those young larks that have a few hours’ advantage in hatching: a full day in several cases in the early spi~ing: will have the advantage in size that will allow them to push to that side of the nest over which the food always comes. They survive; the others may perish. Such occurred in many observed nestings in April.
Predacious enemies cause a greater and greater loss as the season advances into June and July. The optimum season for the welfare of the young is shown to be May.
One case of cowbird parasitism was observed and followed. A lark, which hatched before the cowbird, came to maturity. The cowbird probably did not. It is suggested that the early nesting season and the exposed habitat may mitigate against such parasitism as may also the early departure of the young larks from the nest. however, since the adult lark will tolerate the parasitism and the food of June and July is suitable, other reasons prevent more extensive parasitism at this later season.
The young leave the nest, normally, on the tenth day, some three to four days before they can fly. Their protection during this interval is silence and a very effective “freeze” cxr crouch-concealment. Their plumage is remarkably adapted for this. The actions of the parents, especially the female, with her abandonment concealment, are calculated to take advantage of t.he protective color of nest and young at all ages.
Young leave the nest usually by following a parent that has brought food. One case was noted wherein a female enticed a belated nestling from the nest with a food morsel. The young fly in about five days after leaving the nest. They hop for some days after nest leaving, whereas the adults walk. This hopping may be anatomical or an atavism.
Plumages: The recently hatched nestlings are rather heavily covered with down, a necessary protection against sun and cold in their exposed location. The down is cream-buff in color. At nest-leaving age the young lark is in full juvenal plumage but presents an appearance quite unlike that of the adults; each feather of the upper surface has a triangle of brown at its tip, the under surface is white except the throat, which is gray. [AUTHOR’s NOTE: Subsequent molts and plumages are described under the northern horned lark.1 Food: McAtee (1905), in his extensive account of the food of the horned larks, writes that in August and September many grasshoppers are taken (7.1 and 8.9 percent of the total food, respectively) and that weevils constitute 18 percent of the food in August.. He says further that spiders are taken in every month. The conspicuous weed seeds that he lists (foxtail grasses, smartweeds, bindweeds, amaranth, pigweeds, purslane, ragweed, crab and barn grasses) are probably largely consumed in fall, winter, and early spring. The total of 79.4 percent of vetetable matter taken in the year, as given by McAtee, is made up largely of these weed seeds. He found about 40 percent of food taken in August to be animal matter, 20 percent animal matter in September, between 10 and 20 percent in October, 5 percent or less in November, about 2 percent in December, 1.73 percent in January, and 3.11 percent in February. The animal matter of January and February consisted principally of weevils and cocoons of tineid inoth~. Grain, chiefly waste oats, corn, and wheat, formed 12.2 percent of the food of larks, exclusive of California forms, and much of this would have been taken in the period under consideration.
The Main Subdivision at Evanston, where the most extensive observations were made by the writer, had, in the winter of 1925: 26, great quantities of Agropyron repens (quack grass), Setaria (foxtail), and Arnaranthus (pigweed), all of which had been allowed to mature seeds. Of this the seeds of the quack grass were eaten first and wherever their long stems had fallen over the sidewalks the larks would invariably be found in January and February. When quack grass failed, foxtail was eaten, and lastly Amaranthus was substituted when no other seeds were available. Once or twice larks were noted along the roads feeding on the oats of horse droppings, when snow covered all the weed seed of the subdivision. And again at Ithaca the compost heaps, put out for fertilizer along the garden margins, supplied some food when snow lay deeply over the ground.
At Ithaca, during the spring of 1928, prairie horned larks were observed feeding on Setaria (March 1), on Ambrosia artemi~iaefolia (April 1). A pair of larks were frightened away from an arctiid moth larva (Apantesi8 arge), which I observed the female dig up (March 3). Finally a few adults were colleeted in March at Ithaca (Connecticut Hill), and examination of the stomachs of six individuals showed that the vegetation consisted of oats, Setaria, Ambrosia arternisiaefolia, and waste buckwheat.
In summary of the feeding habits of the prairie horned lark in autumn, winter, and early spring, all that need be said is that the bulk of food taken is that of weed seeds, and the animal food, a much smaller proportion, is almost entirely of those forms harmful to the agriculturist. The lark, in feeding habits, finds for his food those things that appertain to the waste lands he inhabits.
Behavior: Breeding birds, such as females in abandonment concealment of the nest, or males in flight song, exhibit several distinct flights, but at other seasons the flight is of but one definite character. This is a choppy undulation brought about by three or four rapid, even strokes of the wings interrupted by the space of about two beats during which the wings are closed. A note is uttered on the climb of each undulation. Or again, on prolonged flights, the character of the wing beat is as follows: Long strokes are made, one, two, three (or one, two), with a pause of about one wing beat between each stroke wherein the wings are folded. Then come four to six rapid and successive strokes, which cause a climb. At this time the note zee t-it is uttered. Then comes a pause of the length of one or two beats, with wings folded, causing a drop in elevation. These repeat. The bird goes thus: jump, jump, jump, climb (call also), drop, jump, jump, jump.
Voice: The horned lark, like the goldfinch, usually advertises itself in flight by a definite, unmistakable note. Except for an occasional song, this is about the only sound from the birds in fall and winter. The flight and call notes are several in number, some of them appertainiug more especially to the breeding season than to wintering birds, and in that connection they will be considered again. The chief stock in trade of the lark and the one most commonly heard is p-deet or merely zeet. It is uttered casually on the climb of the ordinary undulating flight, especially on long journeys and in flights of young birds. Adults frequently make low flights over the ground without uttering a note. This p-seet is occasionally, sometimes frequently, lengthened to p-seet -it during the flight. When flushed the note is zu-u’eet or zur-reet (long drawn), zeet-eet-it, or zeet-it-a-weet, which is so high-pitched and mournful in character that it makes the birds indeed a part of the winter’s gale that whips them away.
The season of song extended, in the Evanston region, from midJanuary until early in July; in the Ithaca region from mid-February to late in June. With ifight songs used for a criterion, it was found that May was the optimum month. The lark sings both from the ground and in the air, under all conditions of weather, though flight songs are most numerous on quiet, mild days, perhaps a little more numerous when the sky is overcast than when it is clear.
The most vigorous period of song extends through nest building, egg laying, and incubation. Perhaps of this period that portion of it when the female incubates allows most song from the male, since he attends the female carefully during nest building and egg laying. The period of least song occurs when the young are in the nest, for the male assists in feeding. Ground songs are regularly distributed throughout the entire day; flight songs seem to be most numerous toward noon and near sundown.
For three months the prairie horned lark is the only singing bird in the open field; but with the coming and establishment of other migrants late in May and in June many other songs will be heard in that region. On June 16, 1926, at Evanston, the prairie horned lark, the last to begin singing that morning, went into flight song at 4:00 A. M. However, the lark almost always closes the singing at night with a long period of recitative which in mid-June, would not close until after 8:00 r. M.
The literature contains several descriptions of the flight song of the prairie horned lark, that of Langille (1S84) seeming to be most accurate. He describes the flight. The song he describes as “quit, quit, quit, you silly rig and get away.” This is the intermittent type; nowhere in the literature has a description of the recitative been found.
Songs are sung from the ground, from a clod or any other slight elevation, the greatest elevation noted being the roof of a sample apartment put up on the Evanston area; and from the air. The ground songs are similar to the flight songs though rarely as long or as systematically presented. The urge to flight song may come at any time or after an invading male lark has been evicted from occupied territory. Larks will also go into flight song upon approach of a human being or they can be forced to go up by driving them for a time about their territory.
The climb to flight song is distinctive and usually executed without a sound from the bird. The songs, in the air, are of two types: A recitative or rapid monotony of notes usually uttered at the beginning of the flight song, though occasionally at other periods, never over a few seconds in duration, accompanied by a steady beat of the wings; and an intermittent song uttered while the lark sails, about two seconds in duration, followed by a somewhat longer silent period during which the lark flutters up. The recitative can be transcribed as pit-wit, wee-pit, pit-wee, wee-pit; the intermittent as pit-wit, pitwit, pittle wittle, Uttle, litle, le~eeee. Large circles are described overhead during the flight song, or the bird heads into the wind if it is strong. The lark closes flight song by a headlong drop to earth with wings tightly folded.
Female larks seem to be unaware of the males in flight song, though other males note the bird overhead. The territory a bird may occupy in flight song is very extensive. Never were two visible birds noted in such a performance simultaneously. The one in the air is left undisturbed though his performance may carry him over many other lark breeding grounds below. Breeding territories are not vertical for a distance above a few feet; the flight song territory is something quite different.
Of several methods employed to determine the heights of larks in flight song, the most accurate was found to be the use of a binocular with an ocular scale. It was determined thus, through measurement of 25 songs, that the lark sings from elevations that vary from 270 to 810 feet. The average was 464.4 feet. Differences in height seemed to be individual variations or due to weather. Thirty times flight songs varied from one minute to five; the average was 2.34 minutes. Intermittents, regularly given, averaged 11.9 a minute.
An Evanston bird sang from song posts on the ground, which, during one entire day, varied a few feet from the incubating female out to 100 yards. The average was 38.66 yards. Ithaca birds, with bigger territories, sang frequently as far as 150 yards from the nest.
Fall: Young larks flock shortly after nest leaving. If the breeding ground has become untenable owing to vegetation, they seek other regions. Flocks grow larger through addition of adults in August and September and then smaller as migration begins. In flight the flocks are comparatively compact, but they spread widely when the birds alight to feed or pass the night. During autumn and winter they occupy regions essentially like those in which they breed in March and April, that is, semibarren or almost denuded areas, which may be natural or due to some seasonal condition of agriculture. The Lapland longspurs and the shore larks (Otocoris alpestr& alpe8tr~8) are the only other birds that occupy a habitat with conditions just like those in which the prairie horned lark occurs in fall and winter.
Subspecies of the horned lark vary from such highly migratory forms as the northern horned lark to such strictly sedentary subspecies as the California horned lark. The prairie horned lark is intermediate between these extremes and leaves its breeding grounds for a period of one to two months during winter. This bird breeds north to the southern edge of Canada and migrates south to South Carolina, Kentucky, and Texas. From the northern part of this range it is absent during the month of December and part or all of January. Throughout the remainder of the breeding range some individuals are always present.
TEXAS HORNED LARK
OTOCORIS ALPESTRIS GIRAUDI Henshaw
Along the coast of Texas from Galveston Bay to the iiiouth of the Rio Grande, and for a short distance into Tamaulipas, Mexico, we find this race of the horned lark, rather widely separated from its nearest relatives and usually on the salt marshes or not far from the seacoast. It seems to be most closely related to the prairie horned lark. Dr. Oberholser (1902), in comparing it, says: “This race is quite similar to praticola, though considerably more grayish, rather smaller, and with the yellow of throat usually deeper and suffusing also the superciliary stripe. In winter plumage the dark streaking on the breast is frequently heavier. It is fully as gray above as arcticola, but is of course easily distinguishable by its reduced size and yellow of throat and eyebrow. It is so much smaller and more grayish than either koyti or alpestri.s that it does not need special comparison.”
W. E. Grover wrote to Major Bendire (1895) : “The Texan Horned Lark is locally here known as ‘Chippie’ and ‘Road Chippie’, as it is essentially a ground bird. It frequents the level, grassy prairies along the Gulf shore, and may frequently be observed in the wagon roads; hence its local name. I do not know how early it arrives in this vicinity [Galveston]; I noticed a few on April 1, and by May they are abundant. The nest is built in a saucer-shaped hole scratched out by the birds, and here it is nearly always placed alongside of bunches of wild chamomile (Matrica’ria ocronata) growing close to the road; it is constructed of dry prairie grass and lined with thistle down. The top of the nest is even with the surrounding ground.”
Bendire goes on to say: “All the nests of the Texan Horned Lark I have seen are much more substantially built than any of the balance of the subspecies breeding within the United States.” He says of an unusually bulky one: “It was placed in a pile of dry cow droppings near the shore of Aransas Bay. The outer walls are chiefly constructed of salt-cedar twigs (Monanthochloe littoralis) , and the lining consists of dry sea moss picked up on the shore. It measures 6 inches in outer diapieter by 2½ inche~ in height. The inner cup is 3 inches in width by 2 inches in depth.” An average nest is only about 4 inches in outside diameter, “and it is sparingly lined with blades of dry grass, and a few feathers.”
The Texas horned lark lays ordinarily three or four eggs, which are practically indistinguishable from those of other horned larks of similar size. The measurements of 40 eggs average 21.5 by 15.7 millimeters; the eggs showing the four extremes measure 23.4 by 16.0, 22.6 by 16A 19.7 by 15.3, and 21.2 by 14.9 millimeters.
STREAKED HORNED LARK
OTOCORIS ALPESTRIS STRIGATA Henshaw
The above name applies to the horned larks that breed in the Pacific coast belt of Washington and Oregon, west of the Cascade Mountains, and southward to Siskiyou County, Calif.
Dr. Oberholser (1902) writes: “This race differs from merrilhi in much smaller size, deeper and more extended yellow suffusion below, and in the decidedly more brownish color of the upper parts. In autumn and winter, when merrilhi is often brownish above and shows sometimes as much yellow below as strigata, size is the best means of identification. In color it much resembles alpestris, but in summer the back is more blackish, in winter the yellow suffusion is more extensive, while its smaller size will of course distinguish it at all seasons. It differs from lwyti as from alpestris, with the additional character of a deep yellow eyebrow.”
While I was in Seattle, in 1911, Samuel F. Rathbun showed me the breeding grounds of the streaked horned lark, of which he says in his notes: “Almost invariably this lark frequents what are known as gravelly prairies. The gravelly prairies are distinctive. They are of rather limited extent, and have a soil that is mostly of fine, smoothly worn gravel, which at times represents the greater part of the soil. In spring and early summer these prairies are strewn with wild flowers, but when the flowers disappear the prairies look more like sterile areas.”
J. H. Bowles (1900) says that “around Tacoma these little larks are extremely local in their distribution, large areas of prairie being altogether untenanted, while an exactly similar piece of land will be swarming with them.” They nest commonly on the grounds of the Tacoma Golf Club. “The surrounding prairie extends for miles where hardly a dozen pairs of the birds can be found in a day’s walk, while on the links last summer I estimated that fully one hundred pairs must have nested. Indeed, so sociable are they that only an occasional nest is placed more than a few feet from the ‘putting green’ or the ‘tee-off’.”
Nesting: Mr. Bowles (1900) says: “The location of the nest may be almost anywhere on the ground, but the soil must be extremely dry. As a rule the birds scratch out a hole for themselves about two and one-half or three inches deep, both birds working, but I have found nests in the hoof prints of cattle, in cart ruts, holes made by dislodged stones and one that was placed in an unused golf hole.” One very large nest was “well lined with grass, fir needles and feathers.”
Mr. Rathbun (MS.) says of several nests that he has found: “Each time it was in a slight depression of the ground on the dry, open prairie, oftener close to the edge of a dried cow dung, and just within or against the growth of grass, which would be more dense in such a spot. A nest found in very early June was made of dry, gray mosses, bits of dead weed stalks, dead grasses, with various soft substances of plant kind, and it was neatly lined with fine, dry grasses. Its dimensions were: Outside diameter, 4 inches; inside diameter, 21/2 inches; height, 2 inches; depth, 11/4 inches.”
Eggs: Three or four eggs constitute the usual set for the streaked horned lark. These are practically indistinguishable from those of other horned larks of similar size, with the usual variations. The measurements of 32 eggs average 21.0 by 15.8 millimeters; the eggs showing the four extremes measure 23.4 by 16.9, 19.0 by 15.7, and 20.0 by 14.9 millimeters.
Enemies: Mr. Rathbun writes to me that the “crows used to pester the breeding larks, for the grass was so short when and where these were nesting that many a nest was broken up by the crows. Several times I watched the black rascals carefully working over the nesting areas, and I found the torn nests and now and then an eggshell.”
DUSKY HORNED LARK
OTOCORIS ALPESTRIS MERRILLI Dwight
In naming this race in honor of Dr. J. C. Merrill, who collected the type near Fort Klamath, Oreg., Dr. Dwight (1890) described it as “larger, more broadly streaked above, and blacker than 8tr2gata, with less yellow about the head and throat, the nape pinker.* * * This is the blackest-backed of all the races, the dark brown of 8tr~gata having a decidedly yellowish shade, particularly in autumn specimens, whereas merrilli is black-brown in spring and strikingly grayish and streaked in autumn.”
Its breeding range lies chiefly in the Transition Zone from southern British Columbia, through Washington and Oregon east of the Cascades, and from extreme northern Idaho to northeastern California and northwestern Nevada.
Major Bendire (1895) says of its haunts: “This subspecies is essentially a bird of the foothills (the so-called ‘bunch grass country,’ Featuca sp. ~) as well as of the more open and grass-covered valleys and plains occasionally found in the mountains, while it is either rare or entirely absent in the more arid sagebrush plains found interspersed through the same regions.”
Dr. Walter P. Taylor (1912) says that, in northern Nevada, these “horned larks exhibit a very marked preference for the vicinity of the fields and dry meadows, as along Quinn River. The birds were frequently encountered, however, on the most inhospitable deserts, although they were more numerous in pleasanter surroundings. We did not observe them at a greater altitude than ‘1000 feet, although Ridgway noted them as high as 11,000 feet.”
Referring to the Lassen Peak region in California, Grinnell, Dixon, and Linsdale (1930) write: “Horned larks in the eastern part of the section were seen on ground where the cover of vegetation was slight. The birds frequented both cultivated land and unbroken land. Areas of sandy as well as hard ground were foraged over. * * * Single birds were sometimes seen perched in the tops of sage bushes. On July 25, 1928, * * * many horned larks were seen among small and scattered sage bushes. During the hot mid-day hours the birds sought shade beneath these small bushes.”
Nesting: C. E. McBee (1931) has found the dusky horned lark nesting very abundantly on the grassy plateau and on the cultivated lands about his ranch in Benton County, Wash. He gives an excellent description of the nests, which I quote:
The nests of the larks are built in a cup-shaped hollow scratched out by ithe birds, with the top of the nest even with the level of the ground. They :are composed of various kinds of vegetable material; sometimes small rootlets, ~pliable twigs, or pieces of a soft kind of plant which grows on the plateau, ~but more often a good share of the nest Is made up of the thin coverings of ‘wheat stubble and pieces of grass. In only two instances has the writer found ;any evidence of animal material; these were both small pieces of rabbit fur insed as lining. The birds place the nests in the shelter of a clod, small plant, ~or bunch of stubble. The prevailing spring winds are from the southwest and the nests are always placed directly opposite, so that they are open to view from the northeast. This is an Important point to remember when looking for nests; It Is much easier to find them while traveling in a southwesterly direction. Although the birds seem to prefer wheat stubble fields for nesting sites, the writer has found nests in plowed ground, in freshly seeded fields, and in grassy sod of abandoned ranches.
Eggs: IHe says that “the number of eggs to a set is either two or three: very seldom four.” In five years he found “only three sets of four eggs, and the markings on them lead to the belief that one of them was deposited by another female. This was true in each set. Three is the common number, although late sets sometimes contain only two eggs when complete.”
“The eggs,” he continues, “vary a great deal in size, shape and coloration. The shape varies from ovate to elongated ovate, as a rule, and an average egg measures about .82 by .62 inches. The ground color varies from light gray to, rarely, almost white, sometimes faintly tinged with greenish. The markings are usually dots and small spots that vary in color from dark gray to light brown, sometimes sprinkled rather evenly over the entire egg, in others with a distinct ring around the larger end.”
Some eggs of this race that I have seen have a yellowish-white ground color, with a wreath of “buffy brown” and “dark olive-buff” spots around the center of the egg, otherwise finely sprinkled with the same colors. The measurements of 35 eggs average 21.5 by 15.4 millimeters; the eggs showing the four extremes measure 24.4 by 15.8, 21.8 by 16.5, 20.1 by 15.2, and 20.2 by 14.0 millimeters.
Enemies: Mr. McBee (1931) says that “natural enemies of the birds are probably many. Ravens undoubtedly account for some; snakes for many young birds of the hopping age, and the very plentiful ‘Sagerat,’ which travels far and is believed to eat eggs, destroys some.” But, as the birds prefer to nest in stubblefields, spring plowing “leads to the destruction of untold hundreds of nests, eggs and young birds.” He continues:
Allowing a minimum of One nest to every two acres, which the writer believes Is too conservative, various operations during the nesting season on the McBee ranch alone would account for the destruction of some three thousand nests. There Is no possible way of preventing the destruction, except by rescuing those nests which happen to be noticed by the tractor or team driver. Most of the farmers are ~cqualnted with the fact that the larks are valuable Insect eaters, so will not knowingly destroy them. It is a sight, indeed, to see an expensive tractor outfit, or a twenty-horse team, stopped while the driver leaves his place to set a tiny nestful of eggs or young birds to a place of safety. Observations by the writer tend to show that in most cases the parents will return to their young, even though they may be moved several feet; sometimes to heavily Incubated eggs, but very seldom to fresh or incomplete sets of eggs. They will also Immediately start rebuilding their destroyed nests.
Winter: The dusky horned lark is present all the year round in eastern Washington and Oregon. In winter immense flocks of this and the pallid horned lark (arcticola), with perhaps some other forms of the species, form one of the most conspicuous features in the bird life of the open plains. Mr. McBee (1931) writes:
Weed seeds and kernels of wheat are the main articles of food for the larks, and while there Is no snow, these are easily obtained. However, when the ground is covered to some depth, the birds are forced to pick off the seeds from the tumbleweeds which protrude. Soon after the snow falls it is stained In the vicinity of protruding weeds by dirt and pieces of stalk which are thrown off by the birds in the process of obtaining seeds. When all visible seeds have been eaten, hunger forces the larks to the strawstacks, pig-pens and barns of the ranches. During the heavy snow of the winter of 1929: 30, the writer fed the birds cracked wheat and succeeded in tolling them to within a few feet of a large bay window in the ranch-house, where for two days they fed In great numbers. Many interesting observations were made during that time of the feeding habits of the birds. Especially amusing was the occasional display of a terrible temper by the larks. A few members of the flocks were bullies of the first water and absolutely refused to allow any other birds to feed on the square patthes of ground which were cleared for them. One especially bad tempered bird succeeded in keeping for its sole use an entire feeding plot, almost three feet square, for upwards of an hour. At times fifty or sixty birds would be sitting around in the deep snow surrounding the feed, hungry almost to the point of starvatIon, feet heavy with droplets of Ice, but more content to sit and suffer rather than brave the wrath of the terrible tempered one. Newly arriving individuals, upon alighting in the cleared space, soon found the lay of the land and joined their companions in the snow.
Professor Beal (1910) makes the following tribute to the hardiness of horned larks:
The writer has met them on an open prairie when the temperature was nearly 30 degrees below zero, and though a fierce gale was blowing from the northwest they did not exhibit the least sign of discomfort, but rose and flew against the wind, then circled around and alighted on the highest and most windswept place they could find. Probably they remain through the night in these bleak spots, for they may frequently he seen there after sunset. Most animals seek shelter from wind and cold, even though it be nothing but the leeward side of a ridge or hummock, but the horned lark refuses to do even this, and by preference alights on the top of the knoll where the wind cuts the worst. It seems strange that In so small a body the vital heat can be maintained under such adverse conditions, but if one of these birds be examined, its body will be found completely covered with a thick layer of fat, like the blubber on certain marine animals. This indicates that horned larks have plenty to eat, and that their food Is largely carbonaceous.
ISLAND HORNED LARK
OTOCORIS ALPESTRIS INSULARIS Townsend
The island horned lark is the small, dark race of the species that inhabits practically all the Santa Barbara Islands off the coast of southern California, though it is more abundant on some of the islands than on others. It was formerly supposed to be identical with the streaked horned lark and was so listed by some of the earlier writers. Dr. Oberholser (1902) describes it as follows:
Like Otocoris a. 8trigata, but darker, somewhat less ochraceous above, less yellowish on breast and abdomen. * * Nothwithstanding Dr. Dwight’s statement that he could not distinguish the Santa Barbara Islands birds from RtriDa.ta, they constitute an easily recognizable race which, though curiously enough most closely allied to 8tfigata, yet differs in the darker color above, particularly on cervix and bend of the wing; in the more grayish tone of the back and scapulars; the absence of yellow on the breast; and the much more conspicuous streaking on this part. * * * From- merrilii the Island bird differs In smaller and much more reddish coloration; while from ache of the adjacent mainland it may be separnted by Its conspicuously darker coloration throughout.
I made my acquaintance with the island horned lark on San Nicolas and Santa Barbara Islands, far off shore in the southern group of islands, which I visited as a guest of J. R. Pemberton in his power boat. San Nicolas is the outermost island of the group, over 60 miles from the mainland; it is a barren, windswept isle, about 7 miles long and 3 miles wide. It is mainly formed of soft sandstone, and the high cliffs that tower above the wide sandy beaches are broken by deep rocky canyons and are sculptured into all sorts of fantastic shapes by the erosion of sand and wind. The upper part of the island, about 800 feet above sea level, is a nearly level, or rolling, grassy plateau, which serves as grazing land at times for sheep and horses. Here we found the larks very abundant on February 24, 1929. They were mostly in pairs, or trios, flying about as if mating or preparing to nest. As we found no nests we were probably too early for eggs. We saw the larks also on the benches below the sandstone hills, where scanty grass and other low vegetation was growing. On the sloping sand dunes near the beaches there were many large clumps of iceplant in which the larks have been known to conceal their nests.
We landed the next day on Santa Barbara Island, about 35 miles from the mainland, much smaller and more precipitous than San Nicolas. From the only available landing place, a flat rock near a colony of barking sea lions, we climbed up a steep and narrow trail to the top of the island, which at its highest point is over 500 feet above the sea. The top of the island is a rolling grassy plateau on which herds of sheep were grazing. It is broken by numerous deep gullies, which were overgrown with a curious plant called the “tree dahlia”; it looks like a small tree with a thick woody stem and rough surface, often branching like candelabra, with a growth of leaves and a cluster of bright yellow, daisylike flowers at the end of each branch. The Santa Barbara song sparrows, which were very abundant here, are said to build their nests in these plants. On the hills and the open grassy parts of the plateau, the island horned larks were very common but apparently not yet nesting; at least we found no nests.
Nesting: T here are two interesting nests of the island horned lark, preserved with the surrounding vegetation, in the Thayer collection in Cambridge. They were collected by C. B. Linton on San Nicolas Island on May 12 and 14, 1910, and are accompanied by photographs. The first nest contained three eggs and was sunk into the sand under a thick patch of iceplant that was growing in a dense mass on the sloping side of a ravine and only about 50 feet from the ocean. The nest was made of fine grasses and weeds and was profusely lined with white feathers, apparently gull feathers. The pale eggs in the white nest, framed by the pale gray foliage of the iceplant, make a harmonious picture. The second nest, containing four eggs, was placed among a more scattered growth of iceplant on a sandy slope; it was made of very fine grass and bits of iceplant but contained no feathers.
Another nest in the same collection was taken by 0. W. Howard on San Clemente Island on April 4, 1905; this nest, containing four eggs, was located near the summit of the island near a cattle trail; it was compactly made of various coarse and fine weed stalks, fine grasses, lichens, bits of wool, and plant down and was lined with finer bits of the same materials.
Dr. Joseph Grinnell (1897) found a nest on San Clemente Island on June 3, 1897, that “was on the ground in a depression under the broad, obliquely-inclined leaf of a cactus. It was thus well-protected, as no fox could reach the contents without encountering the stiff spines.”
Major Bendire (1895) states that H. W. Henshaw found a nest on Santa Cruz Island on June 4, 1875, that was “placed within the cavity of an abalone shell, one of a large heap lying half overgrown with herbage. The whole cavity of the shell was filled by the material, and the eggs looked very pretty as they lay contrasted with the shiny pearly shells clustered about them.”
Eggs: The island horned lark lays three or four eggs, which are practically indistinguishable from those of the other western horned larks of similar size. What few I have seen are of the finely speckled pale type, but they probably show all the variations seen in eggs of the streaked horned lark and the California races. The measurements of 28 eggs average 20.7 by 15.7 millimeters; the eggs showing the four extremes measure 22.3 by 16.0, 20.9 by 16.2, 18.8 by 15.2, and 19.8 by 15.1 millimeters.
The sequence of molts and plumages, feeding habits, and general behavior are apparently similar to those of the neighboring mainland form, the California horned lark, from which this race probably originated, developing independently certain characters similar to those of the streaked horned lark, with which it was formerly confused.
CALIFORNIA HORNED LARK
OTOCORIS ALPESTRIS ACTIA Oberholser
The California horned lark was formerly considered to be identical with the Mexican horned lark (chr’ysolaema), but Dr. Oberholser (1902) has shown that it differs from that race and has given it the above name. He describes it as “similar to Otocori.s a. cAry8olaema, but upper surface paler, more rufescent; yellow of throat and head of not so deep a shade. * * * From rrubea, with which it intergrades in central California, actia differs in the much more pinkish tint of cervix, rump and bend of wing, as well as in its more grayish back which is usually in more or less abrupt contrast to the color of the nape.” It is somewhat smaller than chrysolaerna. Although there are other races of the horned lark in California, actia seems to enjoy the widest range, which extends from San Francisco Bay south to northern Lower California and east to the San Joaquin Valley.
James B. Dixon (MS.) says of its haunts: “This bird is a common resident in the salt-grass pastures and drier barren areas, from 2,600 feet elevation to sea level. Its favorite nesting locations are the dry, high humps in salt-grass cow pastures that are well grazed and do not have very much cover. They also nest in vineyards at the bases of the vines and in sparsely growing grainfields of all kinds. Their liking for closely cropped barren areas is well demonstrated by their nesting commonly on golf courses.~~ John G. Tyler (1913) says of its haunts in the Fresno District: “In former years, when large tracts of land north and east of Fresno were devoted to grain farming, the California Horned Lark was one of the most abundant birds to be found in the district; but it has not responded favorably to the settlement of the country and is now rare in many parts of the valley. * * * ~~ seems that for feeding and nesting these birds must have dry, barren ground almost free from shrubbery.”
Nesting: My experience with the nesting of the California horned lark has been limited to the finding of two nests on March 20, 1929, while driving over the open plains in southern Kern County with J. R. Pemberton. These rolling plains lie south of Maricopa and east of the Wheeler Ridge; they are covered with a scanty growth of short grass and are much grazed over in places by sheep, of which we saw some large flocks. The larks were abundant here and were mostly running about in pairs. We saw a female running about actively and feeding in a hurried, nervous manner. Mr. Pemberton said that she had just come off her nest to feed and would return to it within 10 minutes. We stopped to watch her, and in about 5 minutes we saw her settle down on some higher ground where there were a number of stones scattered about. We walked up and flushed her off her nest in a little hollow; it was a typical homed lark’s nest, made of coarse grasses and weed stems, mixed and lined with finer grass and a few bits of soft, cottony material; it held three well-incubated eggs. The other nest was found by flushing the bird, as we drove within 5 feet of it; it was sunk into the ground between two small pieces of dry cow manure; it also held three eggs.
Mr. Tyler (1913) says: “Nests of this species are built most often in summer-fallow fields, but sometimes in very young vineyards, hay fields from which the crop has been cut, and on the uncultivated plains. Sometimes they are found at the base of a clod or a small accumulation of trash, but in the majority of cases that have come under my observation a small weed or plant, frequently the California poppy, is chosen, probably more for the shade it affords than with any thought of concealment.”
Judged from the dates at which eggs have been found, this lark must raise at least two broods in a season and perhaps sometimes three.
Eggs: The California horned lark lays two to five eggs to a set, usually three. In 35 sets recorded by Mr. Tyler (1913), there were four sets of two, eight sets of four, two sets of five, and t.he others sets of three. The eggs are similar to those of other horned larks of similar size.
The measurements of 40 eggs average 20.2 by 15.3 millimeters; the eggs showing the four extremes measure 21.5 by 16.1, 21.4 by 16.2, 19.2 by 13.5, and 20.0 by 13.8 millimeters.
Food: According to W. L. McAtee (1905), “the food habits of the California subspecies (Otocoris alpestris actia) were found to differ so remarkably from those of the other horned larks as to merit separate notice. Briefly stated, the difference consists in the high percentage of vegetable: as compared to the animal: food consumed by the California birds.” Based on a study of 26’l stomachs, collected in every month but May, he says that the “vegetable food composes 91.44 percent of the diet of the California horned larks, while them larks in the remainder of the country take less than 80 percent of the same class of food.” McAtee continues:
Of the vegetable matter, weed seed, which is 51.1 percent, is less than the amount of the same kind of food taken by the other horned larks. The rest of the vegetable food, 40.2 percent, Is grain, including that from wild as well as from cultivated plants. * * * Of the grain eaten by the horned larks of California, 31.1 percent consists of oats and 9.1 percent of wheat, corn having been eaten by but one bird. Oats, then, are the favorite food, and on this account the horned larks are liable to damage the crop. However, a great part of the oats consumed probably comes from the wild plants so abundant In all parts of the State, and the destruction of these Is a benefit. * * * The California horned larks consume only 8.56 percent of animal food [ants, grasshoppers, and other insects], while the other forms collectively eat 20.61 percent. * * * It appears that the highest percentage of animal matter Is taken in June. This, however, is only 27.7 percent, not much more than half the highest monthly average for the other members of the species.
Professor Beal (1910) remarks in his summary: “In the final analysis of the food habits of the horned lark there is but one tenable ground of complaint, namely, that it does some damage to newly sown grain. This can be largely remedied by harrowing in immediately after sowing, and can be wholly prevented by drilling. The bird’s insect diet is practically all in its favor, and in eating weed seed it confers a decided benefit on the farmer. It should be ranked as one of our useful species, and protected by law and by public opinion. ~
MAGDALENA HORNED LARK
OTOCORIS ALPESTRIS ENERTERA Oberholser
The horned larks of central Lower California, from Santa Rosalia Bay to Magdalena Bay, have been given the above name. Dr. Oberholser (1907) describes it as “similar to Otocoris alpestris am’mop/tila, but smaller, the upper parts paler and more grayish, the cinnamomeous of nape, upper tail-coverts, and bend of wing more pinkish. * * * This new race is in color very similar to Otocoria alpestris leuoolaerna, but is more grayish above, at least when in good plumage; and has the eyebrow usually more yellowish; furthermore, the greatly inferior size of Otocaris a. enertera distinguishes it at once. From Otocoris alpestris actia, whose range it approaches most closely, it differs very much more than from either Otocoris a. ammo phila or 0. a. Zeucolaeina, being strikingly paler and more grayish throughout, as well as somewhat smaller.”
It seems to be a resident form on open ground, locally, in the Lower Sonoran Zone. Nothing seems to have been published on its habits, which probably do not differ from those of other southwestern subspecies of the horned lark. There is a set of three eggs in the Doe collection, taken by J. S. Rowley near Rosalia Bay, Lower California, on April 23, 1933; these measure 22.8 by 16.5, 22.3 by 15.8, and 21.9 by 15.6 millimeters.
RUDDY HORNED LARK
OTOCORIS ALPESTRIS RUBEA Henshaw
Dr. H. C. Oberholser (1902) says of the restricted range of the ruddy horned lark: “The present race seems to be strictly resident, occupying a comparatively circumscribed area in the region drained by the Sacramento River, passing south into actia at about the latitude of San Francisco, and northeastward into merrilli.”
This subspecies is described by Ridgway (1907) as “most like 0. a. actia, but much more rufescent, t.he occiput, hindueck, shorter upper tail-coverts, lesser wing-coverts, and sides of breast deep chestnutvinaceous or dark vinaceous-rufous, the back decidedly brown, broadly streaked with darker. Adult female most like that of 0. a. oaa~acae, but darker and browner above, with hindneck distinctly rufescent; similar also to that of 0. a. alpestris, but much smaller, back browner with spots less dark, and hindneck and sides of breast more rufescent.”
Dr. Oberholser (1902) says of it: “This form is easily distinguish. able from all the other horned larks by the peculiar color of the occiput and cervix, which is bright brick red with very little tinge of pinkish, particularly in summer; the remainder of the upper surface is much suffused with the same shade, further differentiating rubea from both insularis and strigata, which races in other respects it closely resembles.”
Grinnell, Dixon, and Linsdale (1930) say of the haunts of the ruddy horned lark in the Lassen Peak region: “The main habitat chosen by this race of horned lark within our section was the rocky mesa just above and bordering the Sacramento Valley on the east side [their cut shows a level grassy plain, with numerous rocks scattered over it]. Whenever visits were made to this ‘waste’ mesa land, in April, May, June, or December, this bird was conspicuous for its large numbers and for being almost the only one to live on that rocky type of ground. In winter the compact flocks would start up when disturbed and would fly so near the ground, below the horizon line, as to be nearly indistinguishable against the background of ruddy-hued earth and rocks.”
I can find nothing peculiar mentioned on the nesting or other habits of the ruddy horned lark, which are apparently similar to those of the closely related California horned lark. The eggs are also simliar to those of the neighboring race, though what few I have seen are rather more richly colored, more yellowish. Bendire (1895) mentions a peculiar set of eggs collected by Dr. Charles H. Townsend, that are “so suffused with rich reddish brown as t.o be unrecognizable”; this is evidently a case of erytlirism, which occurs occasionally in the eggs of other species. The measurements of five eggs, all I have been able to gather, are 20.8 by 14.7, 19.8 by 15.2, 18.5 by 14.5, 20.5 by 14.8 millimeters.
MONTEZUMA HORNED LARK
OTOCORIS ALPESTRIS OCCIDENTALIS McCall
Although this race of the horned lark was described and named as long ago as 1851, it was not recognized in the first two editions of our Check-list, published in 1886 and 1895, mainly because several good ornithologists confused it with other races. It was, however, included in the third edition in 1910, as Dr. Oberholser (1902) demonstrated that it is a recognizable race. He says of it:
The geographical variation exhibited by this race has been obscured, since Dr. Coues Included occidentalis In his leucolaema; Mr. Henshaw referred It to arenicola, and Dr. Dwight to a4uata; but the form is weu worthy of recognition * * * From adusta, to which it Is most closely allied, occidentalis differs in its much larger size and decidedly less ruddy colors above, the nape being more pinkish, the back more dusky. It is distinguished from ooxecoe by much paler, less rufescent colors above, and by decidedly larger size; from both act a and clLt-ysolaema by greater size, together with paler, more brownish coloration. Although of the same dimensions as leucolaema, this form may be separated by the darker, more cinnamomeous or rufescent shade of the entire upper surface, this in summer being particularly noticeable on the cervix; and these characters will serve to determine even doubtful specimens at all seasons.
In its nesting habits, molts, food, behavior, and other habits it probably does not differ materially from other southwestern races.
The measurements of 6 eggs, two sets of three, average 20.7 by 15.6 millimeters; the eggs showing the four extremes measure 21.3 by 14.8, 20.4 by 16.6, 19.9 by 16.1, and 20.7 by 14.8 millimeters.
SCORCHED HORNED LARK
OTOCORIS ALPESTRIS ADUSTA Dwight
In southeastern Arizona, from the eastern slopes of the Huachuca and Santa Rita Mountains, vast grassy plains extend eastward toward the valley of the San Pedro River; these plains are much like virgin prairies, pure grass lands, entirely devoid of other vegetation except for the straggling lines of cottonwoods, sycamores, and willows that mark the underground courses of the mountain streams that flow out from the canyons onto the plains at a few points, but they are hardly in sufficient numbers to make any appreciable break in the continuity of the prairies. These grassy plains form the best grazing lands in that region, but the cattle need vast areas of land to support them even here, and water must be provided for them from driven wells, windmills, and watering tanks, to which the cattle resort from miles around.
Here was the typical home of the scorched horned lark, as we found it abundant and apparently breeding in 1922, though we did not succeed in finding a nest. Similar plains extend westward for a long distance from the Chiricahua Mountains to the eastward, and as far north as Wilcox in Cochise County, where J. S. Rowley tells me that he found it breeding “rather abundantly.” A. J. van Rossem (1936) says that it “does not reach its western limit at the Santa Ritas, but continues west along the mesa lands to, and all along, the east base of the Baboquivaris.” Between this latter range, in Pima County, and the valley of the Colorado River there seem to be no suitable breeding grounds for horned larks, as they shun even a scanty growth of bushes or the ordinary desert vegetation.
In naming this race, Dr. Dwight (1890) describes it as “similar to clu-ysolaema [which he then included with what we now call actia], but of a uniform scorched pink or vinaceous-cinnamon above.”
Nesting: Mr. Rowley writes to me that, on the plains near Wilcox on May 24, 1936, he found these birds flying about and mating and located a nest in process of construction, “the female doing a lot of fussing around with a blade of grass in her bill.” Five days later, the nest contained three fresh eggs, and the female was sitting very closely. “The bird did her best to use what shelter was available by excavating under the side of a dried cow dung, to get even a little relief from the hot sun.”
There is a nest of this horned lark in the Thayer collection in Cambridge that was collected by 0. W. Howard in Cochise County, Ariz., on June 10, 1902, and held three eggs. It was placed at the edge of a tuft of grass and was made entirely of very fine grasses, mixed with pappus down; in its somewhat flattened condition, it measures nearly 4 inches in outside diameter.
Eggs: Apparently the usual set consists of three eggs. The three eggs in the Thayer collection are elliptical-oval and have very little gloss. The ground color is dull white, and the eggs are finely and lightly dotted and spotted over the entire surface with very pale “olive-brown”; on two of the eggs there is a concentration of spots of darker brown and drab in a wreath about the larger end. Probably a larger series of eggs would show the usual variations in colors and markings seen in the southwestern races of the species. The measurements of 31 eggs average 21.4 by 15.7 millimeters; the eggs showing the four extremes measure 23.2 by 15.5, 20.9 by 16.6, 19.4 by 14.8, and 19.8 by 14.4 millimeters.
Fall: Referring to the vicinity of the Huachuca Mountains, Harry S. Swarth (1904) writes: “Toward the end of July and early in August, young and old gathered together in immense flocks, and were at this time very restless and difficult to approach, flying a long distance when disturbed. They seemed to depart for the south soon after, for on September 5, 1902, on a drive of over twenty miles over country in which they had bred in the greatest abundance, not a single Horned Lark was seen.” Later he says (1929): “In the fall there proved to be but a small proportion of adusta among the enormous flocks of horned larks that frequented the plains.” The predominant number of oacidentalis at that season indicated “a deserting of the breeding grounds by adusta during the winter months.”
MOHAVE HORNED LARK
OTOCORIS ALPESTRIS AMMOPHILA Oberholser
In describing and naming this subspecies, Dr. Oberholser (1902) states that it is like 0. a. actia but “may be easily distinguished by its very much paler color above, while its decidedly smaller size, conspicuously more cinnamomeous shade of nape, upper tail-coverts and bend of wing render it readily separable from leucolaemz. Compared with occidentalis it is paler, of smaller size, with the cervix, upper tail-coverts, and bend of wing more cinnamomeous, the upper surface less uniform. It is somewhat smaller than adusta and paler, particularly in winter, with the back scarcely or not at all reddish, the demarcation line between cervix and back usually well marked. * ï * The young of ammo phila differ markedly from the young of actia in their paler, much more grayish upper parts; being practically indistinguishable from leucolizema of the same age.”
This subspecies occupies, in the breeding season, a rather limited range in the desert regions of southeastern California, from the Mohave Desert northward to Owens Valley. Dr. Oberholser (1902) says: “This desert race seems to be most typical in the region immediately southwest of Death Valley, California, whence a good series of specimens was brought back by the Death Valley expedition of 1891. The breeding birds in this series were identified as arenicola (= leucolaema) , the winter specimens as Chry8olaelna (= actia) .”
I can find nothing in the report of this expedition, or elsewhere, to indicate that the habits of this subspecies are in any way different from those of the other desert-loving races.
SONORA HORNED LARK
OTOCORIS ALPESTRIS LEUCANSIPTILA Oberholser
From extreme southern Nevada southward along the Colorado River in western Arizona and extreme southeastern California and into northeastern Lower California, we find this extremely pale desert race. In describing it, Dr. Oberholser (1902) says: “This new raco is the palest of all the American horned larks, not excepting pallida itself, from which form it further differs in lacking much of the cmnamomeous tinge of the upper parts, particularly on the cervix and bend of the wing. Other characters distinguishing leucansiptil.a from actia and amnwphila are the more uniform upper surface and the much more pinkish shade of the cervix, upper tail-coverts and bend of wing; from occid~entalis, the decidedly smaller size; from aduata, the conspicuously less reddish upper surface; from leu~olaema, reduced size and more uniform upper parts.”
I can find nothing published on its habits, which probably are similar to those of the other desert races in neighboring regions.