Song Sparrow

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

 

On any list of North American birds selected for their general familiarity, the song sparrow would have few peers. Although relatively small and not very conspicuously marked, this species combines a readiness to dwell near humans and a persistent and attractive song with a breeding range extending from Mexico to the outer Aleutians and from the islands off the Atlantic coast to those off the coast of the Pacific. Further, the territories of the males are small and the suitable habitats extensive, with the result that the song sparrow is abundant in most of its range. Add to the foregoing the fact that these sparrows are readily trapped, and it is not surprising that some populations have been studied in meticulous detail.

Not only is the song sparrow one of our best-known birds; it is also our most variable, with 31 subspecies recognized as occurring within the territory covered by the A.O.U. Check-List (1957) and 3 additional subspecies in Mexico (Friedmann, et al., 1957). Robert Ridgway (1901) writes, “No other bird of the Nearctic Region has proven so sensitive to influences of physical environment,” and Alden H. Miller (1956) cites the song sparrow as “one of the best examples of substantial racial diversification” among terrestrial vertebrates on this continent. Most of the subspecies occur west of the Rocky Mountains and in Alaska. Thus 9 races are found exclusively in California, to which may be added in California 8 other races that are not confined to that state. As a result of this plasticity, the song sparrow figures prominently in literature dealing with the origin of species and with ecologic gradients. The frontispiece in Joseph Grinnell and A. H. Miller’s (1944) work on California birds will repay examination for its portrayal of variations in eight of the races of that state. Ira N. Gabrielson and F. C. Lincoln (1951) put the extent of the intra-specific variation in the following way: “it is probably true that if all the resident Song Sparrows between Kodiak Island and the Imperial Valley in California were suddenly destroyed, there are few observers who would believe that there was any close relationship between the large dusky Aleutian birds and the small pale form about the Salton Sea.”

It will assist the reader if he is aware of the following decisions as to the manner of presenting this life history of the song sparrow:

(1) Most subspecies are treated separately in order to permit the use of the detailed information that is available for some populations and to maintain the integrity of three contributed accounts, Margaret M. Nice’s summary of her seminal study of euphonia, Richard F. Johnston’s report of his investigation of samuelis, and Robert W. Dickerman’s account of fallax.

(2) When two or more geographically proximate and ecologically similar subspecies are believed not to differ in the essentials of their life histories, they are sometimes grouped and information about them is pooled or is otherwise generalized, as indicated.

(3) When published studies have treated some aspect of the species as a whole rather than of subspecies, e.g., its food habits or its molestation by the cowbird, these results are presented under the first subspecific history, i.e., of the nominate race M. m. melodia, which also includes data that cannot be referred to subspecies and material that appears to be of general applicability.

Thus, the life history of M. m. melodia is to a degree broadly descriptive of the species. Mrs. Nice’s treatment of euphonia, on the other hand, contains a wealth of detail about a small population of a widely distributed migratory race. Dr. Johnston’s life history of samuelis treats in similar detail a rather specialized, sedentary race with a very limited range. For a general view of the song sparrow and its “wonderful adaptability” (Taverner, 1934), therefore, the reader might wish to consult the life histories of the races just mentioned, as well as the accounts of the races grouped as “Alaskan song sparrows” and “Pacific insular song sparrows.” Finally, M. m. rivularis might be referred to as an example of the several subspecies inhabiting the deserts of the United States and Mexico.

EASTERN SONG SPARROW

MELOSPIZA MELODIA MELODIA (Wilson)

Contributed by VAL NOLAN JR.

HABITS

The breeding song sparrow of eastern Canada and of the United States west to the Appalachians displays the typical preference of this species for moist ground and for a low, irregular, dense plant configuration considerably exposed to the sun. “No land bird seems more fond of water,” writes E. H. Forbush (1929). Everywhere it is “primarily a bird of the lower lands * * *” (Knight, 1908), along the banks of streams, the brushy shores of ponds, and in shrubby wet meadows or cattail swamps. Even on the central Atlantic coast, where the race atlantica replaces the present subspecies on the beaches, melodia seems to be the song sparrow found back in the thickets (Stone, 1937), and Dexter (1944) has reported a nest of this race in a salt marsh on a tidal inlet at Gloucester, Mass. But these lowland situations are only a first preference, for the bird is tolerant of a wide range of conditions. It is often found in brushy fence rows and along country roads; and it sometimes breeds even in rocky wooded clearings in Maine, in small wooded openings only a few rods in diameter in New York (Eaton, 1914), and in second-growth woodland in Pennsylvania (Todd, 1940). Gardens and yards offer the song sparrow sunny, bushy, moist cover, and the bird is a common nesting species in suburbs and small towns. On Mount Mitchell, North Carolina, the highest point in the eastern half of the United States, Burleigh (1941) observed individuals of the race euphonia as high as 6,300 feet above sea level during the summer.

Spring: Although a few song sparrows winter far north, most withdraw from that part of the range in Canada and northern New England. M. M. Nice (1933) has cited the evidence, from banding records, that some individuals of this subspecies are resident “in regions where most of their kind are migratory.” Hervey Brackbill (1953) has found that the breeding population of Maryland contains both migratory and sedentary individuals. In the Hudson River valley and around New York City the species is a common winter resident. The arrival of migrants, and the return to their breeding territories of birds that have wintered, normally begins in the latter half of February in the southern part of the range, while the first spring arrivals appear in Maine in mid- and late March and in Canada in March and early April. In Maine migration continues into early May (Palmer, 1949).

Carl H. Helms (1959) weighed song sparrows captured in Massachusetts just before and just after a large night migration in April. Birds weighed before the flight averaged 1.41 grams heavier than those that arrived as the result of the movement. These post-flight individuals were noticeably less fat.

The song sparrow sings even on cold clear mornings of late winter, and its voice is a characteristic sound of early spring. Aretas A. Saunders (1947) reports singing in Connecticut when the temperature was —2° F.

Territory: Territory appears to be established principally or entirely by the male. E. H. Forbush (1929) has described behavior that probably includes elements both of territorial defense and of courtship or pair formation: “There is considerable rivalry among the males, but their contests appear to be mainly competitions in song and flight. They chase the females and each other about through the air with fluttering wings, often sailing and singing. Their pursuit seems not to be in earnest, as, notwithstanding the rapid movement of their wings, their progress is slow. Now and then a bird pauses in his flight to sing, supported for an instant on his widespread pinions. Flight-songs also carry them up into the air. Occasionally a battle ensues between two rival males, and sometimes they even roll and tumble in the dust with locked bills and beating wings.” In the section on courtship is a description of behavior that probably has territorial functions as well. The song sparrow’s persistent songs, six to eight per minute at dawn in spring (Forbush, 1929), are, of course, associated with the maintenance of territory.

The size of the individual territory in favorable habitat is less than an acre. Robert E. Stewart and Chandler S. Robbins (1958) give an interesting series of data on the population density of breeding song sparrows in Maryland. (Population densities provide a basis for estimating only maximum territory size, for it does not follow that all the area censused actually fell within the boundaries claimed by the males.) In 19.2 acres of ” ‘shrubby field with stream-bordered trees’ ” were 21 territorial males; in 9.5 acres of ” ‘open hemlock-spruce bog’ (brush-meadow stage * * *)” were 3 males; and in 20.5 acres of ” ‘moderately sprayed apple orchard with infrequently mowed ground cover * * *’ ” were 4.5 territorial males.

Courtship: Witmer Stone (1937) writes as follows of his observations on Cape May: “In late March and April the air seems simply filled with Song Sparrow song and at this time we see male birds flying from bush to bush with neck stretched out, head and tail held high, and wings vibrating rapidly. This seems to be a part of the courtship display and as soon as the bird alights it bursts into song. On March 21, 1925, and April 2,1914, I have noted this performance and the birds were evidently paired * * *.” This behavior probably was associated also with territory defense. To the account from Forbush quoted in connection with territory may be added the same author’s statement that song sparrows “spend much time in the pleasant pastime of courtship. The females seem to be modest and coy.” The duration of the periods of pair formation and between pair formation and nesting seem not to have been recorded. A comparison of the dates given for the height of the return of spring migrants and for the beginning of general nesting in a given locality suggests that a month or more often elapses in these “prenuptial” and “preliminary” periods (Nice, 1943).

The same males and females have been found mated to each other in successive years (Hamill, 1926; Higgins, 1926).

Nesting: Building is carried on principally by the female, but Ora W. Knight (1908) once saw a male apparently assisting his mate. He was “more inclined to shirk his share, picking up material, dropping it and picking it up again, singing meanwhile.” During building, says Forbush (1929), “the male devotes himself more to song than to labor.”

The duration of building is variously given, with 5 days the lowest figure and 10 days a commonly accepted maximum. Weather is known to influence the speed of building; and it may be supposed that the time-advance of the season, the number of nesting attempts already made, and the presence or absence of an earlier brood might all affect the female’s building behavior. As mentioned below, females of the race melodia commonly raise three broods in a season and sometimes deposit the eggs for a later brood in a previously successful nest. Andrew J. Berger (1951) found five nests built in one season by the same female of the subspecies eupphonia; not all succeeded.

The heights of nests range from ground level to at least 12 feet high. Most nests are placed on the gound, usually concealed under a tuft of grass, a bush, or a brush pile; and elevated structures are rare or absent early in the season. Eaton (1914) reports that 99 percent of the nests in New York are on the ground, but most writers use words suggesting only that something over half are located there. Elevated nests are “at a height of generally not over two or three feet” (Knight, 1908), but numerous references to somewhat higher sites can be found. Locations over water are not uncommon.

Plants in which nests are placed are grasses, sedges, cattails, a great assortment of bushes and shrubs, and, more rarely, trees of many species. Nests are occasionally built in cavities. Hollows in old apple trees are apparently the commonest such locations (Knight, 1908; Todd, 1940; Eaton, 1914); but hollow logs and rails, unoccupied buildings such as woodsheds, and even nest boxes (Palmer, 1949) have been resorted to.

The materials used for the outer, bulky part of the nest, as opposed to the lining, are most commonly leaves, strips of plant bark, and weed and grass stems. The lining is of fine grasses, rootlets, and horse or other animal hair. W. E. C. Todd (1940) states that nests “when above the ground * * * are often quite bulky.” Knight reports the dimensions of one nest placed on the ground: the diameter of the cavity was 2½ inches, while the overall diameter varied between 5 and 9 inches; the cavity depth was 1½ inches, the overall depth 4½ inches.

Eggs: The statements in this paragraph are applicable to the song sparrow without regard to race. The female lays from 3 to 6 eggs. They are slightly glossy and range from ovate to short ovate. The ground color of freshly laid eggs is “pale Niagara green” but this fades upon exposure to a greenish-white. Most eggs are very heavily speckled, spotted, or blotched with reddish browns such as “Verona brown,” “russet,” “cinnamon brown,” or “Brussels brown.” Some eggs have undermarkings of “pale neutral grey.” The spottings generally are more or less evenly distributed over the entire surface, sometimes obscuring the ground color and making it appear to be a light huffy brown. They vary considerably both in shape, size, and intensity. The measurements of 400 eggs average 20.4 by 15.6 millimeters; the eggs showing the four extremes measure 25.9 by 17.8, 24.5 by 18.3, 16.9 by 15.4, and 19.6 by 12.8 millimeters.

Turning to the race melodia, the measurements of 50 eggs average 19.5 by 15.1 millimeters; the eggs showing the four extremes measure 21.9 by 14.5, 20.1 by 16.8, 17.8 by 14.2, and 18.0 by 14.0 millimeters.

Incubation: Forbush (1929) states that incubation is “by both sexes, female chiefly” and that “some males assist the females a little * * *.” However, there is no evidence that males have an incubation patch, without which they would be ill-equipped to supply heat to the eggs. Mrs. Nice (1937) found males of the race euphonia lack this patch, and her unequivocal statement that only the female euphonia incubates casts doubt on Forbush’s contention. The role of the male during incubation is probably confined to the defense of territory and nest.

Incubation seems to start sometime not many hours from the laying of the last egg of the set, if this inference may be drawn from the failure of observers to report differences in development in the young of a brood. In euphonia Mrs. Nice observed that most often a clutch hatched over a 2-day span, indicating that incubation had begun before all eggs were laid.

The incubation period is said by Knight and Forbush to be from 10 to 14 days, but it is doubtful if accurate measurements of the period, as it is presently defined, would be less than 12 days, as in euphonia (Nice, 1937). W. E. Schantz (1937) watched a female euphonia incubate three eggs (laid July 10 —12) for 24 days; the eggs failed to hatch.

Young: Most of our knowledge of the development of the behavior of nestling song sparrows comes from Mrs. Nice’s work, devoted chiefly to euphonia. The following paragraph is based on her report (1943). The development of the plumage is described below under the heading Plumage.

Newly hatched song sparrows can grasp, gape, swallow, defecate, and change location “by means of uncoordinated wrigglings.” A feeding note has been heard in 2-day-old birds. The eyes begin to open at age 3 or 4 days. Incipient preening motions appear at age 5 days, as do, rarely, cowering and the ability to utter a location call. At age 7 days many motor coordinations are acquired, and henceforth the bird “is capable of leaving the nest.” Among the behaviorisms of the 7-day-old are cowering, stretching of the wings, head-scratching, yawning, and climbing to the nest rim. Birds 8 and 9 days old acquire new types of wing-stretching, engage in wing-fluttering and -fanning, and body-shaking, and utter new feeding notes.

Both parents feed the nestlings, chiefly on “insects, worms, beetles, grubs, flies, caterpillars, grasshoppers, and similar insects” (Knight, 1908). The period in the nest varies, its minimal limit being given as 7 days by Forbush (1929) and its maximum as 14 days by most writers. Seven days undoubtedly does not represent a natural, undisturbed nestling period, but is probably the youngest age at which nestlings will leave the nest when disturbed. Knight says that young leave ground nests earlier than they do elevated nests, and that this early age is 10 days. At this time they are still unable to fly, and newly fledged birds remain hidden in plant cover. Mrs. Nice (1937) states that young euphonia “when * * * about 17 days old * * * are able to fly and come out of hiding.”

Dependence on the parents continues until after the post-juvenal molt (Todd, 1940). The parental bond may be assumed to be severed at the age of about 28 to 30 days as in euphonia (Nice, 1937).

As in other species, juvenile song sparrows occasionally engage in some of the behavior of nest building (Hoyt, 1961).

As second and third broods are produced regularly, and fourth broods probably occasionally, as far north as Massachusetts, the matter of timing successive families is of interest. “When the young of the first brood are able to fly, the female immediately begins to deposit eggs for the second brood, often in the same nest, leaving the male to care for the first, and he attends them usually until the young of the second brood have hatched, when he leaves them to help feed and care for the younger brood” (Forbush, 1929).

Plumages: In the following description of the plumages and molts, material involving both melodia and euphonia has been used. The sexes are identical in their molts and practically identical in their plumages. Females average a little later than males in date of molt. Minor sexual differences in plumage will be mentioned.

Natal down, described as both sepia-brown in color (Dwight, 1900) and black (Nice, 1943) is present at hatching. Mrs. Nice (1943) writes that this down “is prominent on the dorsal, femoral and occipital regions and on the coverts. For the first two days there is little change except in increased length of down.” The progress of the molt into the juvenal plumage is described in detail by G. M. Sutton (1935), who writes:

* * * the nestling-stage of the juvenal plumage is * * * notable for its dullness, the feathers of the loral, malar, and superciliary regions being still for the most part in their sheaths, and the tertials so short that their rich edgings are not yet apparent. The streaking of the chest is quite sharp, but on the sides it is, if anything, less marked than in later stages. Male and female birds are apparently not distinguishable at this age. The pectoral streaking is so much intensified because the feathers lie close together and are partially sheathed, that the actual width of the streaks is difficult to determine.

By the time the tail is an inch long the feathers of the face are almost altogether unsheathed, the tertials and secondaries are practically of full length, the pectoral plumage is fully fluffed out, and the bird is, therefore, much more colorful in appearance. At this stage males may, with a fair degree of certainty, be distinguished from females by the heavy streaking of the chest. Chapman * * * tells us that the “breast blotch is wanting” in this plumage. While this is no doubt to a considerable extent true, two individuals in a series of eleven specimens at hand show a definite blotch and two others exhibit a tendency toward convergence of streaks in the middle of the chest.

Sutton considers that song sparrows have a rather definite and complete juvenal plumage. “By the time the juvenal rectrices are of full length the body plumage is comparatively complete with all the feathers unsheathed and with no noticeable intrusion of pin-feathers of some subsequent plumage.” He says that “specimens in juvenal feather may be taken during a long period of the summer,” but is cautious about concluding how long the individual bird wears the plumage.

The foregoing is essentially an account of the molt, not the plumage, of which Dwight (1900) gives a good description. The bird “resembles Z. albicollis, but lacks chestnut above” and is “paler on [thel crown and less streaked below. Above, including sides of head, wood-brown or sepia broadly striped on back, narrowly on crown, nape and rump with dull black, the feathers centrally black with a narrow zone of walnut and wood-brown and grayish edgings. Indistinct median crown and superciliary stripes dull olive-gray with dusky shaft streaks. Rictal and submalar streaks black; orbital ring buff. Wings dull black with walnut edgings, the wing coverts and tertiaries buff tipped. Tail olive-brown broadly edged with walnut and indistinctly barred. Below, dull white washed with pale or yellowish buff deepest on the throat and flanks and streaked on sides of chin, throat, breast and sides with dull black. Feet and bill pinkish flesh, becoming dusky with age, the lower mandible remaining partly flesh-color.” Dwight believes this plumage is worn several months; it fades considerably.

The first winter plumage is acquired, according to Dwight, “by a partial, sometimes complete, postjuvenal moult” beginning in some birds in mid-August, in others not until the last of September. These latter will still show new feather growth late in October or early in November, although “the whole period of moult does not cover much more than two months in the majority of cases.” The molt “involves the body plumage and the tail and very often, part at least, of the remiges. The renewal of five or six outer primaries occurs in nearly all young birds of this species and is very likely characteristic of the first brood. * * * The secondaries are rarely found in moult, the tertiaries, alulae and wing coverts regularly so. * * * [Occasionally~ the renewal of primaries, secondaries and even of rectrices, might easily be overlooked as the new feathers are nearly of the same pattern and color as the old and not in contrast * * *.”

In appearance, the first winter plumage is like the previous one, “but is whiter below and richer in chestnut streakings both above and below. The lateral crown stripes are distinct with black streaks, the median and superciliary stripes distinctly olive-gray. Below, white washed with pale vinaceous cinnamon on sides of head, across jugulum and on sides, and streaked, except on chin and mid-abdomen, with clove-brown bordered with chestnut, the streaks becoming confluent at sides of chin and on mid-throat forming three nearly black spots. Old and young become absolutely indistinguishable in most cases, young birds with the wing edgings perhaps a trifle duller and with a yellowish tinge.” In this plumage females are “apt to be more washed with brown or to have a yellowish cast when compared with males” in the same plumage.

The first nuptial plumage is acquired, according to Dwight, by wear, which is marked; “by the end of the breeding season the birds are in tatters. The buff is lost and the streaking below comes out in strong contrast on a white ground.”

The adult winter plumage is “acquired by a complete postnuptial moult beginning usually about the middle of August and completed before the end of September. Old and young cannot be told apart with any certainty, adults however with wing edgings that may perhaps average darker and browner and the throat markings blacker.”

The adult nuptial plumage is “acquired by wear as in the young birds with the same results.”

The following description of the adult plumage is by Robert Ridgway (1901):

Adults, (sexes alike): — Pileum brown (mummy brown to almost burnt umber), narrowly streaked with black and divided by a narrow median stripe of gray, tbis also narrowly streaked with black; hindneck brownish gray, more or less streaked or washed with brown; scapulars and interscapulars black medially, producing streaks of greater or less width, these margined laterally with brown (like the color of the pileum), the edges of the rectrices, more or less broadly, brownish gray; rump olive-grayish, more or less streaked with brown (Sometimes with blackish also); upper tail-coverts browner than rump and more distinctly streaked; tail brown (broccoli brown to russet brown), the middle pair of rectrices with a narrower median stripe of dusky brown, the inner webs of the other rectrices darker brown than outer webs: lesser wing-coverts brown; middle coverts brown, margined terminally with pale brownish gray, and marked with a more or less distinct median streak or spot of dusky; greater coverts brown, margined terminally with paler and marked with a broad median tear-shaped (mostly concealed) space of blackish; tertials mostly blackish, but outer webs chiefly brown, passing into a paler (sometimes pale grayish or almost grayish white) hue terminally; rest of remiges dusky, edged with paler or more grayish brown; edge of wing white; a broad superciliary stripe of olive-gray, sometimes approaching grayish white on lower portion; loral, suborbital, and auricular regions darker olive-grayish, the latter margined above and below by narrow postocular and rictal stripes of brown, these brown stripes sometimes narrowly streaked with black; a broad malar stripe of dull white or pale buffy, margined below by a conspicuous submalar stripe or triangular spot of black or mixed brown and black; under parts white, the chest marked with wedge-shaped streaks of black, more or less broadly edged with rusty brown, these streaks more or less coalesced in the lower central portion of the chest, or upper breast, forming a more or less conspicuous irregular spot; sides and flanks streaked with black and rusty brown, the ground color, especially on flanks, more or less tinged with pale olive-grayish or huffy; under tail-coverts white or pale buffy, more or less streaked with brown; maxilla dusky brown, paler on tomia; mandible horn color; iris brown; tarsi pale brown, toes darker.

Albinism occurs in song sparrows, and Root (1944) reports a banded individual that acquired a considerable degree of whiteness during a 28-day period in early autumn, presumably as the result of molt.

Food: S. D. Judd (1901) has described the diet of song sparrows without regard to race. For the year, animal matter constitutes 34 percent of the total food, the greatest amount being taken from May to August, when insects represent about half the bird’s food. Ground-, leaf-, and dickbeetles, weevils, and other beetles rank first in number; grasshoppers, locusts, larvae such as the cutworm and army-worm, ants, wasps, ichneumon flies, bugs, leaf-hoppers, larvae and imagos of horse-flies, etc., are also taken. The remaining two-thirds of the diet is composed of seeds of crabgrass and pigeongrass, timothy, old-witch grass, barnyard grass, panic-grasses, orchard and yard grasses; knotweeds, wild sunflower, lamb’s quarters, gromwell, purslane, amaranth, dandelion, chickweed, dock, ragweed, sheep-sorrel and wood-sorrel; a little grain, largely waste; and, before the seeds have ripened, wild berries and fruits such as blackberries, strawberries, blueberries, elderberries, and raspberries, wild cherries and grapes, and woodbine berries. The species is very beneficial as a destroyer of injurious insects and weed seeds. Judd says, “Only 2 per cent of the food consists of useful insects, while 18 per cent is composed of injurious insects; grain, largely waste, amounts to only 4 per cent, while the seeds of various species of weeds constitute 50 per cent.”

W. L. Dawson (1923), writing of song sparrows of no specified race, says that a bird “sometimes seizes and devours small minnows.

Behavior when foraging reflects the seasonal dietary preferences already described. Eaton (1914) states that in summer song sparrows cease to feed largely on the ground and sometimes forage for insects among foliage as high as 20 and 30 feet, although usually among bushes and grass. These birds scratch the ground by kicking simultaneously with both feet. Charles H. Blake writes of watching a song sparrow catch winged termites as they emerged from their subterranean colony in early June.

Behavior: Song sparrows are often furtive in manner, and Knight (1908) gives a good description of the behavior of alarmed birds. He states that they prefer to “work downward into the bushes with bobbing tail, hopping along from twig to twig, or skulking through the underbrush, grass and leaves. They do not fly, save from bush to bush, unless closely pursued with evident intention to flush them or do them harm.” Witmer Stone (1937) speaks of “how well adapted [song sparrows] are for a terrestrial life and how rapidly they can run, mouse-like, through the grass.”

Despite their sometimes secretive behavior, birds dwelling near humans often develop considerable tameness. Forbush (1929) states that they may be conditioned by feeding to come when called and tells of one song sparrow that learned to associate the sound of a bell with the fact that food was to be scattered and of another that learned to peck at a window to be fed.

The behavior of females on the nest is often cryptic, according to Knight (1908). Sometimes the bird sits until almost stepped on; at other times, when the male gives the alarm, she slips off, sneaks a few feet away, and then begins to call. Johnston (1957) in long experience with the shrub-nesting race, samuelis, saw only one case of rodent-like distraction display. Both adults protest intrusions into the vicinity of the nest, and Forbush (1929) describes the posture of nest defense as involving “outspread wings and depressed tails.” This threat, if unsuccessful, may be replaced by attack. Birds as large as the catbird and hairy woodpecker happening to approach the nest are attacked, and Forbush mentions a successful attempt by a song sparrow to drive five house sparrows from a feeding station.

Bathing, states Forbush, occurs “during the day whenever opportunity offers” and, if there is water, “every night after sunset.” Puddles, including salt water along the shore, are used; and the song sparrow is one of many species that bathe in drops of water on grass and leaves by striking the foliage with the wings and body and thus throwing water on the plumage.

Scratching of the head by song sparrows is “indirect,” with the foot brought over the wing to reach the head, and Hailman (1959) has observed song sparrows and other emberizines scratching the head against perches.

Anting has often been noted in song sparrows; Whitaker (1957) has summarized the details.

“Helping,” i.e., the feeding of young both of other song sparrows and of other species, has been noted several times and summarized by Brackbill (1952) and Skutch (1961). Perhaps the most surprising instance, reported by Brackbill, involved the cooperative building and joint use of a nest by a pair of cardinals and a pair of song sparrows. Both females incubated, the cardinal sometimes sitting on the sparrow. The cardinal eggs succeeded, and all four adults fed the nestlings. Forbush (1929) describes an instance in which two females laid a total of eight eggs in one nest; one of these birds had its own nest 30 feet away but did not use it. The females took turns incubating, and all eggs were said to have produced fledglings.

The flight speed of song sparrows has been measured by 0. P. Pearson (1961) as 15.9 miles per hour and possibly as much as 21 miles per hour.

Manwell and Herman (1935) found that individual song sparrows displaced and released in spring as much as 1 ~ miles returned quickly to the point of capture; at an intermediate trapping station on the presumed line of flight none were caught.

Body temperatures of 64 individuals of the race euphonia averaged about 109.6° F., varying 10° F. within the sample (Becker and Stack, 1944). There is a considerable literature on song sparrow weights; Mrs. K. B. Wetherbee (1934) gives many data. Mrs. Nice (1935) lists an average weight of 21.3 grams for 267 adults. LeRoy C. Stegeman (1955) reports weight fluctuation amounts at times to 20 percent in 24 hours, with peak weights in the spring recorded in the late morning and late afternoon.

Voice: Male song sparrows sing their variable repertoire not only during the breeding season but at other times. Songs may be heard in much of the breeding range during any month of the year (Saunders, 1947), and dawn singing on clear, cold mornings in January and February is especially noticeable. Regular singing in spring begins in Connecticut in late February or in March (Saunders, 1947) and generally closes, for a time, in the third week of August (Saunders, 1948a). There follows a revival of song, beginning in Connecticut on the average date of September 30 and continuing until November 21 (average). Aretas A. Saunders, who is responsible for these dates, writes (1948h): “This species is the most regular and dependable fall singer of all our birds.”

Forbush (1929) states that the birds sing “no matter how very stormy the weather” and sometimes even “in the darkness of night.”

Six to eight songs per minute is the frequency during the dawn hours of the breeding period (Forbush), with singing continuing, but less frequently, all day. Knight (1908) reports that in summer in Maine most singing occurs during the dawn and evening hours. Forbush says that occasionally molting birds sing a whisper song.

When singing, the male mounts to a position typically between 7 and 15 feet from the ground (Eaton, 1914); trees, shrubs, fences and boulders are among the song posts used. Singing has often been observed in birds on the wing, and Forbush’s description, quoted under Territory, indicates that this form of behavior may have become an element in some displays.

Female song sparrows have been known to sing, and Mrs. K. B. Wetherbee (1935) has written of a female in Massachusetts that sang “a clear series of whistled notes” from April to mid-June.

Not only the vociferousness of the species but also the unusual variability of the repertoires of the individual males are responsible for an abundant descriptive literature. Further, the use of electronic recording and analysis of songs has thrown light on the extent to which song sparrow vocalizations are modifiable as opposed to innate. For the present account, two authorities on bird song are quoted. Aretas A. Saunders wrote Mr. Bent: “I have 885 records of the song, no two of them alike. If we count trills as single notes, the number of notes per song varies from 4 to 20, averaging about 11. The length of songs varies from 1.8 to 5.2 seconds, the average being 2.7. The pitch varies from D” to F””. The pitch interval varies from 1 to 7½ tones, the average about 3½ tones. Each individual song sparrow sings a number of different songs. It commonly sings the same song over a half a dozen times or so, and then takes up a different song. The number of songs per individual varies from 6 to 24, the latter being an unusual bird.”

The same author writing elsewhere (1951b) goes into additional detail and reports the following: Pitch varies from 1150 to 5450 vibrations, in notes audible to man, and pitch intervals are similar to those in human music. There is little variation in intensity. “Quality is usually sweet and musical * * * Consonant sounds are not very noticeable. * * * The song has three parts: strongly rhythmic introductory notes, a central trill, and a final series of rather irregular and indefinite notes. * * * Songs are of five types * * * [differing] primarily in the position and relative pitches of the introductory notes and the trill.”

Donald J. Borror (1961), who analyzed 889 tape recording of songs from 113 different individual song sparrows of the races melodia and euphonia, writes,

The songs of this sparrow consist of a series of different phrases (mostly 1- to 4-noted), and usually a trill; many of the notes are buzzy. * * * A given bird has a vocabulary of a large number of notes and phrases, and these are variously combined to produce up to a dozen or more different song patterns; the different patterns of a given bird are often quite different. The songs of a given pattern may vary * * *.

Song Sparrow songs are of two general types, those beginning with two to four (rarely one or five) similar and equally spaced phrases, and those beginning with four to twenty similar phrases that increase in tempo * * * “. Songs of the first type were much more common, making up 83.8 percent of the Ohio [euphonic] patterns and 86.7 per cent of the Maine [melodic] patterns * * *.

A Song Sparrow apparently has an inborn tendency to sing songs of two general types, but it learns its phrases by listening to other, nearby Song Sparrows. As a result, the songs of different birds in a local population contain similar notes and phrases (but usually arranged differently), while the songs of birds in separated populations contain different phrases. The farther away two populations are, the less likely they are to use similar phrases in their songs.

In a later, very detailed analysis of variation in the songs of Maine song sparrows, Borror (1965) found 544 song patterns represented in 7,212 tape-recorded songs of 120 birds.

A description of autumnal song by Forbush (1929) probably is applicable not to adults but only to birds of the year: Most of the singing un fall] is quite different, ranging from a low connected warble to a song resembling that of the Purple Finch, and (rarely) one like that of the Vesper Sparrow. There is a particularly low, sweet melancholy warble uttered just before the bird departs for the south.” Formless, continuous warbling of the kind described is commonly a stage in the ontogeny of song in passerines (Lanyon, 1960).

Call notes are described as “tchenk,” “tchip,” “tchunk,” “chip,” “tcheek,” “chuck.” Forbush also mentions a note “sst”; a similar note given by California races is regarded by Dawson (1923) as functioning as a flocking or recognition call.

Field Marks: A medium-sized sparrow, the song sparrow is best recognized by the heavily streaked breast, on which the streaks are “confluent into a large central spot” (Peterson, 1947). In flight, which is usually for short distances between perches or into cover, the bird is distinguished by its manner of pumping its rather long, rounded tail up and down. Witmer Stone (1937) describes this flight graphically as “‘brokenbacked’ * * * as if the tail were hinged at the base.”

Banding: The longevity record for banded song sparrows appears to be about 8 or 9 years. Mrs. Nice (infra) reports a male that was at least 7½ and possibly 9½ years old at death. A song sparrow Mrs. K. C. Harding (1943) banded April 27, 1936 at Cohasset, Mass., and recaptured there on April 5,1943, was in at least its 8th year.

Recaptures and recoveries at points other than the original banding station have been reported with some frequency in the journal, Bird-Banding. Some of the most interesting of these follow: The number of such “recoveries” and “foreign retraps” from a total of 3,614 song sparrows banded in Montgomery County, Pennsylvania; was 12 (Middleton, 1956); from 6,109 banded in Groton, Mass., 7 (Wharton, 1953); from over 1,200 banded on Cape Cod, Massachusetts, 1 (Broun, 1933). Some of the Pennsylvania birds were caught in Georgia, North Carolina, Maryland, and New Jersey. The Groton, Mass., birds were caught in Arkansas, South Carolina, North Carolina, Nova Scotia, and New Brunswick. The Cape Cod bird was caught in South Carolina. May Thacher Cooke (1943) reports a bird banded in Massachusetts and captured in Newfoundland. Wendell P. Smith (1942) caught a bird in Vermont in April, and it was recaptured 90 miles southward in New Hampshire in June of the same year, an interesting case of reversal in the direction of migratory movement.

Enemies: Perhaps the most interesting and surely the most well documented hazard in the song sparrow’s environment is exposure to the parasitic brown-headed cowbird and bronze cowbird. Herbert Friedmann’s latest report (1963) on the cowbirds states that the song sparrow (all races) shares with the yellow warbler the claim to being the most frequently reported host of M. ater. Friedmann’s summary of the relations between all races of the song sparrow and the brownheaded cowbird is quoted substantially in full:

The song sparrow is one of the most frequent, if not the most frequent, victim of the brown-headed cowbird. Since the former is sympatric with the latter throughout the entire breeding range of the parasite, it is parasitized probably more often and over a greater area than any other bird. The total number of records is very great. After accumulating over 900, I stopped noting them except for records of special interest. The data came from every province of Canada and every state of the United States included in the breeding ranges of both birds. All three races of the parasite are involved, and no less than 17 races of the song sparrow: melodie, atlantica, euphonia, juddi, montana, inexpectata, merrilli, fisherella, morphna, cleonensis, gouldii, samuelis, pusillula, heermanni, cooperi, fallax, and saltonis. So far, none of the purely Mexican races have been reported as fosterers of the cowbird, but this fact is probably due more to a lack of human observation than to any actual immunity of the bird to cowbird parasitism.

There is no need to detail actual instances for the various races of the song sparrow since such cases already have been given in my earlier summaries [Friedmann, 1929, 1934, 1938, 1943, 1949]. However, a few additional records of infrequently reported races of the host species should be mentioned * * * [viz. cleonensis, fallax, morphna, saltonis, samuelis, and inexpertata].

In recent years, not only many hundreds of additional cases, but also much more quantitative data on the host-parasite relations have become available. Hicks (1934) found that 135 out of 398 nests (34 percent) of this sparrow were parasitized in Ohio. Nice (1937a***, 1937b***), also in Ohio, reported that 98 out of 223 nests (43.9 percent) contained eggs or young of the cowbird (the annual percentage varied from 24.6 to 77.7 percent). Sixty-six unparasitized nests raised an average of 3.4 song sparrows, whereas 28 successful but parasitized broods averaged only 2.4 song sparrows, indicating that each cowbird was reared at the expense of one song sparrow. In one instance Nice *** found that a pair of song sparrows raised a young cowbird together with five of their own young. Apparently here no loss of sparrows was involved. In another paper, Nice (1936) noted that, in all the song sparrow nests which she had watched during a period of five years, adult cowbirds removed 5.7 percent of the song sparrow eggs and nestling cowbirds crushed or starved 3.5 percent of the young sparrows. The cowbird eggs did not succeed as well as those of the host; only 30.7 percent of the former, but 35.8 percent of the latter, reached the fledgling stage. In 1930—31, there was one female cowbird to about 11.5 pairs of suitable hosts, but in 1934—35 there was one to 8.6 pairs of suitable victims.

Of all song sparrow nests parasitized, Nice reported that 70 percent held a single cowbird egg each, 27 percent held 2 each, and 3 percent held 3 each. In the area of study—near Columbus, Ohio—the song sparrow was the most important host of the paeasite [sic]. Norris (1947***) noted that 11 out of 27 nests (40.7 percent) in Pennsylvania were parasitized, and Berger (1951***) recorded 37 out of 59 nests found in Michigan (62.7 percent). In the Detroit area, as reported by the Detroit Audubon Society (1956***), the average frequency of parasitism of the song sparrow was 40.1 percent of all the nests found * * *.

One is drawn toward attempting an over-all estimate of the frequency with which the song sparrow is victimized, but to do so with any feeling of accuracy is difficult because the incidence of parasitism appears to vary geographically (or, at least, the frequency with which it is reported varies). From this it follows that the over-all percentage depends on how many geographically different areal data are used in the estimation. [With one group of studies from the eastern United States] * * *, we come up with a total of 323 parasitized nests out of 804 nests observed, or a little over 40 percent. On the other hand, in southern Quebec (Terrlll, 1961, p. 11), out of 486 nests observed, only 62, or 12.7 percent, were parasitized. If we put all these studies together, we get a total of 382 out of 1,285 nests victimized, or 29 percent. This figure becomes yet smaller when we attempt to include data from other parts of the continent.

[One color-banded song sparrow in a single summer] *** had no fewer than five consecutive nests * * “‘. It would seem that, if none of these nests had been interfered with, there would not have been sufficient time for four or five in one season. * * * It appears that one of the effects of parasitism may be to increase the “nesting potential” of the host. * * *

As many as 7 cowbird eggs have been found in a single nest of this sparrow; there are numerous records of 3, 4, and 5 parasitic eggs to a nest. Occasionally, but not often, song sparrows may partly bury cowbird eggs by building a new nest lining over them—if the alien egg is laid before any eggs of the host.

Salmon (1933, p. 100) has reported seeing a song sparrow feeding three fledgling cowbirds; no young sparrows were mentioned. Lees (1939, p. 121) recorded that near Wetaskiwin, Alberta, he watched a song sparrow feeding no less than five young cowbirds. This must be a record of fledgling success for any host species.

Friedmann (1963) lists no instances of parasitization by the bronze cowbird of subspecies of the song sparrow covered by these life histories. Only the Mexican race mexicana has been involved.

Other enemies of the song sparrow are at least four species of hawks (Munro, 1940; Randall, 1940; Hamerstrom and Hamerstrom, 1951; Heintzelman, 1964) and at least five species of owls (Allen, 1924; Hawbecker, 1945; Johnston, 1956; Fisler, 1960; Graber, 1962). 5. A. Altmann (1956) has reported mobbing of mounted specimens of both screech and great horned owls, and F. Hamerstrom (1957) witnessed mobbing of a tame red-tailed hawk. Forbush (1929) mentions nest defense against snakes and turtles; he does not indicate what turtles may be involved, but box turtles of the middle west (Terrapene o. ornata), at least, have been known to eat birds and their eggs (Legler, 1960).

A curious case in which a garter snake (Thamnophis s. sirtalis) disgorged an adult song sparrow is reported by Carpenter (1951), who suggests that the bird must have been found dead and then eaten. Mahan (1956) discovered a milk snake (Lampropeltis triangulum) eating song sparrow eggs in a nest 15 inches above the ground.

A number of external parasites taken from song sparrows east of the Mississippi River, most of them within the range of rnelodia, have been reported by Harold S. Peters (1936). These include the Mallophaga Degeeniella tndgata (Kell.), Mac haerilaemu.s moA slum (Kell. and Chap.), Menacanthus incerta (Kell.), Philopterus subflavescens (Geof.), Ricimus melospir.ae (McGregor); the bloodsucking hippoboscid flies Ornithoica confluenta Say, and Ornithomyia anc hineuria Speiser (syn. 0. fringihin4) ; the mites Analgopsis sp., Lipony8sue sylviarum (C. and F.), Trombicula bisignata Ewing, Trombicula caeicola Ewing; and the ticks Haemaphysalis leporispa2ustnis Packard, Irodes brur&neus Koch, and Modes sp. Herman (1937) gives further data on the hippoboscids parasitizing song sparrows, as does Boyd (1951); the latter would apparently refer the records of Ornithoica confluerda to Ornithoica vicina, as the parasite of song sparrows.

Nestling song sparrows are among the many species victimized by the maggots of blow flies (Calliphoridae). Johnson (1932) found larval Protocalliphora splendida (Macq.) in a nest, and George and Mitchell (1948) report Apaulina metallica (Townsend) (syn. Protocalliphora metollica).

Blood protozoa found in song sparrows (Herman, 1944) include a number of species of the genera Haemoproteus, Leucocytozoor&, PlOAmodium, Toxoplasma, and Trypanosoma.

Cats, other predatory mammals, and man are often responsible for song sparrow deaths; and the physical environment takes its toll in starvation (Forbush, 1929) and in the flooding of ground nests.

Song sparrows are among the birds whose feet occasionally exhibit large, rough, wart-like swellings. H. and J. R. Michener (1936) have described the appearance and effect of this disease, which they also imply may affect the wings and heads of song sparrows, and which they regard as mildly contagious and epidemic. Their observations were made in California. The disease runs its course between from 1 to 5 months. Apparently it often produces no noticeable after effects, but it may cause the loss of the nails and sometimes the phalanges. These authors quote a pathologist’s histological analysis of a foot of an unspecified species. The opinion was that the lesions were not true tumors but were the result of an unrecognized irritant or infection. Viruses are now known to produce comparable effects in some birds (Herman, 1955).

Fall: In Massachusetts, a few song sparrows begin moving away from their breeding places in mid-July. The fact that nests have been found in New York as late as August 25 (Eaton, 1914) suggests that it may be the young of the year that move at so early a date. There is some conflict in the evidence as to when the fall migration is at its peak. The data of Stewart and Robbins (1958) are perhaps the most recently reported and based upon the most voluminous and varied factual data. These authors state that in Maryland and the District of Columbia the normal period of fall migration is between Sept. 20: 30 and Nov. 20-30, with its peak between October 10 and 30. More northerly latitudes would, of course, be correspondingly earlier, e.g., in Maine “throughout September and most of October” (Palmer, 1949).

Winter: Most song sparrows that pass the winter in Massachusetts (Forbush, 1929) remain near the sea, where there are usually patches of ground clear of snow. In Ontario (Snyder, 1951) and New York (Eaton, 1914) the habitat at this season is principally marshes and swamps. In Pennsylvania, Todd (1940) says brushy thickets and fields with corn shocks are frequented by song sparrows. The birds are not especially social, but they are often seen in loose flocks of mixed composition and, particularly in severe weather, may assemble m small companies with other song sparrows.

South of. the breeding range, in the deep south and southeast of the United States, melodia is found with euphonia, juddi, and, in places, atlantica. Here the birds seek the same brushy, moist, riparian and marshy situations that they prefer for breeding. Sprunt and Chamberlain (1949) describe such habitat in South Carolina and say the song sparrow “often * * * is found with swamp sparrows * * *. The thoroughly characteristic song is delivered throughout the winter except in very cold weather or on freezing days.” In contradiction Arthur H. Howell (1932) in his work on Florida says, “The Song Sparrow, so well known in the North by its cheery song, is practically silent during its stay in the South, except for its metallic, characteristic tchip.” Howell adds that the birds associate “in small loose companies, but not in compact flocks.” George H. Lowery, Jr., (1960) of Louisiana, emphasizes the “entirely different personality” of the song sparrow that winters in the south and describes it as a shy and, it seems, silent skulker that prefers the depths of thickets or “a rank growth of broom sedge.”

DISTRIBUTION

Range: Southeastern Ontario, central Quebec and southwestern Newfoundland south to eastern Texas, the Gulf coast, and southern Florida.

Breeding range: The eastern song sparrow breeds from southeastern Ontario (Muskoka District, intergrades with M. m. euphonia) , central Quebec (Lake St. John, Romaine, Blanc Sablon), and southwestern Newfoundland (Parson’s Pond) south through eastern New York (intergrades with M. m. euphonia in central section) and Pennsylvania to extreme northeastern West Virginia (Halltown) and central Virginia (Lynchburg, Petersburg).

Winter range: Winters from southern Ontario (Barrie, Arnprior), southern Quebec (Montreal), central New Brunswick (Memramcook), Prince Edward Island (North River), and Nova Scotia (Pictou) south to eastern Texas, eastern and southern Louisiana (Kisatchie, New Orleans), southern Mississippi (Saucier), southern Alabama (Petit Bois Island), and western and southern Florida (Pensacola, Flamingo); casually north to Newfoundland (Mobile)

Casual records: Casual in Bermuda.

Migration: The data deal with the species as a whole. Early dates of spring arrival are: Virginia: Lynchburg, March 2. West Virginia: Wellsburg, March 21. Maryland: Laurel, February 19. New York: Monroe County, February 21; Suffolk County, February 22. Massachusetts: Martha’s Vineyard, March 14; Essex County, average of 5 years, March 10. Vermont: Bennington, March 11. New Hampshire: Concord, February 22; New Hampton, March 2 (median of 21 years, March 21). Maine: Lake Umbagog, March 25. Quebec: Montreal area, March 27. New Brunswick: St. John, March 16; St. Andrews, March 17. Nova Scotia: Yarmouth, March 15. Prince Edward Island: North River, March 19. Newfoundland: Codroy River, May 2. Illinois: Chicago, March 5 (average of 16 years, March 15). Indiana: Elkhart, March 7. Ohio: Buckeye Lake, February 27. Iowa: ~Sioux City, March 14 (median of 38 years, March 25). Minnesota: Minneapolis-St. Paul, March 14 (average of 15 years, March 22); Lincoln County, March 14 (average of 27 years for southern Minnesota, March 22). Nebraska-Holstein, March 5; Red Cloud, March 11 (average of 21 years, April 3). South Dakota: Sioux City, March 20 (average of 7 years, March 27). North Dakota: Cass County, March 25 (average, April 2); Jamestown, March 31. Manitoba: Treesbank, April 4 (average of 25 years, April 14). Saskatchewan: Sovereign, April 5. Arizona: Baboquivari Mountains, February 16. Utah : Kanab area, February 21. Montana: Columbia Falls, March 19. Washington: Destruction Island, March 10. British Columbia: Arrow Lake, March 7. Alaska: Kupreanof, April 18.

Late dates of spring departure are: Florida: Enterprise, April 17. Alabama: Jasper, May 10; Wheeler National Wildlife Refuge, May 4. Georgia: Savannah, May 10; Atlanta, May 9. South Carolina: Charleston, May 5 (median of 5 years, May 3). North Carolina: Raleigh, April 29 (average of 14 years, April 4). Virginia: Lawrenceville, April 12. Maryland: Laurel, April 29. Louisiana: Baton Rouge, April 8. Mississippi: Rosedale, May 26; Gulfport, April 4. Arkansas: Washington County, May 4; Arkansas County, April 28. Tennessee: Athens, April 28 (average of 6 years, April 24). Kentucky: Bowling Green, April 29. Missouri: St. Louis, April 19. Illinois: Chicago, May 19 (average of 16 years, May 6). Ohio: Buckeye Lake, median April 5. Texas: Tyler, April 28; Sinton. April 8 (median of 5 years, March 29). Oklahoma: Payne County, May 4; Cleveland County, April 24. Kansas: northeastern Kansas, May 9 (median of 18 years, April 26). Nebraska: Holstein, April 25. New Mexico: Mesilla Park, March 20. Arizona: Tucson, May 2. Utah: Washington County, May 1. California: Death Valley, April 6. Oregon: Netarts, April 14. Washington: Cathlamet, May 22.

Early dates of fall arrival are: California: Fortuna, September 19. Arizona: Tucson, September 18. Nebraska: Red Cloud, September 7. Kansas: northeastern Kansas, September 2 (median of 20 years, October 7). Oklahoma: Payne County, September 27; Cleveland County, October 13. Texas: Sinton, September 17 (median of 5 years, October 1); Midland, September 24. Iowa: Sioux City, September 4. Ohio: Buckeye Lake, median, October 1. Missouri: St. Louis, September 30. Illinois: Chicago, August 4 (average of 16 years, August 16). Kentucky: Bowling Green, October 4. Tennessee: Nashville, October 4 (median of 11 years, October 11); Athens, October 7. Arkansas: Washington County, October 14; Arkansas County, October 15. Mississippi: Rosedale, October 2 (mean of 42 years, October 6); Gulfport, October 24. Louisiana-Baton Rouge, October 23. New York: Tiana Beach, October 10. New Jersey: Island Beach, September 26. Maryland: Laurel, September 20 (median of 5 years, September 30); Ocean City. October 4. West Virginia: French Creek, October 10. Virginia: Lawrenceville, October 14. North Carolina: Raleigh, October 2 (average of 18 years, October 13). South Carolina: Charleston, September 21 (median of 7 years, October 10). Georgia: Athens, October 2; Savannah, October 12. Alabama: Birmingham, September 22; Marion, September 29. Florida: Tallahassee, October 8; St. Marks, October 9.

Late dates of fall departure are: Alaska: Tenakee Inlet, September 29. British Columbia: Arrow Lake, November 29. Montana: Fortine, November 5. Arizona: Oak Springs, October 12. Saskatchewan: Eastend, October 22. Manitoba: Treesbank, October 28 (average of 22 years, October 15). North Dakota: Case County, November 1 (average, October 24); Jamestown, October 23. South Dakota: Lennox, November 4; Sioux Fails, November 1 (average of 6 years, October 12). Nebraska: Holstein, November 28. Minnesota: Minneapolis, November 28 (average of 14 years for southern Minnesota, November 7). Ohio: Buckeye Lake, median, November 7. Indiana: Roanoke, November 19. Illinois: Chicago, November 17 (average of 16 years, November 3). Newfoundland: Codroy River, October 10. Prince Edward Island: North River, November 25. New Brunswick: Scotch Lake, November 4. Quebec: Montreal area, November 22. Maine: Lake Umbagog, October 27; Bangor, October 26. New Hampshire: New Hampton, median of 21 years, November 12. Vermont: Rutland, November 21. New York: Suffolk County, November 5; Ontario County, November 1. Maryland: Laurel, November 19. Florida: Leon County, November 27.

Egg dates: The data deal with the species as a whole:

Alaska: 21 records, April 29 to July 3; 11 records, June 1 to June 26.

Alberta: 51 records, May 18 to July 28; 30 records, May18 to June 6; 13 records, June 17 to July 26.

Arizona: 33 records, April 9 to August 22; 14 records, May 16 to May 29; 10 records, June 2 to June 29.

British Columbia: 15 records, April 7 to June 26.

California: 352 records, February 12 to July 7; 73 records, April 12 to April 30; 163 records, May 12 to May 31; 62 records, June 1 to June 20.

Illinois: 116 records, April 22 to August 7; 58 records, May 10 to May 31; 30 records, June 8 to July 18.

Maryland: 210 records, April 12 to August 21; 105 records, May 7 to June 19.

Massachusetts: 156 records, May 5 to August 17; 68 records, May 16 to May 31; 42 records, June 6 to June 20; 30 records, June 26 to July 10.

Michigan: 205 records, April 21 to August 29; 100 or more records, May 7 to June 14.

New Brunswick: 36 records, May 10 to July 16.

New York: 147 records, May 2 to August 24; 65 records, May 10 to June 21; 26 records, June 4 to June 27.

Nova Scotia: 60 records, April 20 to August 6; 30 records, May 24 to June 25.

Ontario: 140 records, April 28 to August 3; 72 records, May 21 to June 20.

Oregon: 80 records, April 7 to July 15; 36 records, May 1 to May 22; 32 records, May 25 to June 29.

Washington: 82 records, April 5 to July 14; 32 records, April 5 to April 19; 30 records, May 2 to May 29.

ATLANTIC SONG SPARROW

MELOSPIZA MELODlA ATLANTICA Todd

Contributed by VAL NOLAN JR.

HABITS

This race breeds on the ocean beaches and barrier islands of the central Atlantic states and shifts its range only a little southward in winter. Witmer Stone (1937) writes that on Cape May, N.J., the bird inhabits “possibly the inner edges of the salt marshes” as well as the coast islands, and other writers have found atlantica in or near salt marshes. Stone proceeds to give the following interesting ecological information about this race and melodia: “To illustrate how environment affects the distribution of these birds it may be mentioned that a series of breeding Song Sparrows collected in 1891 on the edge of the old Cape Island Sound and on the salt meadows that formerly existed southwest of Cape May are all typical of the Atlantic Song Sparrow * * ‘~ Since * * * the meadows [were filled and] replaced by dry ground with thickets of bayberry, etc., the common Eastern Song Sparrows of the interior have spread out and occupied the area.”

“[M. atlantica is] rarely, if ever, found far from salt water. It nests in myrtle thickets and in willows at the edge of the salt marsh, obtaining much of its food in the marsh itself, somewhat in the manner of the Seaside Sparrow” (Burleigh, 1958). Alexander Sprunt, Jr. wrote Mr. Bent that “the nest of atlantica resembles that of melodia in construction, being made of grasses at low elevations, but always in or near tide water. The eggs are practically indistinguishable from those of melodia.” The measurements of 40 eggs average 19.6 by 15.4 millimeters; the eggs showing the four extremes measure 21.6 by 15.5, 20.1 by 16.8, and 17.8 by 14.2 millimeters.

The voice is similar to that of melodia, according to Sprunt, but Murray (1941) found the song distinguishable “by a buzzing quality, definitely reminiscent of the song of the Bewick’s Wren.”

The winter habitat is much the same as that of the summer, although Sprunt’s note says that in South Carolina the bird “frequents the same habitat as melodia, showing a predilection, however, for salt and brackish marshes and environs.”

Writers disagree as to whether atlantica and melodia are distinguishable in the field. W. E. C. Todd (1924) described the subspecies atlantica as “much grayer above, with the blackish streaking more distinct, and the reddish brown feather-edging reduced to a minimum. More nearly resembling Melospiza melodia juddi, but more grayish above * * *

DISTRIBUTION

Range: Tidelands from Long Island, New York, to Georgia.

Breeding range: The Atlantic song sparrow breeds in the tidelands along the Atlantic coast from Long Island, New York (Shelter Island) south to North Carolina (vicinity of Beaufort), including lower Chesapeake Bay and the lower Potomac River in Maryland (Morgantown) and Virginia.

Winter range: Winters on breeding grounds north at least to Maryland, ranging south along Atlantic coast to South Carolina (Mount Pleasant, Yemassee) and Georgia (Savannah).

MISSISSIPPI SONG SPARROW

MELOSPIZA MELODIA EUPHONIA Wetmore

Contributed by MARGARET MORSE NICE

HABITS

From 1928 to 1936 at Columbus, Ohio, I studied a population of song sparrows on a 40 acre tract known as “Interpont.” The habitat consisted of weeds, shrubs and trees; the area adjoined the Olentangy River. Over 500 adults were color-banded and 353 nestlings given aluminum bands (Nice, 1937). Later a number of nestlings were hand-raised and kept in captivity; several belonged to the eastern subspecies melodia, having been hatched in Massachusetts; the others were euphonia from southern Michigan (Nice, 1942).

Migratory status: About half the resident males proved to be permanent residents on Interpont, and about 20 percent of the females. Six males and one female changed status during the study. Banding of two and three successive generations gave no evidence of a migratory or resident strain. The migratory impulse seemed to be latent in all the birds, cold weather in October stimulating it in the majority of the individuals, mild weather in October inhibiting it in some.

Spring: The spring migration normally showed two main flights: an early migration of breeding males in late February or early March, and the main flight of breeding males and females, and also transients, in the middle of March. The early migration was absolutely dependent on a warm wave the last of February or the first of March, but the main migration was only relatively dependent on a rise in temperature. Severe cold waves stopped migration short. The early males migrated at markedly higher temperatures, an average of 50°  F., than did the later males, which migrated at an average of 43°  F. High temperatures in December, January, and early February never brought a flight, so it is clear that migration was dependent on both increasing day-length and rising temperature. Fifty-seven migration dates for 22 banded males were obtained; 5 birds came consistently early and 6 consistently late, while the others varied in different years depending both on the weather and on their ages, older birds coming earlier than younger ones.

Territory: The holding of territory is a fundamental trait with these song sparrows, enforced by innate behavior patterns consisting of song, display and fighting. This territoriality is essential for the undistrubed carrying out of the reproductive cycle (Nice, 1939). Although highly territorial for over half the year, and inclined, if a resident, to remain on or near his territory permanently, yet in fall and winter the male becomes somewhat social, particularly in times of severe cold and snow. Occasionally females hold territory for themselves; sometimes one helps defend the male’s boundaries; sometimes females ignore the boundaries established by males. Male and female song sparrows were recorded as driving from their territories their own and 21 other species, ranging in weight from 6 to 42 or even 50 grams. (The song sparrow’s own weight averages 22 grams.) Yet several of these species nested among the song sparrows.

Some male song sparrows keep the same territories year after year, while others make slight changes. Females returned to their former nesting territories in 20 of 54 cases, settled next door almost as often, and in 19 instances settled at distances of from 100 to 800 yards. Twenty-two males banded in the nest took up territories from 100 yards to nearly a mile from their birth places, while 12 females banded in the nest settled from 50 yards to nearly a mile from their birthplaces. The minimum size of a territory was some 2000 square meters (½ acre), the average size in a region well filled with song sparrows about 2700 square meters (2/3 acre), and the maximum size about 6000 square meters (1½ acres). In years of low population density, territory size was somewhat larger.

Pair Formation: In pair formation, the territorial male gives the same initial reaction to all intruders of his species, that is, he flies at them. Migrating song sparrows respond by leaving; a male in breeding condition puffs out his feathers, sings, and waves a wing, while a female seeking a mate stands her ground and gives special notes and postures. The male “pounces” upon his mate, sometimes colliding with her, sometimes only swooping over her, then flies away with a loud song; she stands still and either gives her copulation note or a threat note. Copulation, however, never occurs in connection with pouncing. The male stops singing as soon as joined by a mate.

The pair normally stays together throughout one nesting season. Yet a female may sometimes follow her young into a neighbor’s territory and if he is unmated she may remain with him for the next brood. Remating of pairs a second year has been known in only 8 out of 30 possible cases, probably due to the many chances a male has of getting a mate before the return of his last year’s mate. A female finding her former mate already mated to another female does not drive off the new female, as does the brown thrasher (Thomas, 1952), but joins another male. There were 4 cases of bigamy, apparently arising when an incubating female lost her mate and attached herself to a neighboring male.

Nesting: Typically the nest is built entirely by the female. An exceptional male, while unmated, built 2 incomplete and one complete nest; later he helped his mates built 3 nests (Schantz, 1937). The nest is a rather simple affair built largely of dead grass and weeds, with a few fine roots and pieces of grape-vine bark, and lined with fine grass and occasionally horse hair. Renesting regularly occurs if a nest fails, until the nesting season is at its end. The song sparrow shows its adaptability in the nest sites chosen. The requisites are secure support and concealment. In April almost the only situations on Interpont offering these characteristics are on the ground; here under tufts of grass, weed stalks or thistles, and often in a natural depression, nine-tenths of the nests of the first attempt have been situated. One-third of the nests of the second attempt were placed above the ground, as were two-thirds of the third attempt. Few nests were built more than 3 feet above the ground. Replacement nests were built at 10 to 55 yards from the nests replaced, averaging 25 yards.

Reginald F. James wrote Mr. Bent from Willowdale, Ontario: “The Song Sparrow begins to nest around April 20th, and its first nest is usually placed on the ground. The second nest is built about 18 inches above the ground level. If and when a third nest is constructed, it may be placed as much as 3 feet above ground level, usually in a thorn bush.”

One nest, however, he found 9½  feet above ground in a horizontal hole in a willow. One banded song sparrow nested for 3 years in his garden: “On May 18, 1947 it was found sitting on three lightly marked eggs in a perfect replica of an ovenbird’s nest. On June 14th she was found sitting on three heavily blotched eggs and one cowbird egg, eighteen inches from the ground in a wild rose bush.”

A different picture is given in a letter to Mr. Bent by D. J. Nicholson of small colonies of song sparrows observed from June 10 through August at Lake Summit, near Tuxedo, Henderson County, North Carolina, at an elevation of 2,000 feet. Nests were often placed 2 to 4 feet up in small pine saplings. Three nests were found “in very large tall pines 25 or 28 feet above the ground,” and two others at 20 feet; these were built 8 to 15 feet from the trunk of the tree. Around Tuxedo many nests were found “well up in apple trees 6 to 12 feet above ground.”

The nest is usually pretty well completed in two days and lined on the 3rd and perhaps the 4th. A female will engage in building for from 15 to 23 minutes, then interrupt her work for 5 to 8 minutes.

In the matter of nest building, the young female is in every way the equal of the older and experienced bird in choice of site, skill in construction, quality of the finished structure, and excellence of its concealment.

The start of egg-laying with the song sparrows on Interpont was closely correlated with the temperature in April; in one year, 1929, it was also affected by the temperature in the last third of March. In 6 years the first egg was found between April 15 and 19, but in 1929 it was found on April 10, and in 1932, April 23. The start of general laying was closely correlated with temperature. The “normal” date was April 25, but this was accelerated nearly 2 weeks in 1929 and 4 or 5 days in 1930 and 1931, but was delayed 4 or 5 days in 1933 and 1934.

In eastern North America between the latitudes of Maine and North Carolina, the races of the song sparrow normally nest through July and into August. Drought curtails nesting, however, as it did on Interpont in 1930, when adults began to molt two weeks early. George M. Sutton (1960) found the same to be true on the Edwin S. George Reserve near Ann Arbor, Mich., where nests were active in July and August of 1934, 1935, and 1940, but not in the extremely dry summers of 1936 and 1946.

Eggs: The ground color of the eggs on Interpont ranges typically from blue through blue-green to grey-green. The spots are brown to red-brown and rarely lilac, and are arranged in an endless variety from small speckles nearly uniformly distributed over the whole egg to a few large splotches irregularly placed, usually the larger part of the pigment being around the large end, sometimes in quite a regular ring. Measurements of 503 eggs ranged from 17.5 to 22.5 mm. in length and from 14 to 17 mm. in width, the median being 19.9 x 15.5. As to weight, 44 fresh eggs varied from 1.8 to 2.85 grams, the average being 2.28, the median 2.23.

In 211 nests on Interpont sets of 5 eggs were found in 30 percent of the cases, 4 eggs in 50 percent, and 3 eggs in about 20; the average size was 4.1 eggs. In North Carolina in 175 nests D. J. Nicholson discovered only one 5-egg set; 4-egg sets were in the majority, although there were several of 2 eggs and once he collected a single well incubated egg. When a song sparrow’s nest is destroyed the first egg of the next set is laid 5 days later.

The weight of a set was approximately half the weight of the bird that laid it.

Young: Incubation, which is performed by the female alone, usually lasts slightly over 12 or 13 days but rarely has taken 14 and 15 days with unusually inattentive females. The bird stays on the eggs for 20 to 30 minutes, then leaves for 6 to 8 minutes. The total percentage of daylight hours spent on the nest averages 75 to 80. The male guards his territory, nest, and mate, and does considerable singing. He often calls her off the nest with a sudden loud song.

The young are brooded by the female for the first 5 or 6 days of nest life. The rate of feeding increases with the age of the nestlings; in 7 broods the rate for the first 5 days averaged 7.2 times an hour, for the second 5 days 17.8; the average for the whole period was 11.1 times an hour. The weight of the nestling increases more than tenfold in the first 10 days of life. Young usually stay in the nest 10 days, leaving before they can fly. They become independent of the parents at the age of 28 to 30 days.

After a nest was destroyed, the young of the replacement nest were usually fledged just 30 days later. Periods between the fledging of two broods successfully raised ranged from 30 to 41 days. M. m. euphonia might be called three-brooded, although Schantz (1937) reports that one pair raised four broods in one nest in one season. D. J. Nicholson writes of a similar observation near Tuxedo, N.C. in 1956, his only such record in 56 years of experience.

Plumages: Alexander Wetmore (1936b) described euphonia as similar to melodia but distinctly darker above, being grayer, with the dark markings generally more distinct; sides of head grayer, less huffy or brown; tail averaging darker. In common with other races of the song sparrow in the east, many individuals of euphonia show a distinctly rufescent phase. One breeding bird collected has the brown markings (auburn in color) predominating over any other shade on the dorsal surface.

Behavior: Female song sparrows arose about 5 minutes after civil twilight in clear weather and 7 minutes after civil twilight in cloudy weather. Males did likewise in January and early and late fall; but in early spring they arose at civil twilight, and later in spring and in mid-fall when in full song they rose 4 minutes before civil twilight. Roosting took place at about 10 times the light values of the rising tune of the female. From December through May and again in October the last notes of the males came with surprising regularity 13 —14 minutes after sunset in clear weather.

Song sparrows use several methods for defense of nest and young. They may sometimes threaten an enemy with posture and sound. They attack cowbirds that approach the nest site. Some individuals attack small snakes, while others ignore them. Typically solicitude for the nest reaches its peak when the young are ready to leave it. Warning notes from the parents at the approach of an intruder induce silence in the young. Distraction display, in which the bird runs about close to the observer with wings held stiffly erect and tail depressed, was typically shown when young of 6 or 7 days shrieked when being banded. Occasionally parents will try to lure young to safety by bringing food near, then hurrying awa’y with it.

Experiments with enemy recognition, as well as observations in nature, led to the conclusion that owls are recognized by song sparrows largely through an inborn pattern, hawks through their rapid movements, and cats and cowbirds through conditioning. With a handraised bird, memory of circumstances connected with strong alarm persisted after 4 to 19 months, a response that has definite biological value (Nice and ter Pelkwyk, 1941).

Anting was observed in 3 of the hand-raised song sparrows; both ants and sumach berries were used. Anting first appeared at the age of 36 to 37 days (Nice and ter Pelkwyk, 1940).

Although the song sparrow is only slightly gregarious, it shows the basic mechanisms for social integration (Nice, 1943.)

Voice: Eight different kinds of vocalizations were heard from young song sparrows, 16 from adult males, and 15 from females. Songs are given at the rate of 5 to 7 a minute when the bird is singing steadily, but occasionally during encounters involving territory establishment, the rate may be 10 times a minute. Sometimes a primitive-sounding, irregular flight song is uttered. Young birds change often from one song to another; at the height of vigor there is sustained effort and one song may be sung 60 or even 70 times in succession; an old bird may change from one song to another somewhat more often. The greatest number of songs in one hour was 325, and in one day 2,305 from a bird 8 or 9 years old.

Female song sparrows occasionally sing early in the season before nesting begins; the song is given from an elevation and is short, simple, and unmusical. The most energetic singers were also zealous in chasing male neighbors.

There are 5 chief stages in the development of song in the young bird: continuous warbling, advanced warbling with some short songs, predominantly short songs, songs practically as in adult but with repertoire undetermined, songs as in adult with repertoire fixed.

This species, where each male has a quota of songs peculiar to himself, but where there are occasional duplications in a community, offers an opportunity to test the matter of inheritance or learning of song with banded birds. The possession of similar songs was no proof of close relationship between the singers, as neither brothers, fathers and sons, nor grandfathers and grandsons had similar songs, while birds known to be unrelated occasionally had similar songs. With two hand-raised brothers (melodia) from Massachusetts exposed only to records of songs of English birds, the form, length, and timing of their songs were typical of the species, but the quality was atypical and may have been suggested by the recorded songs. The following year, a nestling euphonia hatched 600 miles from the hatching site of the two brothers, heard one of them sing a small amount in the fall but heard no other singing; in December this euphonia burst into song with all of the 6 songs he had heard and with nothing else. It is evident that the pattern is innate, but that quality may be imitated. Particular songs may be improvised, or they may be adopted from some other song sparrow.

Enemies: The song sparrow, like other small passerines, suffers from a multitude of adverse factors: weather; predators, native and introduced; the brood parasite, the cowbird; and man with his destruction of habitat, his inventions: windows, rat traps, automobiles, lighthouses, ceilometers at airports: shooting by his offspring in mistake for “English sparrows.” The introduced predators: cat, rat, and dog: appeared to be more destructive than native predators: snakes, birds, and mammals. On Interpont about 26 percent of the song sparrow nests found were parasitized by the cowbird in 1930 and 1931, 58 percent of the early nests in 1932, 36 percent in 1933 and 69 to 78 percent during the next 3 years. From 0 to 5 song sparrows were raised in 28 nests along with 1 cowbird, from 0 to 2 song sparrows along with 2 cowbirds. Sixty-six successful nonparasitized nests raised an average of 3.4 song sparrows, while 28 successful parasitized nests raised an average of 2.4 song sparrows. So each cowbird appears to have been raised at the expense of one song sparrow.

D. J. Nicholson found no cowbird eggs or young in any nests during 9 years of observation in North Carolina. But in July, 1954, he saw a fledging cowbird being fed by a Mississippi song sparrow. “Apparently cowbirds are extremely scarce in that region of North Carolina and have just come there,” he wrote.

Turning to egg and nestling losses from all causes, 44 percent of the eggs laid left the nest as fledged young during the first 2 years of the Interpont study and 29 percent during the next 4 years; 36 percent of the eggs yielded fledglings during the 6 years. The first 2 years, during which conditions for reproduction were favorable, correspond well with results of other studies on altricial young in open nests. The poor success afterwards reflected the disturbed conditions of the environment. The percentage losses suffered by eggs and nestlings of 906 eggs laid during 7 years was: flood 2.8 percent, predators 36.7 percent, cowbird 6.1 percent, sterile and addled eggs 5.6 percent, parental failure 2.4 percent, man 4 percent, parents killed 3.7 percent, young starved 3.1 percent. For all eggs laid these figures represent a death of 40.9 percent in the egg stage, and of 23.5 percent after the nestling stage was reached: 64.4 percent in all.

The survival of the adult breeding males averaged over 60 percent during the first 2 years, but after that dropped to 48, 23, 30, and 20 percent. The loss of breeding males from April to June averaged 15 percent during the first 3 years, 30 percent during the next 3; the loss of breeding females averaged 30 percent and 35 percent during the same periods. The proportion of first year males in the population ranged from about 26 to 55 percent.

Out of 317 fledged nestlings 26 males and 14 females were later found as breeders. The late arrival of young females doubtless raises difficulties in the way of return to the vicinity of the birth place. The percentage of yearly return of birds, without regard to sex, ranged from 4.5 to 20, averaging 12.6; and 4.5 percent of the eggs laid produced birds that returned to breed on the area. If the population is to be maintained it is estimated that an average of 20 percent of the fledged young should survive to adulthood. It is believed that 50 to 60 percent of the young song sparrows that survived to breed returned to the place of birth to do so.

In a well-situated group of song sparrows the average life of the breeding males was 2½ to 2¾  years. Individuals have been known to have reached 7 years of age, while one male was at least 7½ , and perhaps 9½  years old at his death.

Fall: There was always singing in the fall from the males on their territories, much in mild weather, little in bleak. Usually the summer residents sang for only a few days the last of September and first of October, but some residents might sing well into November. In each of 5 years one banded male sang for periods lasting 40 to 65 days. The character of the autumn singing differed little from that of the height of the nesting season, except that there were more incomplete songs.

The fall migration of the transients in central Ohio takes place in late September and throughout October. Summer resident females left by the middle of October, males a little later. Bleak weather tended to hasten migration, mild weather to delay it or even occasionally to supress it.

Winter: Resident song sparrows stay on or near their territories throughout the year, although they do not defend them in winter. At this season they may range over an area 6 to 10 times as large as their breeding territories. In cold spells they sometimes come more than a quarter of a mile to a feeding station. In snowy weather they assemble in loose flocks which are neither family parties nor neighborhood groups, as they are composed of both residents and winter residents. (Families break up at the end of the nesting season.) The resident males start their singing and take up territories during warm weather in late January or early February, but a return of winter brings a return to winter behavior.

I never had a recovery away from Interpont of the 870 birds banded. A song sparrow banded June 8, 1932 near Cleveland, Ohio, was taken in Janesboro, Ga., Dec. 25, 1933.

[Note: Because Mrs. Nice’s account of euphonia is essentially a summary of her banding study of an Ohio population, it is desirable to add a word about the race in other places, and especially in the Appalachian Mountains. Indeed, the old vernacular name Mississippi Song Sparrow was rejected by some (see Burleigh, 1958) as being less appropriate than “Appalachian Song Sparrow.” — V.N.]

Arthur Stupka (1963) writes that this race is common in the lower altitudes of the Great Smoky Mountains National Park but breeds at any heights there. The habitat above 6,000 feet is “brushy openings in these moist highiands.” In the mountains of Georgia, on the other hand, Burleigh (1958) states that the bird is fairly common in “thickets and underbrush in the valleys. It has no liking for thick woods and will never be seen on mountainsides or in wooded ravines.

Interestingly, both the foregoing authors have noted an extension of euphonia’s range. It appears to have moved into the high altitudes in only recent years, and Stupka discusses the possibility that large-scale habitat changes attributable to lumbering and chestnut blight have thinned the forest overstory and produced the changed distribution of the bird. In Georgia, the breeding range of the species has been extended even more markedly. “Less than fifty years ago, it was not known to breed in Georgia * * * (Burleigh, 1958), but by 1945 it had penetrated southward beyond the mountain valleys to the northern edge of the Piedmont.”

Milton B. Trautman (1940) has given a most comprehensive description of the habitat preferences of this race in a large and varied area around Buckeye Lake in central Ohio:

In winter the bird was abundant in brushy thickets, fallow fields of rank weed growth, weedy fields of uncut corn, brushy edges and openings of woodlands, brushy fence rows, brushy and weedy swamps, weedy edges of the canal, and cattail marshes. It was less numerous or entirely absent from the bleak snowswept fields and pastures, from the vicinity of farmhouses containing little shrubbery, and from orchards and woodlots which had no ground cover. During spring and fall it was also noted in small numbers in more barren localities, such as pastures, meadows and last year’s forage and grain fields.

In the nesting season, from late March to early August, the species was found in the greatest concentrations in lowland weedy situations. Many pairs nested on the islands and lowland shores of the lake where herbaceous plants and brush were present, about weedy and brushy borders and isolated brushy areas of cattail swamps, in weedy and brushy inland swamps, in brushy edges and openings of woodlands, along brushy fence rows, in fallow and weedy fields, and in the openings and borders of brushy thickets. A few also nested along weedy or brushy fence rows, in fields of smaller grains and forage crops, and in woodlots containing little ground cover. It was only in the barren upland fields and on the crests of wooded hills that nesting birds were absent.

DISTRIBUTION

Range: Northern Wisconsin, northeastern Michigan, southern Ontario, and western New York south to south-central Texas, the Gulf coast, and southern Georgia.

Breeding range: The Mississippi song sparrow breeds from northern Wisconsin, northeastern Michigan (Marquette, Whitefish Point), central southern Ontario (Bruce County, Hamilton), and western New York (east to Keuka Lake) south through southeastern Minnesota and Iowa to northeastern Kansas (Bendena), southwestern Missouri (Jasper County), and northwestern and north-central Arkansas (Winslow, Newport); through western Pennsylvania, western Maryland (Accident), West Virginia (except extreme northeast), southwestern Kentucky (Paducah, Glasgow), southwestern Virginia (Pulaski, Marion), southeastern Tennessee (Chattanooga, Crab Orchard), northeastern Alabama (Valley Head), and western North Carolina to northern Georgia (Milledgeville) and northwestern South Carolina (Clemson); casual in summer in south-central Kansas (Harper) and northern Louisiana (Tallulah).

Winter range: Winters from southern Wisconsin (Yiroqua, Green Bay), southern Michigan (Alicia), southern Ontario, and western New York southwest and south through southeastern Nebraska, eastern Kansas (Douglas County), and central Oklahoma (Norman) to south-central Texas (Fort Clark, Matagorda), southern Louisiana (Main Pass), southern Mississippi (Bioxi), southern Alabama (Petit Bois Island), southern Georgia (Grady County, St. Simons Island), and South Carolina (Kershaw County, Mount Pleasant); casually in northern Michigan (MeMilan) and western Kansas (Seward County).

DAKOTA SONG SPARROW

MELOSPIZA MELODIA JUDDI Bishop

Contributed by VAL NOLAN Jr.

HABITS

This breeding race of the great plains of Canada and of the northern United States withdraws from much of its range in winter and spreads southward and eastward as far as Florida. Early migrants return to Canada in mixed flocks with tree sparrows, juncos, fox sparrows, and other fringillids in March and April (Houston and Street, 1959).

Thomas S. Roberts (1936) has described the habitat in Minnesota: “Bushy meadows and the banks of lakes and streams are the chosen dwelling-places, but it is not confined to such surroundings and may be found almost anywhere about prairie groves, upland fields, clearings, and gardens, shunning only the deep shade of heavy timber.” Roberts also quotes T. Martin Trippe, a pioneer Minnesota bird student, who in 1871 noted that this shy bird of the brush prairies and thickets, near water, was changing from “almost the wildness and timidity of a wild-duck” to a confiding neighbor of man.

Territory sizes under favorable conditions would probably approximate those stated above for euphonia and suggested for melodia. However, Beer et al. (1956) report that song sparrows nesting on small islands in Basswood Lake, Minn., regularly hold territories as small as 0.04 acre, which is the total area of each of two of the islands. Indeed, territories of 0.05 acre each were held by two pairs present on a single island, but one male was “definitely subordinate.” Suthers (1960) measured lake-shore territories on Lake Itasca, Minn., “to retain the effects of shore line on size” while avoiding the probable effects of insularity; territory sizes varied from 0.34 to 0.68 acre, averaged 0.47 acre, and were thus intermediate between those of island and mainland territories. Comparison with work of Tompa (1962) on the race morphna, below, is suggested.

M. m. juddi is among the first birds to sing in spring, and descriptions of the song resemble those of the eastern races. However, Louis B. Bishop (1896), who described the race, found the song “quite different,

* * * clearer, sweeter, more powerful. * * * I could not believe [it] was a song sparrow until I had the bird in my hand.”

No differences between the reproductive cycle of juddi and of melodia and euphonia are apparent in Roberts (1936) and other literature. Beer and his colleagues (1956) state that on the small islands in Basswood Lake, Minn., “Nesting, while normally on the ground, may take place in holes in trees or in small evergreens as much as seven feet off the ground where there are not suitable open areas.”

Roberts describes an interesting instance of predation ov a nest by a garter snake which suddenly appeared, and seized and made off with a newly hatched nestling “in spite of a vigorous attack by one of the parents. When pursued the snake quickly swallowed the tiny nestling.” Roberts opened the snake (a gravid female) and found a second song sparrow nestling in it. As garter snakes are common in the nesting habitat of song sparrows, the episode was probably typical.

L. B. Bishop (1S96) described juddi as being similar to the race presently known as M. m. melodia, “but with the ground color of the upper parts paler, especially the superciliary streak and sides of neck, and the white of the lower parts clearer; the interscapulars with the black center broader, the reddish-brown portions narrower, and the gray edgings paler; the dark markings on the breast restricted, and more sharply defined against the ground color.”

DISTRIBUTION

Range: Northeastern British Columbia, southern Mackenzie, northern Manitoba, and northern Ontario south to southern Texas, the Gulf coast, and central Florida.

Breeding range: The Dakota song sparrow breeds from northeastern British Columbia (near Peace River), central southern Mackenzie (Great Slave Lake), northern Saskatchewan (Lake Athabaska), northern Manitoba (Knee Lake; casual at Churchill), and northern Ontario (Fort Severn, Attawapiskat Post) south through the plains of Alberta (Grand Prairie, Calgary, Milk River) and eastern Montana (Miles City) to northern Nebraska (Sioux County, Dakota City), northwestern Iowa, southern Minnesota, extreme northwestern Michigan (Baraga County), and southwestern Ontario (Amyot); casual in southern Nebraska (Red Cloud).

Winter range: Winters from southeastern Montana (Miles City), South Dakota (Yankton), and southern Minnesota (Cambridge) south and east to western and southern Texas (Fort Davis, Boquillas, Del Rio; Atascosa County; Longview), Louisiana (Lake Charles), southern Mississippi (Petit Bois Island), Georgia (ha, Tifton), central Florida (Enterprise), and southwestern Virginia (Blacksburg); casually to Manitoba (Burnside) and southern Arizona (Tucson).

Accidental on Banks Island, Franklin District.

MOUNTAIN SONG SPARROW

MELOSPIZA MELODIA MONTANA Henshaw

Contributed by VAL NOLAN JR.
HABITS

This is the breeding subspecies of most of the Rocky Mountains of the United States, from the states bordering Canada to those adjoining Mexico. Although it may nest at least as high as 9,000 feet (Betts, 1912), the song sparrow is replaced in the Canadian life zone by the congeneric Lincoln’s sparrow and is ordinarily found at somewhat lower altitudes. In Yellowstone, for example, Milton Skinner (1925) found montana only up to 6,500 feet. At the other extreme, some individuals breed on the plains of Colorado at the eastern edge of the mountains (W. W. Cooke, 1897; Niedrach and Rockwell, 1939).

The habitat of montana is water-edge vegetation, whether it be in marshes around mountain lakes as at Lake Tahoe, Nev. (W. W. Price in Barlow, 1901), in streamside willows as in eastern Oregon (Peck, 1911), or in boggy areas of cultivated fields and meadows as in Colorado (Rockwell, 1908; Niedrach and Rockwell, 1937). Alexander Wetmore (1920) describes the habitat at Lake Burford, N. Mex., as clumps of dead tule (Scirpus occidentalis) fringing the lake, although occasionally birds ventured up into sagebrush to feed or nest.

In the marshes of the Grand Tetons of Wyoming, Salt (1957) studied the ecologic relations of song sparrows (believed to have been montana, because of the location), Lincoln’s sparrows, and fox sparrows. Song sparrows occupied the dense thickets 6 to 10 feet high along open water, where they foraged “often with their feet in the water.” Population density was approximately 3 individuals per 10 acres, and Salt estimated plant food comprised 60 percent of the summer diet.

Spring: Some birds winter throughout the breeding range, while others migrate as far south as the northern states of Mexico. Reoccupation of breeding territories takes place in late March in Colorado (Cooke, 1897) and extends into early April in Montana (Saunders, 1921).

Nesting: Nesting habits are like those of the races already discussed. Ground nests appear to be the commonest and are built in grass, alfalfa, in hollows under sage and willows, among ferns under fallen trees, and in cattails at slight elevations. Elevated nests have been found as high as 11 feet, and willows and lodgepole pines are numbered among the nest trees. The nest itself is described in the same terms as are the nests of the eastern races and apparently does not differ significantly. As an example, M. S. Ray (1910) has reported a nest in a flooded alfalfa field. It was made “of grasses and weed stems with a lining of horsehair, was placed on slightly higher ground,” and well concealed, and contained an unusual clutch of 6 eggs. Nests are sometimes built over water (Rust, 1917). Jean M. Linsdale (1936), in testing the hypothesis that colors of nest linings and downy plumage of nestlings are adapted to the degree of exposure to sun, studied these points in the Toyabe Mountains of Nevada, where the breeding race of the song sparrow is montana. Linsdale found five nests “close to the ground” and “covered” so they were not exposed to sun. The down of the nestlings be classed as intermediate between light and dark in color, but the nest linings were dark, with the result that they would absorb warmth rather than reflect it.

Nesting continues until mid-summer, e.g., in Oregon until at least mid-July.

The male is said by Niedrach and Rockwell (1937) to help in nest building.

Eggs: The measurements of 40 eggs average 19.9 by 14.9 millimeters; the eggs showing the four extremes measure 22.6 by 15.2, 20.8 by 15.5, 18.3 by 14.2, and 19.3 by 13.5 millimeters.

Incubation: Courtship feeding by the male of the female as she sits on the eggs has been attributed to this race by Niedrach and Rockwell (1937). However, because these authors give no details and because Mrs. Nice (1943) did not observe such behavior in euphonia, confirmation seems desirable. David Lack (1940b) in his review of courtship feeding states, “Typically [it] seems absent in * * * American sparrows, including the well-studied * * * Melospiza (Song Sparrow).”

Plumage: Ridgway (1901) states that montana is similar to melodia, “but wing, tail and tarsi averaging decidedly longer, bill smaller and relatively more slender, and coloration grayer; young with ground color or under parts dull white or grayish white, instead of more or less buffy, that of upper parts less tawny than the young of * * * melodia.” H. W. Henshaw (1884) states that fall specimens of montana are “browner” than the race fallax with the markings generally less distinct, i.e., more diffused. The black streaks of the back are always present. M. S. Ray (1913) found a partially albino nestling in a brood of three; “the entire underparts were pure white and iris light reddish.”

Fall and winter: Some birds are found in fall above the breeding altitude, e.g., at 10,000 feet in Park County, Colorado (Warren, 1915). Those that remain in the breeding range through the winter apparently survive by staying close to the moderating influences of water, including warm springs where these occur. Rockwell and Wetmore (1914) found this race common in the mountains along Clear Creek near Golden, Cob., on Nov. 14 “though there was six inches of snow and the bushes were veritable snow banks.” Stanley G. Jewett (1912) has written of seeing several M. m. montana that in December frequented the warm spring flats of the Wood River in Idaho and obtained insects from the muddy ground. The birds were “often seen feeding in the shallow water, while on all sides the snow was piled four feet deep.” Birds that migrate to southeastern California inhabit “thickets of arrowweed, and willows and reeds at the edges of ditches and river courses. The birds forage short distances out into grassy or weedy places and about root tangles and piles of driftwood” (Grinnell and Miller, 1944).

DISTRIBUTION

Range: Northeastern Oregon and north-central Montana to southeastern California, northern Sonora, central Chihuahua, and western Texas.

Breeding range: The mountain song sparrow breeds from northeastern Oregon (Union and Wallowa counties), central western Idaho (New Meadows), and north-central Montana (Missoula and Teton counties, intergrades with M. m. merrilli) south to eastern Nevada (Toiyabe Mountains, Lehman Creek), southwestern Utah (Pine Valley Mountains, Kanab),central eastern Arizona (White Mountains), and northern New Mexico (Santa Fe, Raton).

Winter range: Winters throughout the breeding range and south to southeastern California (Death Valley, Riverside Mountain), northern Sonora (Caborca, headwaters of Bavispe River), central Chihuahua (Chihuahua), and western Texas (Fort Davis, Ingram); east casually to western Nebraska (Crawford), western Kansas (Trego County), and western Oklahoma (Cimarron County).

YELLOWHEAD OR RILEY SONG SPARROW

MELOSPIZA MELODIA INEXPECTATA Riley

Contributed by VAL NOLAN JR.

HABITS

Although Al. m. inexpectata occurs as a breeding bird on the coast and inner islands of the southeastern extension of Alaska, the range of this race also includes much of interior British Columbia and reaches into the mountains of Alberta. This distribution and a difference in its nesting ecology are responsible for the separation of this life history of inexpectata from the combined account of most other subspecies that occur in Alaska.

Both on the breeding and the winter ranges, which do not overlap, this race seems to prefer a marshy or brushy habitat like those, for example, of melodia or juddi. George Willett (1928) writes that inexpectata occurs in southeastern Alaska in inland locations several miles from salt water. “Nests * * * found at Ketchikan were often placed several feet up in trees.” Apparently nests were built near the ocean, as well, for the same writer found a nest “inside a roll of wire netting that was lying just above the high tide line.” Harry S. Swarth (1922) found this song sparrow in relatively few places in the upper Stikine River valley in British Columbia, but near one tributary creek and in the marshy meadows a mile or so back from the river it was abundant. Between July 8 and 26, 1921, Swarth considered full grown young more numerous than adults; the latter were apparently engaged in tending second broods, many probably still in the nest. The adults were very shy and stayed in the nesting areas, which were sloughs grown up with reeds and surrounded by willows. By mid-July, the presumed first-brood young were spreading farther and farther from the marshes and down the river. Another example of the dispersal of young from the breeding habitats is Swarth’s collection of juveniles between July 26 and August 8 in fireweed patches near Flood Glacier where no suitable nesting areas were observed.

Swarth (1924) reported that on the Skeena River near Hazelton, British Columbia, this song sparrow found little suitable habitat and nested only in scattered pairs along small streams. Nesting seemed in progress on May 26, and a young bird was seen being fed by an adult as late as Aug. 29. Most birds had left by the third week of September. Swarth found no song sparrows nesting in the mountains.

Jewett et al. (1953) describe the winter habitat in western Washington as “alder brush and berry tangles of second-growth localities, as well as brush along roadsides and in partly cleared places.”

J. H. Riley (1911) described inexpectata as being “similar to Melospiza melodia rufina, but the browns of the upper parts lacking the reddish tinge, thus giving to the back a gray cast; below not so heavily streaked; averaging smaller.”

DISTRIBUTION

Range: Southeastern Alaska and southern Yukon south to northern Oregon.

Breeding range: The yellowhead song sparrow breeds from the coast and inner islands of southeastern Alaska (Glacier Bay, Admiralty Island, Revillagigedo Island), southern Yukon (Squanga Lake), and northwestern British Columbia (Atlin, rarely) southeast through interior British Columbia to lat. 510 N. (Horse Lake, Yellowhead Pass) and the mountains of southwestern Alberta (Henry House, Banff).

Winier range: Winters from southern British Columbia (Comox, Alta Lake, Okanagan Landing) south through Washington to northern Oregon (Portland, Prineville); casually north to Caribou district, British Columbia (Indianpoint Lake).

MERRILL’S SONG SPARROW

MELOSPIZA MELODIA MERRILLI Brewster

Contributed by VAL NOLAN JR.

HABITS

This migratory race breeds in the interior of the three northwestern states adjoining Canada and of the two southwestern provinces of the latter country. As is true of so many other subspecies of MeIospiza melodia, individuals can be found in winter in locations virtually throughout the breeding range, while others migrate, in the case of this race, to the southwestern United States. Descriptions of the habitat would be generally applicable to those of most of the other inland subspecies: tangles of willows, cottonwood, and alder; open, sedge-grown, brushy meadows; hawthorn brush bordering clearings; and lakeside marshy associations. H. J. Rust (1919) tells of seeing these birds hopping from pad to pad of water lilies and wading into shallow water “reminding one of the Water Ouzel except in color.” At the opposite extreme, they occur in Washington, possibly as breeders, in brush succeeding deforestation at altitudes of 4,000 feet (Jewett et al., 1953). James C. Merrill (1898), for whom the bird is named, states, “There is nothing in their general habits to distinguish them from the Song Sparrows in other parts of the country,” but he notes their very marked preference for the immediate vicinity of water in the type locality, Fort Sherman, Idaho.

Spring: Rust (1919) believed birds that wintered near Fernan Lake, Idaho, to be principally old males, and these started to sing on sunny February days. Females began to join males in mid-March, and the earliest nest was found on Apr. 12, 1918, a year with an unusually warm spring. A like date is given for Washington by W. L. Dawson and J. H. Bowles (1909). At these dates few other species are nesting in the song sparrow’s habitat.

Nesting: Dawson and Bowles (1909) describe a pattern of nest placement that prevails for many other races: First nests are usually built on the ground under cover of vegetation, and elevated nests (up to 20 feet high) become commoner as the season advances. Occasional ground nests may be found at any date, e.g., on Aug. 3, in Washington. On the other hand, Rust (1919) gives an interesting portrayal of a different kind of seasonal change in nest location, with heights tending to move downward from elevated positions to the ground. At Fernan Lake, Idaho, the onset of breeding coincided with high water levels, and early nests invariably were in spirea shrubs and willows whose bases were partly submerged. Receding water in mid-May exposed clumps of sedge, which were then used for some nest sites, although shrubs and willows also continued to be selected throughout the season. J. C. Merrill (1898) never found a nest on the ground, and most were in bushes growing in water. Dawson and Bowles and also Merrill describe clumps of debris stranded in willows by flood water as common nest sites.

J. C. Merrill (1898) writes that nests near Ft. Sherman, Idaho, were “unusually large for a song sparrow” and were built largely of dead leaves and strips of cottonwood bark, deeply cupped and with finer materials for the linings. Rust found sedge the most used plant material and reported that nests built later in the season were more compact and less precariously wedged in forks and supports; horsehair was a common lining.

Eggs: The measurements of 40 eggs average 20.1 by 15.0 millimeters; the eggs showing the four extremes measure 22.4 by 15.5, 20.8 by 16.3, 18.3 by 15.2, and 19.3 by 13.7 millimeters.

Rust (1919) found 4-egg clutches more numerous than 5-egg clutches at Fernan Lake, Idaho. Near Ft. Sherman, Idaho, Merrill (1898) reports 5 as the common size of the first clutch, 3 or 4 as the usual later sizes.

Incubation, young: The incubation period is set by Dawson and Bowles (1909) and by Rust (1919) at 12 days, but Rust’s nestling period, 14 to 16 days, seems remarkably long in view of the periods known for euphonia, 10 days, and melodia, 10 to 14 days. Dawson and Bowles’ statement that young can “fly” when 12 days old should be compared with Mrs. Nice’s finding that 17 days is the age at which young of euphonia commonly begin to fly. Fledglings remain in dense underbrush and sedge until fairly strong on the wing.

Plumage: Ridgway (1901) states that merrilli is “very similar to * * * montana but slightly darker and more uniform above, with the grayish edgings to the interscapulars and scapulars less strongly contrasted with the darker mesial streaks, the latter usually with more brown than black.”

Food: Jewett et al., (1953) tell of the methodical dismemberment, limb by limb, of a moth by a merrilli song sparrow.

Enemies: Both Merrill (1898) and Rust (1919) regard nest losses to flooding as very numerous. As enemies Dawson and Bowles (1909) list the usual nest predators of small passerines, i.e., mammals, snakes, and corvids.

Fall and winter: Individuals wintering in the breeding range resort to tule beds, thickets, brush piles, and the vicinity of outbuildings. Outside the breeding range in California, Grinnell and Miller (1944) report the habitat as “weed thickets, old rice fields, tule beds and willow tangles. In general favors riparian growth and damp places.”

DISTRIBUTION

Range: Southern interior British Columbia and southwestern Alberta south to southern California, southern Utah, and northern New Mexico.

Breeding range: The Merrill’s song sparrow breeds from southern interior British Columbia (south of lat. 51° N.; Alta Lake, Shuswap Falls) and southwestern Alberta (Waterton Lakes Park) soutb to eastern Washington, east of the Cascade Range (Yakima, Wallula; intergrades with M. m. fisherella) , northern Idaho (South Fork Clearwater River), and northwestern Montana (Flathead Lake).

Winter range: Winters from southern interior British Columbia (Okanagan Landing) and northwestern Montana (Fortine) west to western Washington (Destruction and Orcas islands) and south to southern California (Altadena, Victorville), southern Nevada (Charleston Mountains), southern Utah (Santa Clara), and northern New Mexico (Las Vegas, Hot Springs); casually to southern Arizona (Quitobaquito) and northern Sonora (Upper Bavispe River).

MODOC SONG SPARROW

MELOSPIZA MELODIA FISHERELLA Oberholser

Contributed by VAL NOLAN JR.

HABITS

A song sparrow of the interior of the northwestern United States (as far south as Nevada), M. m. fisherella appears in its life history to resemble closely merrilli, one of the races with which it intergrades. Many individuals winter throughout the breeding range, but others migrate, occasionally as far as northern Sonora.

Breeding altitudes in California range between 300 and 8,000 feet above sea level. But everywhere “riparian vegetation, marshes, and lake borders” are sought out, for “the combination of dense low cover and surface water with wet ground is essential for this race” (Grinnell and Miller, 1944). Philip C. Dumas (1950) has documented the foregoing statement about the habitat in a series of detailed density indices for this and other birds in the various environments of southeastern Washington.

Moisture-loving plants such as willows and cattails are the most usual nest sites, which in California “commonly are above ground in the branch-work of bushes” (Grinnell and Miller, 1944). Near Pullman, Wash. (Jewett et al., 1953), nests are often placed between 8 and 12 inches high in willows. Plants of dry associations, e.g., small conifers and shrubs, are also sometimes used for nests, which have been found as high as at least 7 feet (Mailliard, 1919b). Ground nests are also found. Nests are described as loosely woven of weeds and grasses. L. R. Dice (1918) once found a structure in which some of the material had been picked green. W. L. Dawson (1923) reports a most unusual nest in Modoc County, California; it had been built into the side of a large paper wasp nest hanging 5 feet up in a willow, and scraps of newspaper were among the materials used in the construction of the outer surface. In Washington full sets of fresh eggs have been found as early as March 30, and L. R. Dice (1918) found well feathered young in a late nest on July 13.

Feeding “occurs on the moist ground or at the water’s edge, or on plants over the water or floating in it,” state Grinnell and Miller (1944); and Jewett et al. (1953) describe some birds as “almost semiaquatic” in their foraging on the floating green leaves of water plants. H. C. Bryant (1911) found only insects in one stomach he examined.

Grinnell and Miller state: “In winter, restriction to riparian growth is less rigid, but seldom are the birds found far from moist situations and the associated plant species.”

H. C. Oberholser (1911) described fisherella as being similar to heermanni, “but larger; upper surface paler, less rufescent; streaks on lower parts less blackish (more brownish).” M.m. fisherella differs from montana “in its darker upper parts, more blackish brown streaks of under surface, heavier bill, and shorter wing.”

DISTRIBUTION

Range: Northeastern Oregon and southwestern Idaho south to southern California.

Breeding range: The Modoc song sparrow breeds from northeastern Oregon, east of the Cascade Range and west of the Blue Mountains (The Dalles, Pendleton; intergrades with M. m. merrilli in southern interior Washington, and with M. m. montana in central Baker County, Oregon), and extreme southwestern Idaho (Weiser, Jordan Creek) south to south-central Oregon (Medford), north-central and central eastern California (Hayfork, Red Bluff, Mohawk, Olancha), and western Nevada (Santa Rosa Mountains, Fish Lake Valley).

Winter range: Winters throughout the breeding range and south to western and southern California (Laytonville, Hayward, El Monte, Calipatria); rarely to northern Sonora (Caborca) and southern Arizona (Tucson).

SONG SPARROW: ALASKAN SUBSPECIES

MELOSPIZA MELODIA (Wilson)

The following subspecies are discussed in this section: Melospiza melodia maxima Gabrielson and Lincoln, M. m. sanaka McGregor, M. m. amaka Gabrielson and Lincoln, M. m. insignis Baird, M. m. kenaiensis Ridgway, and M. m. caurina Ridgway.

Contributed by VAL NOLAN JR.

HABITS

The six races grouped for the purposes of this life history have in common the facts that they breed only in Alaska and that they are largely birds of the ocean beaches. Two of the races, kenaiensis and caurina, are migratory to some extent and breed, in part, on the mainland, whereas the other four races are almost exclusively island inhabitants and are resident. Otherwise the life histories of these six subspecies are probably much alike. Information referable to a particular race is so indicated.

Two additional races that breed both in Alaska and in British Columbia, inexpectata and rufina, are treated separately, because of the differences in their habitats, which are sometimes “inland localities several miles from salt water * * ~ according to George Willett (1928). The same experienced observer states that the other Alaskan races (recognized at the date he wrote) “appear to be strictly beach birds, nesting and feeding within a few yards of the beach.”

Gabrielson and Lincoln in their work (1959) on Alaskan birds write:

The Alaskan Song Sparrows have become rather specialized birds, adhering very closely to the sea beaches throughout most of their range, and only in the southeastern district from Yakutat Bay south do they show any resemblance to the habits so familiar to bird students elsewhere. * * * The species has one of the most curious ranges of any in Alaska. It is found along a narrow coastal strip from Dixon Entrance [at the boundary of Alaska and British Columbia] to Attu [the outermost Aleutian] * * * and only as straggling individuals in the Bristol Bay area, on the Pribilofs, or on the north side of the Alaska Peninsula.

It is also one of the most variable of Alaskan birds. The races uf southeastern Alaska are dark and relatively small * * ~’ï Northward and westward from Yakutat, however, the birds become larger and paler, reaching the grayest race in the Alaska Peninsula and in the easternmost of the Aleutian Islands [sanaka], and then becoming browner but not much darker in the western Aleutians [maxima].

Ridgway’s measurements of the length, although taken from dried skins, give some indication of the variation in the size of these birds as one passes to the west. Males of the southeastern race (rufina) average 159 mm. in length while those of the Aleutian Islands (maxima) average 187 mm. * * * These big birds * * * with the normal markings greatly obscured and softened look entirely unlike their more sharply marked relatives to the east; and yet, they sing about the same song. It is an unusual experience for an ornithologist to watch one ef these comparatively gigantic Song Sparrows behave exactly as do their small counterparts far to the south.

Spring: Those individuals of the two migrant races that have wintered in southeast Alaska, British Columbia, and the Pacific states of the United States probably return to the breeding range in March and April. Thus caurina is found in California until March, and Swarth (1911) took specimens of that race in the Alexander Archipelago just south of the breeding range in middle and late April. Gabrielson and Lincoln (1951) mention a specimen of kenaiensis collected in southeastern Alaska outside the breeding range on Feb. 6.

The spring behavior of the resident races may perhaps be typified by that of maxima on Attu, as described by Sutton and Wilson (1946) for the period Feb. 20 to March 18. “As a rule we saw the birds in twos, and we believe that most of these were actually mated pairs. They were not in breeding condition, however (the gonads of specimens examined being unenlarged), and we saw little in the way of courtship, few pursuit flights of any sort, and no copulation. Singing we heard now and then on windy days, but it was especially noticeable in calm, sunny weather.”

Nesting: Richard F. Johnston (1954), from a study principally of egg collections, fixes the duration of nesting in the Alaskan Peninsula and the Aleutians (presumably, sanaka and maxima, and possibly kenaiensis, insignis, and amaka) at about 7 weeks and indicates that breeding in Alaska begins on about May 15, reaches a peak on about June 10, and ends about July 5.

Gabrielson and Lincoln (1959) say,

Nests are almost invariably placed either on the ground or very close to it in a clump of grass. Those in the Aleutians and along the south side of the Alaska Peninsula are usually built in the beach grass just above the high tide line or in the same grass in the little bays and stream bottoms which it occasionally follows for a short distance into the interior. In southeastern Alaska, it builds in similar situations, although sometimes the nests are woven around the taller grass stems, and elevated a few inches above the ground. The nests are rather bulky and rough on the outside, and are usually built of coarse grass stems not too well put together. They are usually lined with finer dried grasses that are more carefully woven into the interlining.

Joseph Grinnell (1910b) describes two nests of M. m. kenaiensis. Both were placed in beach grass growing on sand spits and both were alike in structure, composed principally of coarse, dry, mildewed grass stalks, arranged concentrically but not intertwined, “so that if roughly handled the nest would readily fall to pieces. The inner lining is thin and of fine, round, yellow grass stems, the majority approaching a position parallel to the rim of the nest. Although many stems are also incorporated cross-wise, the interweaving is not a conspicuous feature. The inner wall of the nest has a slippery feeling because of the smoothness of the grass-stems and the ease with which they slide, one over another; and it is extremely porous. Yet there is a moderate firmness about the whole structure.” The dimensions of one nest were taken after shipment and may have been inaccurate; the diameter externally was 142 millimeters and internally 70 millimeters, while the depths were 67 and 32 millimeters externally and internally.

Two ground nests of sanaka that Swarth (1934) examined were made of grass, one built in a hollow, fairly well concealed by overhanging ferns and salmon berry on a steep mossy bank, the other on a hillside 30 feet above the water. One of these had an external diameter of about 150 millimeters and an internal diameter of 80 millimeters. R. C. McGregor (1960) describes a nest of san,aka in similar terms; it was in the face of a low cliff and had external and internal diameters of 140 and 160 millimeters and depths of 100 and 60 millimeters, respectively.

J. C. Howell (1948) found eight nests of insignis on the ground on Kodiak Island.

Eggs: The measurements of 20 eggs of M.m. sanaka average 24.1 by 17.4 millimeters; the eggs showing the four extremes measure 25.9 by 17.8, 24.5 by 18.3, and 21.8 by 16.3 millimeters.

The measurements of 20 eggs of M.m. insigni.s average 22.5 by 16.7 millimeters; the eggs showing the four extremes measure 23.9 by 16.3, 23.4 by 17.3. 21.3 by 16.8, and 21.8 by 15.8 millimeters.

Johnston (1954), from 17 records of clutch size of unspecified Alaskan races, found an average first clutch of 4.00 eggs and an average second clutch of 4.33 eggs, with the mean clutch size of the sample being 4.17 eggs. The nest of sanaka McGregor found, described above, had three eggs, and those found by Howell contained from three to five eggs.

Young: Swarth (1934) tells of finding families of young sanaka hopping about the face of a cliff and among boulders on the shore of Unalaska on June 15.

Plumage: Tbis summary of descriptions of the races under consideration is taken from the paper by Gabrielson and Lincoln (1951), which described maxima and amaka and reviewed the Alaskan song sparrows, and from Ridgway (1901).

M. m. maxima: “Separable from sanaka, to which it is most nearly related, by the following characters: bill slightly heavier and averaging somewhat longer, especially in the males; in breeding plumage back and head distinctly brownish in tone rather than grayish. This is due to the wider and heavier brown stripes in the center of the feathers of the back and to a darker brown color of the head. In specimens of sanaka in comparable plumage, the brown feather markings are narrower and more obscured, so that the general effect is an over-all grayish tone of the head and back.

“The brownish appearance also is conspicuous in the fall, * * * [and] a comparable difference is noticeable in the juvenal plumage” (Gabrielson and Lincoln, 1951).

M. m. sanaka: Ridgway’s (1901) description of sanaka (at that date cinerea) is that the “general color above [is] olive-gray (almost ash-gray in summer), the back broadly streaked with brown (usually inclosing narrow blackish shaft-streaks), the pileum usually with two broad lateral stripes of light vandyke or mummy brown (these often obsolete in worn summer plumage); streaks on chest, etc., varying from light grayish brown to rusty brown. Young similar to the young of * * * insignis, but paler above and streaks of under parts grayish brown instead of sooty brown.” The bird is similar to, but larger and grayer than, M.m. insignis.

M. m. amaka: “Resembles maxima from the western Aleutians in color and extensive brown markings, but somewhat more heavily marked with brown than that race both on back and breast * * * Closer in color to maxima than to the geographically closer race sanaka. Bill short and stubby as in sanaka” (Gabrielson and Lincoln, 1951).

M. m. insignis: “This race is somewhat smaller than sanaka and is darker, with a sooty wash that noticeably obscures the markings and tends to make the color more uniform. it is, however, paler and grayer than [kenaiensis] * * *” (Gabrielson and Lincoln, 1951).

M. m. kenaiensis: Ridgway (1901) states that this race is smaller and browner than in.signis with the streaks on the chest and elsewhere darker. It is intermediate between kenaiensis and caurina, being larger with the upper parts more uniform in color and less streaked than caurina. “Young, much resembling * * * insignis but more heavily streaked below; much paler and browner above than young of * * * caurina, with streaks on back much narrower, those on chest, etc., much browner.”

M. m. caurina: This subspecies is intermediate between kenaiensis and rufina, which is a sooty brown form of the outer islands of Alaska and British Columbia. Caurina “is smaller and darker, with the streaks more distinct on the back [than kenaiensis], whereas it has a longer bill and grayer coloration than rufina” (Gabrielson and Lincoln, 1951).

Food: Although the song sparrows of Alaska eat seeds when they are available, Gabrielson and Lincoln (1959) note that they take a considerable amount of small marine life, particularly in western Alaska. “Gabrielson has seen them picking up small mollusks or crustaceans as well as such seeds and berries as may be found close to the water line.” Reported food items are beach fleas, crowberries, seeds of wild rye grass, and “many of the smaller alpine and tundra plants that grow close to the water’s edge.” The birds also undoubtedly eat insects. G. M. Sutton and R. S. Wilson (1946) state that maxima on Attu in February and March feeds on tidal flats side by side with rock sandpipers. A specimen examined smelled like the sandpipers and had eaten tiny snails. Swarth (1912) quotes Allen Hasselborg as saying that winter song sparrows around Juneau were very fat and had masses of unrecognized slimy matter in their stomachs, food that had been gathered on the beaches.

Behavior: Ornithologists familiar with song sparrows in other environments will find the beachcombing habit of the Alaskan races of special interest. Thus Gabrielson and Lincoln (1959) state that “the Song Sparrows most frequently seen at Seward * * * are those that live along the beach and under the main dock of that town. On the west side of the Kenai Peninsula, [they] * * * are usually seen only around the wharves and waterfronts of * * * towns * * From Kodiak west through the Aleutians and the other islands * * * it is most frequently seen as a beach bird which, when disturbed, flies up into the rocks or disappears into the openings of a talus slope.” The same authors state that the birds of southeastern Alaska, presumably inexpectata, rufina, and perhaps caurina, are sometimes found in the same bushy habitats, including ornamental shrubs around buildings, frequented by more southern races. The behavior of such birds is correspondingly like that of the southern forms.

Sutton and Wilson (1946) give a notably interesting and detailed report of the behavior of maxima on Attu in February and March, when

The only vegetation is coarse grass, most of which has been blown Hat and buried under the snow, and the bare stalks of high weeds which offer little shelter. * * * Its manner is not quite orthodox [for a song sparrow]. It seems to move too slowly and it keeps under rocks too much of the time. * * *

Most of them lived along the shore, spending virtually all their time between the water’s edge and the snow: a coastal strip varying in width from a few feet at high tide to a hundred yards or so at low tide. * * *

Certain pairs lived about scrap heaps of metal, piles of gasoline drums, and so forth, as Song Sparrows might be expected to live about brush piles. * * * The usual roosting place was a niche or crevice on the face of a big rock, or under a tussock on a turfy headland. * * * Between 5 o’clock in the afternoon and dark, * * * we frequently saw the sparrows going to roost or flushed them from their roosting places. One which we saw asleep was fluffed up, with the bill stuck between the back and scapular plumage, chest against the bare rock, and tail out, down and slightly spread. Others, which flew out as we climbed about the rocks in the twilight must have been sleeping under the rocks or on the bare ground.

Voice: Gabrielson and Lincoln (1959) state that the song is “rather persistent” during the breeding season, but not heard at other times. On the other hand, Sutton and Wilson (1946) heard it in February from males of maxima whose gonads were not in breeding condition. The song is variable and to Gabrielson and Lincoln “enough like that of the same species on the Atlantic coast * * * to be easily recognized * * *ï” (These authors make this statement only about birds of the western Aleutians [maxima] but presumably select this most remote race in order to make the general point.) Sutton and Wilson speak of a slight initial unfamiliarity in the song of maxima but seem to attribute this to the unfamiliar environment and conclude that the vocalization is “fairly average after all * * They also state that an ornithologist from the eastern United States will instantly recognize the Aleutian song sparrow’s husky, alto “chirp of alarm.” “lie will recognize also a higher pitched chip, which indicates greater excitement, and certain beady cries which accompany pursuit flights.”

Fall: A specimen of sanaka collected July 28 had a very short, new tail, and a male and female of that race collected Aug. 3 had half-grown tails, new feathers of primaries 3 through 7, and were undergoing body molt (McGregor, 1906).

The resident races of the western islands leave the immediate vicinity of the shore after the young have been raised and wander some distance up the side of the mountains until snows force a return to the beaches (Gabrielson and Lincoln, 1959; R. D. Taber, 1946, referring to maxima).

Winter: The behavior of maxima, as Sutton and Wilson (1946) describe it, may be inferred to be typical of that race during the winter and not likely to differ much from that of any of the other sedentary Alaskan races. Wilson, in another paper (1948) states that on Attu song sparrows usually stayed in pairs the year round and that he noted them only “occasionally singly, never in a flock.”

Those birds of caurina (and apparently kenaiensis) that migrate remain essentially shore dwellers on the winter range. Thus Alfred Shelton (1915) reports collecting two specimens of caurina in Oregon after he saw one fly up to elude a breaker and then light again at the surf’s edge. A similar observation made by C. I. Clay is reported by Grinnell (191 Oa), who also indicates that the bird was adhering closely to a winter foraging range 200 yards long on a strip of shore in California. The bird was seen repeatedly feeding among driftwood, and it could never be driven beyond the bounds of its range. Its flights were short and apparently undertaken reluctantly, and it preferred to skulk on foot. Interestingly, song sparrows were said by Allen Hasselborg, quoted by Swarth (1912), to have been distributed singly along 200-yard stretches of stony beach at Juneau, Alaska, in winter.

DISTRIBUTION

Giant Song Sparrow (M. m. maxima)

Range: The giant song sparrow is resident in the Aleutian Islands, Alaska, from Attu Island to Atka Island.

Aleutian Song Sparrow (M. m. sanaka)

Range: The Aleutian song sparrow is resident in the eastern Aleutian Islands, Alaska (Seguam Island to Unimak Island), the Alaska Peninsula east to Stepovak Bay, and the islands south of the Alaska Peninsula from Sanak Island to the Semidi Islands.

Casual records: Casual in fall and winter on the Pribiof Islands (St. George) and the coast of western Alaska (Nushagak).

Amak Song Sparrow (M. in. amaka)

Range: The Amak song sparrow is resident on Amak Island, north of the western end of the Alaska Peninsula.

Bischoff’s Song Sparrow (M. in. insignis)

Range: The Bischoff’s song sparrow is resident in the Kodiak Island group (Barren Islands to Sitkalidak Island) and the adjacent Alaska Peninsula (Kukak, Katmai).

Kenai Song Sparrow (M. m.kenaiensis)

Range: Cook Inlet to southeastern Alaska.

Breeding range: The Kenai song sparrow breeds on the coast of southern Alaska from Cook Inlet (Seldovia, Hope) to the mouth of the Copper River.

Winter Range: Winters in the breeding range and southward in southeastern Alaska (Sitka); rarely to the coast of western Washington (Marysville).

Yakutat Song Sparrow (M. m. caurina)

Range: Coast from Yakutat Bay in southeastern Alaska to northern California.

Breeding range: The Yakutat song sparrow breeds on the coast of southeastern Alaska from Yakutat Bay to Cross Sound.

Winter Range: Winters from southeastern Alaska (Chichagof Island, Juneau, Wrangell, Howkan) south along the marine shore lines of British Columbia, Washington, and Oregon to northern California (Fortuna); rarely to central California (Bay Farm Island).

SOOTY SONG SPARROW

MELOSPIZA MELODIA RUFINA (Bonaparte)

Contributed by VAL NOLAN JR.

HABITS

This race, although it breeds on the outer islands of southeastern Alaska and British Columbia, is not confined to the beaches. George Willett (1928) states that rufina occurs “in inland locations several miles from salt water * * *.” Near Sitka, the same observer (1914) found the bird common in summer “in brush and grass lands on islands and along the shore.” Individual birds occur in winter in the breeding range, but some migrate as far south as western Washington. George Willett (1921) reports that rufina and caurina both remained at Craig, Alaska, through the winter of 1919, but that in 1920 at Wrangell, rufina had departed by early November. “This latter place, though only about a hundred miles distant from Craig is, by virtue of its proximity to the mainland, considerably colder.”

Willett (1928) says all nests of rufina are “either flush with the ground or in short grass a few inches up.” Gabrielson and Lincoln (1959) quote the same observer as having found nests with eggs or young from June 2 to July 22.

The post-breeding and winter habitats of rufina are substantially unlike those of the six races reported immediately preceding. Of birds found in September on Mt. Rainier, Washington, Jewett et al. (1953) write: “It seemed to prefer the alder and huckleberry brush of moist, spongy meadows, as well as willows about lake margins.” They add: “Coastal records of the sooty song sparrow are notably scarce, though it appears to be of general occurrence in winter on the islands of Puget Sound and at certain lowland localities in western Washington away from the coast.” Specimens have been collected in Washington between Sept. 3 and Feb. 20.

Ridgway (1901) describes rufina as being “Similar to M. c. morphna but decidely larger (except bill), with coloration darker (sooty rather than rusty), and more uniform above; general color of upper parts deep sooty brown or bister, brightening into rusty brown or chestnut on outer webs of greater wing-coverts and tertials, the back obsoletely streaked with darker, and the median crown-stripe indistinct or obsolete; streaks on chest, etc., deep prouts brown.”

DISTRIBUTION

Range: Outer islands of southeastern Alaska and of central British Columbia south to western Washington.

Breeding range: The sooty song sparrow breeds on the outer islands of southeastern Alaska (Chichagof to Forrester and Duke islands) and of central British Columbia (Queen Charlotte Islands, Porcher Island, Spider Island).

Winter range: Winters in breeding range (north to Sitka), ranging south to western Washington (Whidbey Island, Toledo).

RUSTY SONG SPARROW

MELOSPIZA MELODIA MORPHNA Oberholser

Contributed by VAL NOLAN JR.

HABITS

Another race from the Pacific Northwest, M. m. morphna breeds in southwest British Columbia and western Washington and Oregon. Bird banders have found many wholly sedentary individuals, but there is also a migratory element. The winter range reaches to California and, rarely, Nevada.

The habitat is well described by Jewett et al. (1953): “Distinctly a ground bird, it prefers to hide in the brush along the bank of some trickling creek, though it is often observed in dry brushy localities at some distance from water, and has been noted in such diverse places as weedy lots and about dooryards in towns, on the tide flats, among the logs and brush of a windfall, in a log jam on a river, among stranded dry logs on the sea beach, along roadsides and in partly cleared places, in tule swamps, about beaver ponds, and even in the dense Douglas fir forest. Everywhere it keeps fairly close to the ground, and in general it remains in wet or marshy places.” Frank S. Tompa (1962), in an important study of the population on Mandarte Island, B. C., found that the birds defended territory in shrubby growth, included grassland within their home ranges, and foraged in the tidal zone as a common feeding ground.

Territory: Tompa’s work on the sedentary population of Manderte Island provides the following data: Territorialism revived in late January, when singing and chasing began in males. At this time adults with previous breeding experience were occupying their territories and home ranges of the year before, substantially without change in the former boundaries. Land left vacant by mortality was occupied by birds hatched the preceding year, and a surplus of such young birds was distributed in loose groups of five to ten individuals. With the revival of territorial behavior, most young birds that were still unsettled on a site emigrated; one was found 1,300 meters from its hatching site and another 6 kilometers. Pair formation occurred in February and March.

Territory size of mated males averaged 288 square meters for 47 cases, and home range averaged 473 square meters. This average territory area is one-tenth the minimum size Nice reported for territories of euphonia in Ohio. Unmated males on Mandarte Island had territories averaging only 82 square meters. Tompa suggests that the amount of shrubbery defended determined whether a male could attract a female.

Territorial behavior gradually declined during the breeding season until it reached its minimum in late July and August when molt began in adults. Song was very rare by late June and early July, except in unmated males. Aggressive behavior revived in October. Some young of the year became territorial in late summer, and from late August until October there was an emigration of certain of these birds, apparently as a result of mutual aggressiveness and the shortage of vacant habitat on which to settle. Aggressiveness declined again after October and reached the minimum in November and December.

Tompa’s investigation establishes that the upper limit of territory size in this local population is fixed primarily by territorial behavior.

Details of early spring behavior of birds at Vancouver, B.C., are reported by William M. Hughes (1951). Males sing on bright winter mornings, and other signs of territoriality appear early in the season. Hughes remarked aggressiveness in birds in January and color-banded three, which he sexed as males. He found that these marked birds fed amicably with other song sparrows (he caught all three again on Feb. 8, together in the trap) but that each then returned to a particular area in which it attacked and pursued others of the species. This territorial behavior persisted even in a late-February and early March period of cold and snow, during the sunny intervals. Song first became frequent on March 13, courtship of females was first noted on March 17, and by March 27 the three banded males all had mates that were carrying nest material.

Song posts of males are 10 to 15 feet high (Jewett et al., 1953).

Nesting: On Mandarte Island, B.C., Tompa found that the breeding season “normally extends from the second half of March to late July.” As many as three broods are raised by some pairs.

In Washington, eggs have been found as early as April 2, and a fresh set of four eggs has been recorded on July 16. Gabrielson and Jewett (1940) set extreme dates of fresh eggs as April 15 and July 10 in Oregon. G. W. Gullion (1951) states that breeding dates of song sparrows in the southern Willamette valley, Oregon, range from Feb. 28 to Aug. 13; the subspecies is not identified but is inferred from the locality to have been morphna.

Nests are said to be very well concealed, and are located both on the ground in grass or tules or by a log, in shrubs such as blackberries, in trees such as spruces, and in brush piles. The seasonal pattern of nest location already described for melodia and many other races, in which the elevated nests are to be expected later in the season, prevails in morphna. Heights are rarely greater than 5 feet, more commonly 1 or 2 feet (Jewett, 1916; Gabrielson and Jewett, 1940). William H. Kobbe (1900) measured an elevated nest made of grasses and found its external diameter was 5 inches, its internal diameter 3 inches, its external depth 3½ inches, and internal depth 2 inches. The same author, as well as Alexander Wetmore (quoted in Jewett et al., 1953), described nest construction in terms closely resembling descriptions of nests of melodia. Hughes (1951) found a most unusual nest on April 16; it was the previous year’s nest of a Swainson’s thrush, 6½ feet high in a yew.

Eggs: The measurements of 40 eggs average 20.2 by 15.1 millimeters; the eggs showing the four extremes measure 21.8 by 15.2, 20.3 by 16.0, 18.0 by 14.2, and 18.8 by 13.8 millimeters.

Young: Song has been noted in young of the year as early as July 9, in a bird that could not have been more than 2 months old. William E. Sherwood (1929) felt that the first songs he heard were subdued in volume, but within a few minutes the songs had become indistinguishable from those of an adult. This early beginning of song correlates well with Tompa’s observation of the onset of territorial behavior after the post-juvenal molt.

Outside the breeding range, in California, morphna “Has been noted specifically in weedy thickets, along grassy ditch banks, in old cornfields, and in wooded or brush-bordered gardens. This race is not noticeably restricted to the vicinity of water in the winter season in California” (Grinnell and Miller, 1944).

Mortality: Tompa found that 29 of 55 birds (53 percent) banded as adults survived from one summer until the beginning of the following breeding season. During the next approximately 10 months, from the onset of breeding until the end of January, adult mortality was only 22 percent. Emigration of the young apparently prevented accurate determination of first~-year mortality. One year Tompa recaptured 21 of 113 young that he had banded the preceding season, 19 on Mandarte, 2 on adjacent islands. Egg and nestling losses on Mandarte Island, where nest predators and parasites were virtually lacking, were less than 40 percent.

Plumage: Ridgway (1901) describes morphna as being similar in color to the race cleonensis, “but much larger and colors more uniform above, the rusty brown or chestnut streaks on back, etc., less strongly contrasted with the rusty olive ground color and the black mesial streaks less distinct (often obsolete); under parts with the chestnut streaks on chest, etc., usually without blackish shaft-streaks, and the flanks olivaceous rather than tawny. Young, slightly rufescent bister brown above, the back streaked with blackish; beneath dull whitish or very pale buffy grayish, the chest, sides, and flanks more or less tinged with buffy or pale fulvous and streaked with sooty brownish.”

Food: L. M. Huey (1954) collected a female of this subspecies near San Diego, Calif., on Oct. 13, and states that “the entire digestive tract was found to be thoroughly stained red from a diet of the ripe [Opuntia] cactus fruit.” Tompa mentions caterpillars and lacewings as food items.

Fall: Mainland birds show a tendency to wander in the fall, as indicated by records from altitudes of 4,000 feet in the Cascade Mountains of Washington and from heights in the Olympics, as well as from records east of the Cascades. Most of these wanderers are thought to be young birds (Jewett et al., 1953), as Tompa’s work suggests.

Winter: Winter habitats are varied and include most of the places itemized by Jewett et al. (1953), as quoted in the second paragraph of this life history. Gabrielson and Jewett (1940) regard morphna as a conspicuous member of the mixed winter flocks of sparrows in Oregon. Songs are heard there on bright winter mornings. In another paper, Jewett (1916) says that birds of this race “are very plentiful on the ocean beach, feeding amongst the driftwood.

DISTRIBUTION

Range: Southwestern British Columbia to northern California.

Breeding range: The rusty song sparrow breeds from southwestern British Columbia (Alert Bay, Chilliwack) south through western Washington (Tatoosh Island, Longmire) to southwestern Oregon (North Santiam River at 3,400 feet, Grants Pass, Wedderburn).

Winter range: Winters chiefly in the breeding range, extending south to northern California (Paicines, Snelling), rarely to southem California (Riverside, Yaqul Wells) and western Nevada (Fallon).

MENDOCINO SONG SPARROW

MELOSPIZA MELODIA CLEONENSIS McGregor

Contributed by VAL NOLAN JR.

HABITS

This permanent resident of the coastal districts of the extreme southwest corner of Oregon and of three counties of northern California inhabits a variety of low dense cover, listed by Grinnell and Miller (1944) as “blackberry patches, ceanothus clumps, bracken, weeds and brush-piles in logged or burned-over land, pasture fencerow tangles, baccharis brush, willow thickets, and fresh- and saltwater marshes. Within the narrow coastal range of this race prevailing fogs and rain supply amply the moisture requirements of Song Sparrows even in cover some distance from streams or marshes. Undergrowth in forests is generally not occupied, the birds apparently seeking brush in openings and at forest edges.” Walter K. Fisher (1902), commenting on the abundance of the bird, writes that it “fairly swarms in some places, and is the commonest bird in deforested areas.”

Ridgway (1901) describes cleonensis as being “similar in size and proportions to M. m. samuelie, but averaging slightly smaller with larger legs and feet, and coloration very different, being much more rufescent; general color of upper parts deep rusty olive, conspicuously and broadly streaked on back, etc., with dark rusty brown, or chestnut, and black; streaks on chest, etc., dark rusty brown or chestnut (black medially), and sides, flanks, and under tail-coverts strongly fulvous.

DISTRIBUTION

Range: The Mendocino song sparrow is resident in the coastal district of extreme southwestern Oregon (mouth of Pistol River) and northwestern California (Del Norte, Humboldt, and western Mendocino counties, south to Gualala).

Casual record: Casual in Mann County, California (Olema).

MARIN SONG SPARROW

MELOSPIZA MELODIA GOULDII Baird

Contributed by VAL NOLAN JR.

HABITS

Except in the marshes around San Francisco Bay, the song sparrow of the long coastal district of central California is this resident race. Here it inhabits fresh water marshes, fog-drenched brush on westward-facing slopes down to the shore or to the edges of salt marshes, streamside growth such as willow clumps and shrubby weedy tangles, and garden shrubbery. As Grinnell and Miller (1944) observe, “An essential combination of dense, tangled vegetation, and moist ground or surface water is provided by each of these types of habitat.” Toward the interior the birds are limited to streamsides and freshwater marshes, but near the coast the fogs, seepage, and damp ground provide enough moisture to free them of such restrictions.

Nests are built “on the ground or in bushes a few feet up” (Ray, 1908). Richard F. Johnston, in his study of the breeding seasons and clutch sizes of western song sparrows (1954), indicates that the season extends from about February 25 to about June 25 in south central California and from about March 25 to about July 5 in the northern part of the range; the peaks of breeding are in mid-April in both sections with a second, greater, peak in the first half of May in the north. Mean clutch sizes in the sets Johnston examined were 3.71 and 3.53 in the south and the north, respectively; first clutches were somewhat smaller and second clutches larger than the means.

Grinnell and Miller (1944) say that cover “is often hunted through in almost wren-like fashion; crevices, holes and branch tangles are entered and inspected for insect food.” H. C. Bryant (1921) stated at Berkeley, “The worst egg eater yet discovered in my aviary is a Santa Cruz Song Sparrow [now M. m. gouldii].”

“In fall and winter some scattering to drier situations is noted, especially to thickets of dead grass and annuals in fairly open fields” (Grinnell and Miller, 1944).

Grinnell (1909) describes gouldii as: “Similar to M. m. cleonensis, but less rufescent, the black element much stronger on feathers of back, scapulars, and exposed quills, that of pileum taking form of streaks alternating with browns; the streaking of underparts also more decidedly black, often scarcely rufescent on edges.” The race is depicted by Dawson (1923).

DISTRIBUTION

Range: The Mann song sparrow is resident in the coastal district of central California (exclusive of tidal and brackish marshes of San Francisco Bay area), from interior Mendocino County (6 miles southwest of Laytonville), northern Sonoma County (Cazadero), and Lake County (Blue Lakes) south through San Mateo and Santa Clara counties to northern San Benito County (Paicines); east to the edge of Sacramento Valley (Stonyford, Vacaville).

SONG SPARROW: SAN FRANCISCO BAY MARSH SUBSPECIES

MELOSPIZA MELODIA (Wilson)

The following subspecies are discussed in this section: Melospiza melodia maxillaris Grinnell, M. m. samuelis (Baird), and M. m. pusillula Ridgway.

Contributed by RICHARD FOURNESS JOHNSTON

[Nowhere have song sparrows become adapted more interestingly to a specialized environment than they have in the salt and brackish marshes ringing San Francisco Bay. Here, as Joe T. Marshall, Jr. (1948) has written, “the spatial isolation of different habitats, particularly bay salt-marsh from upland fresh-water growth is correlated with a marked differentiation of very local races.” Marshall’s paper analyzes the differences in the ecologies of these sedentary marsh song sparrows, but their essential similarities have led to their being grouped for present purposes. The race samuelis is selected to represent the group because of Dr. Johnston’s knowledge of it. A few details regarding other races are inserted into the Johnston account; these are set apart in brackets: V.N.]

HABITS

Melospiza melodia samuelis is one of the many distinctive morphological segregates of this widespread species found in the central Californian region. One of the smaller, darker subspecies (Marshall, 1948), it is closely restricted to a peculiar habitat-type found only on the salt marshes fringing the northern reaches of San Francisco Bay. As these marshes are inhabited by no other subspecies, any song sparrow seen there is almost certain to be M. m. samuelis.

The salt marshes of the north part of the bay are flat expanses of alluvial soil seldom more than 7 feet in elevation above mean sea level. They are exposed to a varying amount of wetting each day

by tidal flux of the brackish bay waters. The marsh vegetation in response to these conditions is arranged in three zones, corresponding roughly to the amount of daily submergence each zone experiences. Fringing the lower edge of the marsh adjacent to the bay is a broad expanse of cordgrass (Spartina foliosa), and on the higher reaches of marsh grows pickleweed (Salicornia ambigua). Lastly, along the raised banks of tidal sloughs, which carry water into and out of the marsh, grows a gumplant (Grindelia cuneijolia). This gumplant, which reaches the size of a small bush, affords a great deal of cover when mixed with the pickleweed; here the marsh song sparrows find conditions most suitable for their existence.

The tides thus set the pattern assumed by the dominant plants on the marsh, and in so doing strongly influence the areas in which the song sparrow lives. High tides also have other effects on the birds. High spring tides, which can flood nests of song sparrows, occur in late April, May, and June in nocturnal hours; high winter tides, which may influence the amount and direction of movements of song sparrows, occur mainly in December during daylight hours.

Nesting, territory, eggs: In lowland California song sparrows tend to begin breeding activities fairly early in the year, ordinarily in late March, but the song sparrows that breed earliest are those living on salt marshes (Johnston, 1954). The first date for a completed clutch of M. m. samuelis is February 28, and the modal date for population-wide completion of clutches is March 28; corresponding dates for song sparrows living in the hills around the San Francisco Bay area are March 25 and April 15, respectively. The marked earliness of breeding in the salt marsh birds probably is an adaptation to tidal conditions, for if these song sparrows bred at the same time that the upland birds do, spring flood tides would destroy many eggs and young (Johnston, 1956a). Birds hereditarily endowed for earliness of breeding thus leave more offspring than those breeding late. [M. m. pusillula also breeds early: —V.N.]

Salt marsh song sparrows are double-brooded; almost all pairs nest twice in a season. If replacement nests are considered, each pair will nest on the average 2.5 to (rarely) 3 times each season.

The song sparrows place their nests in clumps of pickleweed, among stalks of cordgrass, or in crotches of gumplant. All nests are low to the ground, as the average height of marsh vegetation is less than 2 feet. Nests in fact average only 9½ inches above the ground; they are higher on the lower marsh, averaging there about 12 inches high. Nests are not used more than once, and the two or more nests of any one season are built at different sites within a territory.

On one occasion (out of about 130 possible occasions) a song sparrow gave a “rodent run” distraction display as it left the nest when the observer visited it.

IThe measurements of 40 eggs of samuelis average 21.5 by 15.9 millimeters; the eggs showing the four extremes measure 24.4 by 16.3, 22.9 by 17.3, 19.1 by 15.8 and 20.1 by 15.0 milhimeters: W.G.F.H.I Mean clutch-size for salt marsh song sparrows is 3.20±0.05 eggs, on the basis of 157 records collected between 1950 and 1955 (Johnston, 1956a). The range in mean clutch-size is from 2.91 to 3.42 eggs. In any one season the earliest clutches are relatively small (to 2.83 eggs), clutches completed in mid-season are relatively high (to 3.66 eggs), and those of the late season low (to 2.60 eggs). Pairs of samuelis averaged from 7.5 eggs to 9.1 eggs per season from 1950 to 1955. Owing to unpredictable mortality to eggs and young, these figures are not a reliable guide to productivity, which is best defined as the number of fledglings per pair of adults per season. On the salt marsh this productivity is from three to five fledglings per pair per year.

[M. m. pusillula also has a low mean clutch size, 3.3 eggs: V.N.]

Territorial relationships of salt marsh song sparrows are similar to those of the species in general. In late winter and spring, male singing and chasing increase. By late February and March most males have established themselves on territories; pair-formation ordinarily has already taken place, but those birds not yet mated now form pairs. Singing decreases in late March and April, but territorial strife continues sporadically throughout the remainder of the breeding season. Territorial activity ceases in July, but becomes evident again in late August and September. Although in many instances true territories are not staked out the quarters most birds, adults and juveniles alike, occupy in September are identical with those of the following breeding season.

The territories of salt marsh song sparrows are small, and the borders of some sloughs may support as many as 8 to 10 pairs of birds per acre; most territories are thus about 30 feet wide by 150 feet long, a little over 0.1 acre. Territories are larger where vegetation is sparser at the heeds and mouths of the sloughs than along the middle reaches.

Adults show a strong tendency to remain permanently in the territories they take up their first autumn. Very few birds move more than 10 meters from this first autumn territory, no matter how long they live. The longest movement recorded on banded birds (Johnston, 1956a) is 35 meters.

The incubation period is about 12 (12 to 14) days. Young remain in the nest 9 to 12 days, but will leave prematurely at 8 days if disturbed. Fledglings remain under parental care for 5 to 8 more days; this gives a total parental period of attention per brood of about 30 days. The adults start second broods in short order, and the first young then gradually move out into other parts of the marsh.

Dispersal by the young occurs from time of fledging until autumn, when the adults again begin to show territoriality. The median distance of dispersal in these young is 185 meters. The longest distance on record is 960 meters (Johnston, 1956a).

Enemies: Adults are preyed upon by marsh hawks, short-eared owls, and probably Norway rats. Much predation must occur for which there is little record, for although adult mortality is about 42 percent per year (Johnston, 1956) very few birds are known to have died through a specific agency. Only four skulls of song sparrows appeared in 491 pellets of short-eared owls collected in a 4-year period (Johnston, 1956b).

Eggs and nestlings are subject to higher mortality than full-grown birds, and for these the mortality factors are easier to determine. In a sample of 504 eggs and young, about 20 percent of eggs and young were killed by some predatory agent (probably Norway rats), 11 percent by flood tides, 5 percent by desertion of adults, 5 percent by rainstorms, and about 4 percent by accidental loss of eggs from the nest and parasitism by cowbirds (Molothrus ater). Total mortality was 50 percent of all eggs laid.

Plumages: The sequence of plumages in salt marsh song sparrows has not been studied, but probably is the same as for other subspecies of the species. The annual molt begins in early July, almost immediately following cessation of breeding, at which time adults are in extremely poor feather owing to heavy abrasion on rough marsh plants. The annual molt is completed in late August or early September, and at this time most juveniles have completed the post-juvenal molt.

[Grinnell (1909) described maxillaris as resembling the race “gouldii closely in coloration and M.m. heermanni in general size; differs from M.m. samuelis * * * in having the browns more extended and of a deeper tone (bay rather than hazel) and in much greater size and, especially, bulkier bill; differs from M.m. gouldii * * * in much greater size throughout; and from M.m. heermanni * * * in that the base of the maxilla is more swollen, the black streakings everywhere broader, and the general tone of coloration darker.” Ridgway (1901) states that samuelis is “exactly like” the race heermanni in coloration, but is “much smaller, with the bill more slender.” He describes pusillula as being “Most like samuelis but still smaller, the wings and tail decidedly so; coloration much less rusty, the general color of upper parts olive-grayish, the black dorsal streaks not distinctly, if at all, margined with rusty brown, the lateral crown-stripes and wings less distinctly rufescent, under parts more heavily streaked (streaks usually wholly black) and flanks paler fulvous; under parts usually more or less tinged with yellowish. Young much paler and grayer than that of N. c. samuelis, with the broad black streaks on back and scapulars much more strongly contrasted with the ground color; ground color of under parts dull yellowish white or pale yellowish buff, without brownish tinge on breast or sides.” — V.N.]

Behavior: foraging — Salt marsh song sparrows forage on the marshes in the same fashion as do song sparrows in upland situations. Upland races frequently scratch vigorously with their strong feet at the ground surface to expose invertebrate and plant foods within and under the surface litter. Salt marsh song sparrows perpetuate this foraging mannerism; these birds scratch, typically with both feet in unison, on the soft slough mud, and peck frequently at the disturbed surface. One bird, for a period of 40 seconds, alternately scratched 3 to 6 seconds and pecked 1 to 5 times between each scratching period. The bird was hunting small snails, and it furrowed and turned over the top 1/32 inch of mud covering about 8 square inches. Foraging on harder mud comprising the true marsh surface most closely resembles the scratching performed by song sparrows in upland habitats. The birds also forage in “typical” fashion, i.e., slowly progressing by short hops accompanied by wing flicks and tail flicks and punctuated by frequent pecks at possible food items.

Another type of foraging on soft mud could be called “terrestrial flycatching.” In midsummer on salt marshes large numbers of various Diptera occur. Some of these frequent exposed mud at low tide, flying just above the mud or resting on it. Song sparrows catch these flies, by making short, jerky hops or runs with tail elevated and sometimes with wings half outstretched, but without leaving the ground. Aerial flycatching is also a small part of foraging behavior in these birds, but the half-outstretched wings are probably used to maintain balance rather than to remain ready to resume flight.

Salt marsh song sparrows feed at dried heads of the gumplant (a composite) and eat the peeled seeds, perching on the edges of the larger heads or on nearby stalks or heads. They look much like siskins or goldfinches when foraging in this fashion.

In autumn these song sparrows eat fleshy fruits and seeds of pickleweed or salicornia. The birds perch quietly in a patch of salicornia and slowly and methodically chisel the fruits out of the succulent, cylindrical spikes with their bills. They seem to prefer plants within 6 feet of the tidal sloughs, and they consume most of the available fruits along the sloughs in 2 or 3 weeks, ordinarily at the end of November. This feeding is possibly of some importance as a source of free water; it comes at a time when insect foods with their free water are at an ebb.

The birds also eat the almost unbelievably enormous numbers of seeds of salicornia released when the spikes wither in winter. The birds probably take these seeds whenever they find them, but their consumption is especially noticeable after the high winter tides when the loose seeds have gathered in great, floating windrows up to a foot thick and form a concentrated food source. Song sparrows spend almost all the morning hours foraging, mostly on the masses of salicornia seeds; they also eat invertebrate animals they find within the seed masses.

Reactions to high tides. — It is possible to mention only the diurnal behavior evident during the daytime high tides in December; presumably equally important behavior patterns occur when breeding adults and young contend with the night high tides in spring, but they have not been witnessed. It is known that nestling song sparrows about 8 days old escape flooding by climbing up into vegetation above their nests.

The tallest salt marsh vegetation is leafless in December. When tide water rises within a foot to a few inches from the tops of these plants, little cover remains for song sparrows. When no potential avian predators are in evidence, the sparrows usually maintain an active, vigorous foraging pattern. Occasional birds splash in the water in typical bathing routines, and others loaf. A few birds apparently wander beyond their autumnal territories, and occasional examples of territorial strife are evident. Such occurrences end in a chase, and the intruder moves back to his presumed point of origin. One banded bird was seen under such circumstances to have moved about 150 yards, and thus made a round trip of 300 yards.

I reviewed the influence of winter high tides on two distinct populations of salt marsh song sparrows before completing my observations on the behavior of M. m. samuelis. In a marsh where extensive, manmade levees existed, the song sparrows accumulated on them, but on a marsh where only emergent vegetation and floating timbers were available, no such concentrations were seen. Later observations on this marsh from a 6- by 30-foot raft floating at high tide showed that the song sparrows there did indeed congregate; at one time immediately after a marsh hawk passed, 17 song sparrows were perched on the raft. The birds apparently felt exposed in the thin emergent vegetation, and those from as far as 60 yards away streaked toward the raft in full, powered flight. After the hawk passed the birds gradually left the raft, but each time a hawk approached they returned to it. At the same time, the foraging or loafing song sparrows practically ignored short-eared owls; this difference in reaction to the raptors lends support to the idea that short-eared owls are unimportant predators on song sparrows.

The sparrows reacted to marsh hawks just as strongly during normal tides. They always detected an oncoming hawk well before the human observer did and, giving thin, chip alarm notes, moved down to soil level, quite out of sight and reach of any hawk.

DISTRIBUTION

Suisun Song Sparrow (M. m. maxillaris)

Range: The Suisun song sparrow is resident in brackish marshes surrounding Suisun Bay in central California (Southampton Bay, Grizzly Island, Port Costa, Pittsburg).

Casual record: Casual in Santa Clara County, California (Palo Alto).

Samuel’s Song Sparrow (M. m. samuelis)

Range: The Samuel’s song sparrow is resident in central California in salt marshes on the northern side of San Francisco and San Pablo bays (Richardson Bay to Vallejo) and on the south side of San Pablo Bay (southwest to San Pablo Point).

Egg dates: San Pablo Salt Marsh, Contra Costa County, California: 157 records, February 28 to June 18; 78 records, March20 to April 10.

Alameda Song Sparrow (M. m. pusillula)

Range: The Alameda song sparrow is resident in salt marshes surrounding the south arm of San Francisco Bay, California (San Francisco, Alviso, Stege).

MODESTO SONG SPARROW

MELOSPIZA MELODIA MAILLIARDI Grinnell

Contributed by VAL NOLAN JR.

HABITS

This non-migratory race is confined to the central lower basin of the Great Valley of California, the lowland between the Coast Range and the Sierra Nevada Mountains. Grinnell and Miller (1944) state that it breeds “chiefly, perhaps entirely, below 200 feet elevation” but may occur rarely at greater altitudes along streams of the Sierran foothills. The same authors state that the habitat is “Freshwater marshes and riparian thickets. Predominant plant cover consists of willow and nettle thickets and growths of tules and cattails.” No other material on the life history has been found.

J. Grinnell (191 ib) described the race mailliardi, giving as diagnostic characters the generally large size, large bill, and broad and dark markings; “resembles Melospiza melodia maxillaris closely in these respects, but shape of the bill different, more nearly like that in M. m. heermanni.” He considered the bill the distinctive feature of maihiardi. Although the bills of mailliardi and maxillaris appear practically identical when viewed from the side, “when viewed dorsally the bill of maihiardi presents a very much narrower outline, there being scarcely any indication of the lateral swellings of the maxilla characterizing the bill of maxillaris * * “. In coloration mailliardi is very much darker than heermanni, having the streaking everywhere broad and black, with edgings of deep bay, the latter color showing dorsally to the almost entire exclusion of ashy marginings, there being mere traces of the latter. In coloration, mailhiardi differs from maxillaris only in being a trifle less heavily marked on an average * *

DISTRIBUTION

Range: The Modesto song sparrow is resident in the Central Valley of California, from Glenn and Butte counties (Glenn, Biggs) south to Stanislaus County (Modesto, Lagrange); west to the deltas of the Sacramento and San Joaquin rivers.

Casual record: Casual in western Nevada (Fallon).

HEERMANN’S SONG SPARROW

MELOSPIZA MELODIA HEERMANNI Baird

Contributed by VAL NOLAN JR.

HABITS

Still another non-migratory song sparrow from California, this race lives between the north-south mountain ranges in the San Joaquin valley. In this arid region the bird adheres closely to stream-, lake-, and marsh-side vegetation at altitudes of from 100 feet to 5,000 feet. In 1944 Grinnell and Miller reported that “numbers have greatly increased in the last thirty-five years owing to development of irrigation systems in previously unoccupied parts of its general range.” In an earlier paper (1911) Grinnell had noted the absence of song sparrows in great stretches of dry prairies in the San Joaquin valley and predicted range extensions along canals.

The breeding habits of heermanni appear to fit the generalized pattern for riparian song sparrows. Grinnell (191 la) describes two nests, at 2½ and 4 feet, built in low vegetation in which drift trash had lodged. He also reports finding two females with nests, but only one male, at a reservoir 3 miles distant from any other song sparrow habitat. Among other possibilities was that of polygyny attributable to the surplus of females.

The measurements of 20 eggs average 20.4 by 15.4 millimeters; the eggs showing the 4 extremes measure 23.1 by 17.2 and 19.6 by 12.8 millimeters.

The food habits of heermanni are among those Beal (1910) investigated and reported in the life history of M. m. samuelis and the other salt marsh song sparrows.

Ridgway (1901) describes heermanni as being similar to melodia “but smaller and coloration much darker and browner, the black streaks on back, etc., averaging broader, and streaks on chest, etc., darker (black or brownish black in summer); young similar to that of * * * melodia but deeper tawny brown above with black streaks on back broader, the under parts more or less tinged with brownish buff, especially on chest, where the dusky streaks are broader.” Grinnell (1911a) states that heermanni has a “much paler ‘ground color’ * * ‘i’, narrower black-streaking both above and below and * * * slightly smaller bill” than mailliardi.

DISTRIBUTION

Range: Heermann’s song sparrow is resident in the southern San Joaquin Valley of California, from Merced and Mariposa counties (Los Bafios, Yosemite Valley) south to Kern County (Fort Tejon, Walker Basin); east to Kings Canyon (Zumwalt Meadow).

SAN DIEGO SONG SPARROW

MELOSPIZA MELODIA COOPERI Ridgway

Contriluted by VAL NOLAN JR.

HABITS
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Melospiza melodia cooperi is a resident race in southern California and northern Baja California in the valleys of the coast ranges and on the Pacific slopes; it extends eastward to streams penetrating the Mojave and Colorado deserts. Its habitat is river bottom vegetation, fresh water marshes, “at least margins of salt marshes” (Grinnell and Miller, 1944), and garden shrubbery. Settlement of the coastal plain, with a consequent development of water systems, has contributed to an increase in numbers in relatively recent times. Breeding occurs from sea level to altitudes of 5,000 feet, and vagrants have been collected in late summer at 7,500 feet. Nests found in pampas grass (Myers, 1910) and 10 inches high in a dock plant (Chambers, 1917) have been described.

R. F. Johnston (1954) has reported that the breeding season begins about Feb. 5 and lasts until about July 5, with the great majority of nesting records falling between late March and early June, the peak about May 1. First clutches averaged 3.74 eggs, second clutches 3.58; the seasonal mean was 3.69 eggs. The interval between completion of the first nest and the laying of the first egg has been reported for one nest as about 5 days (Lamb, 1922). W. C. Hanna (1924) weighed 48 eggs; the extremes were 2.87 and 2.05 grams, the average 2.41 grams.

A number of diverse but interesting points have been recorded about cooperi. Josephine R. Michener (1926) describes her efforts to raise a young bird unable to fly. Among details of its behavior is the fact that its response to the first standing water it saw was to hop into the dish and bathe, at an age inferred to be about 20 days. E. L. Sumner, Jr. and J. L. Cobb (1928) in the fall and winter of 1927 displaced 25 banded birds about 4 miles and recaptured 3 at the original trapping stations within a few days; 8 birds were released 34 miles west of the point of capture, to which none was known to have returned. More interestingly, 2 were still present at the point of release 7 days later.

J. Mailliard (1919a) observed that a bird that habitually flew against a window pane on cool mornings was in fact gathering sluggish house flies apparently attracted by the warmth of the glass on the preceding evenings. R. S. Woods (1932) saw song sparrows of this race drinking sugar solution put out for hummingbirds. W. L. Dawson (1923) picked three ticks from the head of a cooperi song sparrow in April 1917, but gives no further details.

Ridgway (1901) describes cooperi as being similar to heermanni “but slightly smaller and coloration much lighter and grayer; prevailing color of the back, etc., grayish olive, the back broadly streaked with black, the black streaks with little, if any, rusty external suffusion; young similar to that of M. m. montana.”

DISTRIBUTION

Range: The San Diego song sparrow is resident in the coastal districts of southern California from Santa Cruz County (Santa Cruz) southward to northern Baja California, as far as lat. 30° N. (San Fernando); east to streams penetrating Mohave and Colorado deserts (Manix, Palm Canyon, Vallecitos, east base of Sierra San Pedro Mártir).

SONG SPARROW: PACIFIC INSULAR SUBSPECIES

MELOSPIZA MELODIA (Wilson)

The following subspecies are discussed in this section: Melospiza melodia micronyx Grinnell, M. m. clementae Townsend, M. m. graminea Townsend, and M. m. coronatorum Grinnell and Daggett.

Contributed by VAL NOLAN JR.

HABITS

On each of the San Miguel, San Clemente, Santa Barbara and Coronados Island groups off the California and Baja California coasts lives a sedentary subspecies of the song sparrow. These four races, about whose life histories not a great deal is known, are treated together here.

Although the habitats these islands afford are not uniform, m general the birds dwell in a sparsely vegetated environment characterized by its aridity. Cover is afforded by herbs and coarse grasses, cacti, and, particularly on San Clemente Island, brush and shrubs. On Los Coronados Islands no fresh water is to be found (Grinnell and Daggett, 1903), and fresh moisture must come only from the condensation of fog. Indeed fog seems to supply most of the moisture needs for all these races (Grinnell and Miller, 1944).

C. B. Linton (1908) writes that M. m. clementae was abundant on San Clemente. He continues: “Common in the yards at Howland’s, nesting in the scrub cacti and vines within a few yards of the hacienda. March 31, three nests were found in the corral near the stables; one contained four young one week old, the others having incomplete sets. These nests were built a few inches from the ground in the center of the cacti beds, which, being covered with a thick growth of vines, completely hid the nests.”

Wright and Snyder (1913) report M. m. graminea occurs on Santa Barbara wherever bushes provide cover, and Grinnell and Miller (1944) add that “Bushes are used for nesting.”

Grinnell and Daggett (1903) visited Los Coronados Islands on Aug. 6 and 7,1902, and collected the type specimen of coronatorum. The following quotations are from their account of their field work:

Juvenals were seen along the path * * *. An old and weather-beaten nest was found under a bush. * * * As we landed, an individual was fearlessly hopping close at hand among the boulders almost at the edge of the surf. Most of the Song Sparrows, however, were seen higher up toward the crest of the island, where they were haunting the sparse growth of shrubs on the shaded northeast slope. We saw no trace of fresh water anywhere, and the scanty vegetation presented anything but an inviting appearance. Yet here we heard the familiar notes and full song of these birds which on the mainland keep so close to verdant water courses and damp lowlands. The Rock Wren, always a bird of the dryest localities, did not seem out of place, but the Song Sparrow seemed altogether foreign to such surroundings.

Nelson K. Carpenter (1918) described and photographed two nests of coronatorum, one of grasses placed 3 feet up in a bush, the second on the ground and “constructed entirely of feathers and the skin of a lizard * * *ï This nest was found in the midst of the large colony of California Brown Pelicans and Western Gulls.” Pingree I. Osburn (1909) found three young just able to fly on April 8.

Joseph Grinnell (1928) describes micronyx as the race with grayest coloration among song sparrows in general: “brown or brownish tones almost wanting; dark markings black and sharply contrasted against gray of dorsal surface or white of lower surface; bill, feet and especially claws weak; wing showing extreme of bluntness (longest primary not much longer than outermost).” M. m. micron yr is most nearly like graminea of Santa Barbara Island, “but differs from it in broader and blacker dark streaking everywhere, in grayer ground-color dorsally, especially on the pileum, in paler flanks, in decidedly smaller claws, in blunter wing, and in slightly greater size.” From clementae the race micronyx differs as it does from graminea, “only for the most part (save as to general size) in greater degree. Especially on the top of the head is the greater amount of grayness apparent; the broad brown capital side-stripes in clementae are in micron yr reduced to very narrow ones, which play out altogether on the nape instead of extending back to blend (in clemerttae) with the brownish tone of the dorsum. The broad gray occipital area is in micronyx lined sharply with black shaft streaks which are thus thrown into conspicuous contrast.”

Ridgway (1901) describes clementae as being similar to cooperi “but slightly larger and coloration still grayer, the back light olive-grayish, with black streaks narrower, the black streaks of chest, etc., also narrower; young similar to that of cooperi but paler above.”

Ridgway (1901) describes graminca as being similar in coloration to clementae, but much smaller.

J. Grinnell and F. S. Daggett (1903) describe coronatorum as “most nearly resembling in coloration melospiza clementae, and general size about the same, but tarsus decidely shorter and bill smaller; differs from * * * cooperi of the adjacent mainland in much paler ground color, narrower streaking and smaller bill.”

DISTRIBUTION

San Miguel Song Sparrow (M. m. micronyx)

Range: The San Miguel song sparrow is resident on San Miguel Island, Santa Barbara County, California.

San Clemente Song Sparrow (M. m. clementae)

Range: The San Clemente song sparrow is resident on Santa Rosa, Santa Cruz, Anacapa, and San Clemente islands off the coast of southern California.

Casual record : C asual on the California mainland (Santa Barbara).

Santa Barbara Song Sparrow (M. m. graminea)

Range: The Santa Barbara song sparrow is resident on Santa Barbara Island, Los Angeles County, California.

Coronados Song Sparrow (M. m. coronatorum)

Range: The Coronados song sparrow is resident on the four islands of Los Coronados group off northern Baja California.

TUCSON SONG SPARROW

MELOSPIZA MELODIA FALLAX (Baird)

Contributed by ROBERT WILLIAM DICKERMAN

HABITS

The pale reddish desert song sparrow race, fallax, is a resident of riparian and marsh associations at low to moderate elevations from extreme southeastern Nevada and extreme southwestern Utah southwards to central Sonora. In the period before the introduction of cattle, the rivers of central Arizona had lush bottomlands that supported beaver marshes and their attendant wildlife. Trapping by man, overgrazing by cattle, and, most recently, the demand for irrigation water have reduced the major portions of these rivers to dry eroded beds. As a result, some populations of fallax are extinct (those of the Santa Cruz River and of sections of the San Pedro and Salt Rivers) and others much reduced. The largest populations are now found along the Salt River above Tempe at Coon Bluff and in marshes along the Gila River near Palo Verde, both locations in Maricopa County.

Allan R. Phillips (1943) described a subspecies bendirei from the Salt River at Tempe Butte, Maricopa County, Ariz., as a population intermediate between fallax and saltonis. Later, when it was realized that saltonis migrates regularly to southeastern Arizona, the type of bendirei was reexamined and proved to be a fresh-plumaged migrant of saltonis. Examination of a large series of fresh-plumaged birds from Arizona and Sonora has revealed no positive geographic variation within the population here considered fallax.

J. T. Marshall, Jr., and W. H. Behle (1942) describe the habitat of song sparrows in the Virgin River valley as “the vicinity of cattail swamps with standing water and with brushy thickets such as mesquite or rose in the immediate dry-land surroundings. Both the thickets and the cattails are frequented by the Song Sparrows, the latter being resorted to especially for greater protection * * *. Bird associates are Marsh Wrens, Yellow-throats, Yellow Warblers, and Red-winged Blackbirds * * *”. The swamps where these birds do occur are few in number and widely scattered. If the swamps are of any extent at all, Song Sparrows are numerous in them, but the total * * * population cannot be great. This spotty distribution of suitable swampy habitat * * * makes for discontinuous distribution of the colonies within the general range of the race.”

Nesting: The nesting habits of fallax are similar to those of other populations in more temperate areas. The nest is always built close to open water. Francis C. Willard (1912) found a nest in a low bush. The same author (1923) writes “The Song Sparrow sometimes deserts the ground and low bushes in favor of a tree, and the desert subspecies (Melospiza melodia fallax) also has this trait. One nest was built fifteen feet up in a large willow tree, on a horizontal branch. The bird was on the nest when I found it and remained until I was nearly up to it. * * * There were four of the song sparrow’s eggs and four of the Long-tailed Chat * *

Herbert Brandt (1951) describes the site of a nest he found on the Slaughter (San Bernadino) Ranch 20 miles east of Douglas, Cochise County, Ariz., in an “irrigation project near cattail pond in grassy meadow; surrounding flora, small mesquites, sacaton and other grasses, beyond which are rows of black willow and cottonwood, and in near-by pond, cattails and other water plants; date, May 23, 1948. Nest situated 18 inches up in a low dense mesquite brush, 25 feet from artesian pond rim; nest placed in a four-pronged fork of dead wood; made in a crude, bulky manner of coarse plant stems and leaves; rim substantial but ragged and irregular, shaped to fit the supports; lining of brown rootlets and white cow-tail hair arranged circularly; interior well cupped. Apparently the only pair present in the area. Contents, 3 eggs, unevenly incubated 6 to 8 days.” A female taken March 26 near La Casita (27 miles south of Nogales, Sonora) contained ova measuring 7 and 4 millimeters. Full grown young have been collected by May 30 (Feldman, San Pedro River).

Eggs: There are 15 sets of eggs of fallax in the United States National Museum and one set in the University of Arizona collection. These sets contain 2, 3, and 4 eggs, with 3 being the most frequent number and the average. These eggs measure in millimeters 14.1 to 15.7 by 17.4 to 21.7. The average of the 48 eggs is 15.00 by 19.03.

Plumages: The molts and plumages of the desert populations are similar to those of the species elsewhere, save for the light coloration of the plumage. Adults are in heavy molt in September, and first molt may continue until December, but the time of instigation of these molts is not known. Young in juvenal plumages are to be found in September and possibly later. Wear and fading are extreme in the desert subspecies. This is probably due to coarse vegetation, sun, and possibly higher alkaline content of the soils. Adults taken after March are virtually worthless for taxonomic purposes.

Ridgway (1901) describes fallax as being similar to montana in the slender bill, “but wing and tail averaging decidedly shorter and coloration conspicuously paler and more rusty, the rusty streaks, both above and below, without blackish shaft-streaks, or else with these merely indicated on the interscapular region; young dull brownish buffy or pale wood brown above, the back streaked with rusty brown or dark brown; beneath buffy white, the chest streaked with rather light brown.”

Winter: The Tucson song sparrow winters throughout much of its breeding range, but some movement does occur, especially from the northern part of the breeding range. The breeding population at Tucson has been extinct for more than 5 decades, but an occasional winter specimen is taken there, and a winter specimen of fallax has been taken at Bard, Calif.

DISTRIBUTION

Range: Southeastern Nevada and southwestern Utah south to Arizona and northeastern Sonora.

Breeding range: The Tucson song sparrow breeds from southeastern Nevada (Pahranagat Valley) and southwestern Utah (St. George) south in the Virgin River Valley and the Colorado River Canyon of Nevada and north-central Arizona, and locally through the lowlands of central and southeastern Arizona to northeastern Sonora. It intergrades with saltonis along the Colorado River from southern Nevada to Topock, Arizona, along the Big Sandy River, and a little south of Arlington along the Gila River. In central Arizona it nests below the Mogollon Plateau (Indian Gardens, Oak Creek Canyon, and possibly formerly Walnut Creek north of Prescott), along the Verde and Salt Rivers, the Gila River (Geronimo, San Carlos, Safford, Phoenix, Palo Verde), the San Pedro River (near Feldman, formerly south to Fairbank), along the Sante Cruz River (formerly at Tucson), and at Picacho Reservoir. In extreme southern Arizona it has nested at Patagonia and San Bernardino. In Sonora it breeds along the Rio Magdalena (Caborca) and along its tributaries (Rancho la Arizona, Saric, Magdalene, La Casite, and Agua Caliente), along the Rio Sonora (Hermpsillo, Ures, Arizpe), the Rio Moctezuin (Oposura, Cumpes), and the Rio Bevispe (Husebes, Grenados, and Pilares).

Winter range: Winters apparently over most of the breed range, but in reduced numbers northward.

DESERT SONG SPARROW

MELOSPIZA MELODIA SALTONIS Grinnell

Contributed by VAL NOLAN JR.

HABITS

Melospiza melodia saltonis is a resident of the lower Colorado River valley. The summary of its habitat in California by Grinnell and Miller (1944) gives an idea of its life: “Riparian plant associations, most notably those dominated by arrow-weed (Pluchea), guatemote (Baccharis) and young willows, and tule beds and cattails in marshes, overflow sumps and along irrigation systems. Nests are placed in the vegetation above the mud which marks flood level. Development of irrigation has undoubtedly increased the total population of this race in the last 30 years. Although usually sharply limited to water-seeking plants, and most abundant in cover growing over or at the edge of water, this Song Sparrow has occasionally been noted in mesquite thickets at some distance from water.”

The measurements of 40 eggs average 18.9 by 14.9 millimeters; the eggs showing the four extremes measure 20.8 by 15.2, 18.5 by 15.5, 17.8 by 15.0, and 18.0 by 14.0 millimeters. Robert W. Dickerman reports three sets of eggs of this race from Yuma, Ariz., in the collection of the University of Arizona, two of which contain eggs of the brown-headed cowbird.

J. Grinnell (1909) described saltonis as resembling fallax, “but very much paler throughout, the ‘ground-color’ being white ventrally and ashy dorsally, with streakings of pale hazel; supercilliary stripe wholly white; general size much less than in either M. m. fallax or M. m. montana.”

DISTRIBUTION

Range: The desert song sparrow is resident in the lower Colorado Valley in extreme southern Nevada (east of Searchlight), southeastern California, western Arizona (east to Big Sandy River at 2,000 feet; Alamo), northwestern Baja California (Mexicali, mouth of Hardy River), and northwestern Sonora (Colorado River delta), extending northwest through the Imperial Valley of California (Mecca, Calexico).

Casual records: Casual in the desert area of southeastern California (Death Valley, Oro Grande), south-central Arizona (Tucson), and northwestern Sonora (Sonoyta River, Caborca).

BROWN’S SONG SPARROW

MELOSPIZA MELODIA RIVULARIS Bryant

Contributed by VAL NOLAN JR.

HABITS

This race inhabits “west-flowing stream courses” (Grinnell, 1928) in a very limited region of central Baja California. Here, as Griffing Bancroft (1930) writes, “there are several systems of dry river beds which have an important influence on the biology of this region. Even though the country be arid beyond anything known in the United States there is still enough rainfall to provide some moisture. This water, as well as a part of that from the cloud-bursts that come once in a decade, finds its way to the sea by means of a subterranean flow. * * * Occasionally the subterranean flow encounters bed rock formations which force the water to the surface.”

G. Bancroft (1930) gives the following description of the breeding biology of rivularis in this restricted range:

This light-breasted type of Melospiza occurs wherever there are poois of water with tule or willow. * * * It is by no means as abundant * * * as is, for instance, M. m. cooperi in the willow bottoms of southern California. Still it can hardly be regarded as rare.

The birds begin to lay the latter part of April and continue for at least six weeks. Most of the nests are bulky affairs of tule, usually lined with palm fibre or cow hair. They are normally placed in tule about four feet above the water. * * * [Some were] surrounded by loose and fairly long dead tule leaves. These more than equaled the bulk of the nest * * * . Some of the nests found were in willow trees and one was in a thick weed clinging to a rock and overhanging a pool. In general, excepting the unusual size of their nests, the habits of the Brown Song Sparrow wore much like those of the San Diegan form [cooperi]. They lay either two or three eggs, with four the record.

The eggs themselves are unlike those of any other Song Sparrow I have examined. They are more brilliant than those of the northern birds, a brighter blue, and altogether lacking in the common tan type, in which the spots are so close together as to give the appearance of a reddish egg.

G. Bancroft’s measurements of 35 eggs from San Ignacio yielded an average size of 21.2 x 15.9 mm., and of 250 eggs from El Rosario an average size of 20.1 x 15.2 mm.

Ridgway (1901) describes rivularis as being similar to fallax “but larger, with longer, more slender, and more compressed bill, still less strongly contrasted markings, and duller, less rufescent colors.”

DISTRIBUTION

Range: The Brown’s song sparrow is resident in south-central Baja California (Santa Agueda Canyon and San Ignacio south to Comondti).

Casual record: Casual in the Cape District of Baja California (Todos Santos).