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

The tremendous energy which drives small birds toward their breeding grounds in spring is well illustrated by the way in which some of them overshoot their destination. Witness the occurrence of the eastern race of the white-crowned sparrow on Fletcher’s Ice Island (T—3) on June 16, 1957, when it was at 82°37′ N., 99°50′ W., only 150 miles from the North Pole and some 2,000 miles beyond the nearest nesting habitat of the species, as reported by Spencer Apollonio (1958).

Because unusual observational opportunities are required to recognize such events, we are less aware of them than the facts probably warrant. But it is this sort of random exploration of extralimital territory that helps account for the ability of so many species to colonize new territory as soon as it becomes suitable. We know, today, that the margin of the tundra which forms the northern limit of the white-crown’s range has been both farther north and farther south during the last 10,000 years, and that the range and the population of this species have varied accordingly.

The eastern white-crown is best described as a subarctic and alpine zone bird, but so extensive is its range that only ecological characterization is really helpful. This has seldom been done and as a result “life zone” pigeon-holings are rampant with apparent contradictions. The only full awareness of the ecology of this bird in all the material before me is Harrison F. Lewis’ excellent description, in a letter of July 1, 1963: “In coasting along the north shore of the Gulf of St. Lawrence from west to east, in latitude slightly north of 50°, one enters the breeding range of the white-crowned sparrow abruptly on passing Saint Genevieve Island, the easternmost of the Mingan Archipelago. The reason for the abruptness of this boundary is that the Mingan Islands, which border the coast for some 50 miles, are formed of limestone, as is, also, much of the adjacent coastal mainland. Eastward from Saint Genevieve, the mainland and the coastal islands are of acidic pre-Cambrian rocks, such as granite and gneiss. The vegetation of the limestone belt is much superior to that which grows on the adjacent pre-Cambrian rocks, and the bird life reflects this; for example, the ovenbird nests on the limestone of the Mingan region, but not beyond. West of Saint Genevieve I have only a few records of sporadic occurrence of the white-crown in the nesting season.”

This Mingan region is an outlier of Paleozoic rocks. In his monumental photoreconnaisance of the vegetation and physiography of the Labrador-Ungava peninsula, F. Kenneth Hare (1959) marks it as the eastern terminus of the Laurentide massif and considers it an outlier of the coastal tundra that fringes the southeast coast of Labrador. Climatologists bring the 55° F. July isotherm, which transects the peninsula from west to east, to the coast just east of this region, and forms a rough boundary between the open subarctic woodland to the north, the closed-crown forest to the south and west, and the scrubby forest characteristic of the eastern tip of the peninsula at this latitude.

It is open, stunted tree growth and brush that attracts nesting white-crowned sparrows. At Goose Bay, Labrador, for example, I found them nesting only in the open, often burned, black spruce and dwarf birch on the high, sandy delta of the airport plateau. In better sites nearby, white-throated sparrows nested. An awareness of such temporary ecological changes in distribution allows me to view with interest the nesting reports of white-crowns at North Bay, Ontario, and at Bale Comeau and Godbout on the north shore of the St. Lawrence, as well as near Godthaab, Greenland, in 1824 (Salomonson, 1950); though I cannot credit them for lack of corroborative details on the ecology of the site.

Despite the fact that it was the first known and is the most widely distributed race, the eastern white-crown has been one of the least studied. No one has yet reported in any detail on the biology of the alpine populations of the Rocky Mountain massif, and the eastern group has only recently begun to attract the attention it deserves. In a study of geographic variation in the entire leucophrys group, Richard C. Banks (1964) reduces Harry C. Oberholser’s (1932) Cordilleran race oriantha to synonymy with the eastern race, discounts W. E. Clyde Todd’s (1953) proposal of the name nigrilora for the “ultratypical” Labrador peninsula population, and considers the slightly larger, reddish-backed and black-bred nominate leucophrys to be the ancestral population. “The species,” he writes, “appears to be essentially a northern one which has extended southward only where summer conditions approximate those found in subarctic regions. Thus, in the western * * * United States, White-crowns are found only in high mountains and along the cool Pacific coast.”

Spring: The interaction between internal and external factors in the eastern population remains almost unreported. Only Marshall B. Eyster (1954) has shown that, compared to such congeners as the white-throated sparrow and the junco, the white-crown is much more prone to pre-migratory nocturnal unrest (zugunruhe).

Viewed from the central wintering grounds of the southern Great Plains, the spring migration involves a radiation northward. One population segment goes northwestward into the Rocky Mountain uplands, another more or less due north to the Cypress Hills of Saskatchewan and the Hudson’s Bay country, but the largest segment trends far to the east of north, to summer in northern Quebec and western Newfoundland.

In an analysis of 198 recoveries and 9,107 returns from nearly 232,000 white-crowns banded between 1920 and 1963, Angelo J. Cortopassi and Richard L. Mewaldt (1965) show that migration is of a broad-front type and not oriented to landmarks: of 6,000 birds banded while on migration, not a single individual returned to the place of banding. Migration is by hops of at least 200 miles, with one 310-mile hop recorded in spring, and with a daily mean of about 50 miles. One bird banded by Ralph K. Bell at Clarksville, Pa., May 6, 1962, was recaptured 1,500 miles to the north at Battle Harbor, Labrador, on June 12, having averaged 40 miles per day.

The rapid northward surge of this species results in “a high fidelity of timing” as Cortopassi and Mewaldt put it —94 percent of the birds pass through 1,000 miles of the northeast’s most populous countryside between May 2 and 18, with May 11 the median date for banding migrants in this region. Milton B. Trautman (1956) gives the principal movement at Buckeye Lake, Ohio, as May 8 to 20. In a letter to Ralph K, Bell, Gordon Wilson of Bowling Green, Ky., who has chronicled the birds of his region for 45 years, gives average departure as May 10, with May 28 as a late date.

The only evidence of physiographically-controlled spring migration I have seen is a letter from Clark S. Beardslee to Mr. Bent in 1951. reporting that in spring northbound white-crowns move eastward through Ontario, cross into New York State in the Buffalo-Niagara isthmus, then presumably circle eastward around the south side of Lake Ontario before moving northward into Quebec. Harold D. Mitchell has kindly confirmed these observations for me.

Dispersal into the subarctic breeding grounds on the interior Quebec-Labrador plateaus normally occurs during the last week of May, though late ice and snow may sometimes delay arrival a whole week. I was impressed by the influence of seasonal conditions when I arrived at Knob Lake in interior Quebec on May 18, 1957. A raging blizzard made it plain that winter’s grip was still firm. Returning to within 15 miles of the St. Lawrence above Seven Islands, I found white-crowns abundant and obviously “dammed up” by weather, for they were occupying all sorts of habitats that are not usual for them. They sang softly while awaiting better conditions for concluding their migration. The first birds arrived at Knob Lake on May 23 that year, but even so, they had to spend several days feeding along plowed roadsides until the snow melted from the territories they were to occupy. Harrison F. Lewis’ notes show that conditions were even more severe a decade earlier; his June 4, 1947 journal entry records that this “was an abnormally cold, late spring, with consequent heavy loss of life among small migrant passerines.” He recorded 120 obviously delayed migrant white-crowns at Seven Islands, south and west of the breeding range.

Territory: Territorial behavior has not yet been described for this race of the white-crown. I have twice seen fights in early June in the Knob Lake region, the birds facing each other breast to breast, then jumping, clawing, and flying at each other. Such jousts are usually short-lived.

The white-crown does best in “hybrid” habitats, where disturbance of the surface by fire or mechanical means has increased diversity and the shrub growth is still young. Under such conditions I have found two nests only 300 feet apart, and computed territory sizes as between .88 and 1.85 acres each. On the other hand, a breeding bird census of 18 1/3 acres of open lichen woodland: the most extensive vegetative type of central Labrador —revealed a low density of two nests, or about 1 territory per 9 acres. This would be about 70 pairs per square mile, but Thomas H. Manning (1949) estimated that in western Jugava the population was between 10 and 30 pairs per square mile, the higher counts being in burnt areas.

Courtship: The sexes being alike, it is difficult to follow territorial disputes and courtship until birds have been color-marked. No one has yet done this early enough in the season to unravel this phase of the life history of the eastern subspecies.

Within a week after arrival on the breeding grounds a good deal of territorial song is heard from about 5:00 a.m. to 10:00 a.m. and again in the evening. For a week or so thereafter the birds utter much high-pitched trilling with depressed crowns, either from the ground or from no more than 4 feet above it. This trill is quickly communicated to the whole nearby group, several other birds then repeating it. The trilling bird often attracts a companion who approaches with crest high and loud pete notes, as though alarmed, and they fly off together. Three-party nuptial chases are common at this season. More specifically, the bird that emits the high, trilling dreeeee note often crouches low, with head up, and flutters its wings as though rotated from the “wrist.” I did not notice the tail-spreading and the spasmodic wing opening and closing that caught the attention of Francis Harper (1958) in this same area. As I proved by collecting, the female gives a loud chatter during these mating chases (this female contained oocytes up to 1 millimeter in diameter). Two weeks after the population arrived, while trilling was going on everywhere, I saw a female crouch and trill (I wrote whimper), and the male then mounted quickly three times. Soon afterward song fell off noticeably.

The male of the eastern white-crowned sparrow seems to share some notes and postures heretofore (Nice, 1943, and Blanchard, 1936) ascribed to the female only. On June 13, 1957, I collected a trilling, wing-flipping bird that turned out to be a male. Error is easy under the stress of collecting in close quarters, and a male that entered unobserved may have been the inadvertent victim of my search during the few seconds involved; but, again, on June 24, 1958, I saw a color banded male, the mate of a female with eggs in the nest, rotate its wings “at the wrist,” in the slow wing-flutter that the female gives while trilling in invitation to copulation (Blanchard, 1936).

Nesting: Whereas Alfred 0. Gross (1937) could write that “the height of the breeding season of the White-crowned Sparrow on the Labrador coast is during the first two weeks of July,” for the Labrador peninsula as a whole, most eggs are laid by mid-June. This varies greatly from season to season, however, and from region to region; the season in the higher southern third of the vast peninsula, for example, is usually later than in the lower areas farther north.

In mid-morning on June 16, 1957, I surprised a bird forming a nest. It had pulled out mixed hairy-cap mosses and Cladonia lichens for a proper cup in the shade of a dwarf birch. From a total of some 30 nests examined in situ, it seems safe to consider my description (in Todd, 1963) as typical: “four inches in outside diameter and two and five-eighths inches inside. The inside depth was one and one-half inches. The main body of the nest is woven of fine grass stems, the outside is made up of mixed moss stems, and the bottom is lined with very fine root fibers, or in one case with white ptarmigan body-feathers.” Harrison F. Lewis (in litt.) once found a nest lined with light gray hair, perhaps that of a dog. Most nests set into the moss-lichen or lichen-crowberry mat are partly concealed by overhanging branches of dwarf birch or Labrador tea; not infrequently a nest is neatly tucked into the lower side of a moss or crowberry (Empetrum) mound. Much less frequently, nests are built into moss hummocks on a string of heath shrubbery some distance out in a sphagnum bog. Although I have never seen an elevated nest, Arthur A. Allen wrote me that whereas all the nests he had found in the Churchill region were on the ground, on the Labrador Coast (North Shore) “the few nests I saw were in small firs.” Earlier, Oliver L. Austin, Jr. (1932) had quoted Moravian missionary Perrett’s notes from Makkovik, Labrador, concerning a nest “about three feet from the ground in bushes thrown over a boat to protect it from the sun.”

On June 30, 1957, I found an otherwise typical white-crown nest in a “hybrid” habitat near the airport at Schefferville (Knob Lake), Quebec, which contained 8 eggs and had 3 birds in attendance. On July 2 the eggs and the two birds that came to incubate between 10:45 a.m. and 1:30 p.m. were collected by Don R. Oliver of the McGill University Subarctic Laboratory and shipped to me at Brown University, where William Montagna dissected them. Both birds were females in comparable stages of gonadal regression, with equally developed brood patches. One bird showed two clear follicles and what appeared to be two coalesced follicles; the other had three clear follicles and one questionable follicle. Although polygyny has been reported in Emberizines before (e.g., corn bunting) this appears to be the first American record and is unusual in that only one nest was involved.

Some pairs may mate a second time. On July 10, during their 7th day of caring for young, the banded pair I had under observation at Schefferville, Quebec, performed elements of the nuptial display. The female trilled and fluttered her wings as she left the nest; later I heard her trill while out of sight behind my blind and thought that the sounds indicated a mating chase; that same day the male twittered on crossing her in flight as they exchanged visits to feed the young. E. P. Wheeler, II (in litt.) once found a bird with unhatched eggs as late as July 30 on the Labrador coast.

Eggs: The white-crowned sparrow lays three to five, and rarely six eggs. They are ovate, though some may tend toward either short or elongate ovate, and are slightly glossy. The ground is pale greenish or creamy white and is heavily marked with spots and blotches of reddish browns such as “Natal brown,” “Mars brown,” “chestnut,” “Verona brown,” or “russet.” There is quite a range of variation; frequently the spottings obscure the entire ground, while in other cases considerable ground is showing with the markings concentrated toward the large end where they may become confluent. On eggs with much ground showing, undermarkings of “pale neutral gray” may be discernible. The measurements of 50 eggs of Z.l.leucophrys average 21.5 by 15.6 millimeter; the eggs showing the four extremes measure p24.1 by 16.5, 21.6 by 17.0, 18.9 by 15.8, and 19.8 by 14.5 millimeters.

Of 29 sets recorded, 23 sets had four eggs, 5 had five eggs, and only one had six eggs. I therefore assume the three-egg clutches occasionally reported are probably incomplete.

A color-banded pair had four eggs on June 24, 1958, when discovered, and hatching occurred July 4, so incubation is at least 11 days. The female did all the brooding of the young, so it seems likely that she also did all the incubating of the eggs, as I found her on the nest after dark. Another bird was so bothered by the wind flapping my blind during a 3-hour watch that she was off the nest as much as she was on; she changed every 1½  minutes as a rule, and her longest periods on the eggs were 6 to 7 minutes.

Young: I found that for the first and second days after hatching, the female turns the young, just as she turned the eggs earlier, at 10- to 20-minute intervals. Her brooding schedule is controlled by the male’s visits, for she gets off the nest when he brings her food. He comes silently and directly to the nest, whereas she lands 10 to 15 feet away, always on the same side, then hops in slowly, nearly always using the same perches. On reaching the nest she gives a few alerting chips, to which the young make no vocal response until the 5th day, and the male responds to their trilling only after the 7th day. In 1957 I noted:

“It is surprising to see how well the pale lining of the female’s feathering causes her to blend unobtrusively with the straw border of the nest as she broods. The young grow so quickly that on the third day they push up the female, beg, and gape without parental provocation. They nestle deeply into the nest when the sun dips in late afternoon. On the fourth day their eyes are open during shady intervals, and fully open the next day. They hug the nest when the female scolds and no longer gape at my touch. The tail feathers are now one quarter inch long.”

“By the seventh day the female has trouble brooding since the young toss and turn. The next day they scratch and preen for the first time, and anticipate the parent’s return by trilling.”

Feeding and nest-sanitation are shared almost equally by the parents, the male making slightly more than half the feedings, and removing slightly more than half the fecal sacs. Usually the old birds carry the fecal sacs away, but they occasionally eat the small ones. On their 8th day the young receive feeding visits on an average of one every 10 minutes. On the 9th day they will leave the nest if disturbed, but probably stay on another day or two when not pressed.

Plumages: One-day-old young are fluffy in mouse-gray natal down which covers capital, dorsal, alar, and femoral tracts, but not the ventral tract.

My Labrador notes contain a description of a fledgling about 2 days off the nest. It had dark brown eyes, the bill was brown, and the gape and most of the commissure corn-yellow, the tarsus lilac, and the toenails light horn gray. Francis Harper (1958) describes a young bird as having vermilion mouth linings, with commissure and tomia corn-yellow.

Richard R. Graber (1955) describes in detail the juvenal plumage on the basis of a bird from Colorado and another from Labrador:

“Forehead and crown streaked throughout with black. Crown and forehead white medially, brown laterally. Occiput dark, mottled brown and black. Nape mottled, white and black. Back streaked, black and buff. Rump and upper tail coverts rusty-buff, streaked with black. Rectrices and remiges black. Primaries edged with buff, secondaries and tertials with dull rust color. Uppermost (proximal) tertials edged and tipped with buffy white. Lesser coverts gray; medians black, edged with white; greater coverts black, edged with buff, tipped with white. Two white wing bars. Lores dark, brownish or gray. Narrow white supra-ocular stripe from eye to nape. Auriculars buff-tinged gray, post-auriculars like nape. Chin and throat white, flecked with black, and with black “mustache” marks. Under parts white, or lightly tinged with buff on chest, sides, and crissum. Chest, sides, and flanks heavily streaked with black. Belly and crissum immaculate. Leg feathers dark brown, edged with white.”

Jonathan J. Dwight (1900) presumes that the first winter plumage is “acquired by partial postjuvenal moult, probably in August on its breeding grounds, which apparently involves the body plumage and the wing coverts partly but not the rest of the wings nor the tail.” This is the “brown Livery,” the conspicuously more buffy immature plumage, in which the head is marked by broad reddish-brown stripes instead of the black and white of the adult.

Dwight also writes that first nuptial plumage is “acquired by a partial prenuptial moult beginning the end of March which involves chiefly the head and chin and a few scattering feathers elsewhere. The black and white crown is assumed, which soon shows nearly as much wear as the rest of the plumage. This becomes grayer and the stripes clearer. Old and young become practically indistinguishable” at this stage. Furthermore, “adult winter plumage is acquired by a complete postnuptial moult.”

Robert A. Norris (1954) reports an extensive prenuptial molt during March and April, except for “the alulae, the primaries, secondaries and their greater (outer) coverts, the ten outside tail feathers, and some of the feathers of the “wing lining.” He felt it was “fairly certain that non molting birds such as my mid-March specimens would show * * * overlap between periods of molt and migration,” assuming that completion of the molt requires 2 months.

Amelia R. Laskey of Nashville, Tenn., wrote Mr. Bent that she noticed unusually early beginning of crown molt during the first week of November 1932 and again in 1934. In the experience of both Mrs. Laskey and Ralph Bell, crown molt is normally complete by late April. Bell’s notes show that crown molt takes at least 20 days.

Food: Perhaps the most interesting item on the food of white-crowns is Francis Harper’s (1958) discovery that in spring on the breeding grounds, these birds eat the green capsules of Polytrichum juniperinum, the hairy-cap moss. In 1957 I watched one bird pick and eat 120 capsules in exactly one minute. A female collected on June 8 had nothing but these capsules in her crop, and though all the white-crowns were utilizing this food at the time, some birds also picked up small brown seeds, a few sand grains, and black flies (Simulium sp.).

Like other terrestrial passerines, the white-crown is an opportunist. The hairy-cap moss capsules it consumes in late May and early June, when snows have just melted but before many insects emerge, are at that time the most available food. I have watched them eat the new green catkins of willows. The young are fed insects as nestlings, and themselves catch flies, mosquitoes, and spiders, as everyone who has studied them has noticed. In winter they are primarily seed eaters, and in spring or any other time these are available, they take fleshy fruits such as the red mulberry or the crowberry.

Behavior: The white-crown has long had the reputation of being an aristocrat among the Emberizines. His neat attire, striking crown, and his habit of stretching his head upward to look around have probably combined to earn him this title.

But nobility should imply natural dominance. It was not until I saw white-crowns nesting adjacent to white-throated sparrows at Redmond Lake south of Schefferville, Quebec, that I realized that the white-crown is more mousy than regal in bearing, at least in summer. I found white-throats more deliberate, less upset by intrusion, their alarm notes a quiet announcement of awareness, and their flights shorter. White-crowns, on the other hand, were much more high-strung, always running while on the ground —even if they did this by hopping with both feet —and their alarm notes were more insistent, sharp, or nervous. These differences have an environmental basis, as the white-crowns occupy open country where they are more exposed to potential enemies and pressed by the wind, whereas the white-throats occupy the sheltered brushy borders of the closed crown, Canadian-zone woodland that here reaches a northern limit.

This demeanor changes on the wintering grounds. Albert F. Ganier of Nashville, Tenn., who has studied their ways since the turn of the century, writes me that in his region the white-crowns remain in compact groups of 10 to 20 birds, hugging the same habitat week after week, either in a weed- and brush-grown fence row along some little frequented road, or in an abandoned piece of farmland where plant succession is throwing up clumps and patches of herbaceous and sapling growth. “Here,” he writes, “they feed quietly on the ground, but when intruded upon rise to the top of the low growth and eye the intruder with apparent curiosity, rather than with fear. They crane their necks to get a better look and it would appear that they have not yet evolved a fear of man to the extent of most other birds.” They are apparently not easily flushed out of these preferred coverts, as are the white-throats that fly ahead of the intruder.

Woodward H. Brown (1954) was impressed by the aerial feeding of white-crowns and saw them “occasionally spring 15 —18 inches in the air, returning to former positions on grape vines, catching gnats or other small insects.”

In Quebec-Labrador I was impressed by the fact that females when disturbed always sneaked off the nest for 10 or 20 feet in a sort of “mouse run,” but without the wing-dragging display that shore birds and other species often add before sounding any alarm. On July 3,1958, while crossing a very wet sedge bog near Lake Matamace north of Schefferville, I watched a white-crown, which I took to be a male, catch insects by running in water up to its “knees,” throwing up its tail, raising its white crest high, and “flashing” its wings much as a mockingbird does to flush out the insect life from the bog mat.

On July 10,1944, at Goose Bay, Labrador, I flushed a pair of anxious adults and after 10 minutes of hunting finally drove out a close sitting fledgling which a companion and I captured with difficulty. It was interesting that three adults drove at us frantically as we chased the young bird. They decoyed boldly with the “broken wing” feint, and drew me 40 feet from the young one, returning to me each time I hesitated in following.

In 1958 at an elevation of 2,600 feet on Irony Mountain in central Quebec-Labrador, Henri Ouellet of the Canadian National Museum watched a white-crown feed a still-downy willow ptarmigan. E. P. Wheeler II, who has shared notes made during many years of patient field work in Labrador, noticed that white-crowns sometimes scratch for food very vigorously, using both feet at once as the fox sparrow does. This trait, and the tendency of white-crowns to take easily to man’s newly created habitats in the northern wilderness, as song sparrows and their western congeners do farther south, raises interesting questions about the relationship of all these species. Raymond A. Paynter, Jr. (1964), in the course of a review of some North American Emberizinae, reaffirms the view that the basic differences between the song sparrows, the fox sparrows, and the crowned and white-throated sparrow groups are not clear-cut, and therefore urged that the genera Melospiza, Passerella, and Zonotrichia be merged. The existence of hybrids among the white-crowns (Miller, 1940), between the white-crown and the white-throat (Abbott, 1958), and between the white-crown and the song sparrow and the white-throat and the junco (Dickerman, 1961) lends impressive weight to the Paynter proposal.

Voice: Of ten as I sat in the dusk in my tent or some miner’s shack in the iron ore belt of interior Labrador in 1957, the song of the white-crown reminded me of that of a diminutive eastern meadowlark. “Especially is this so in chorus,” I wrote, “and the first two notes are often like the black-capped chickadee’s sweet-ee call.” I syllabified one common version as, “teu-dee * * * et tu aklavik,” a phrase which will mean more if pronounced in French. Everyone recognizes this song as like an imperfect white-throated sparrow song. To Ludlow Griscom it had the quality of a black-throated green warbler song, while Francis H. Allen wrote Mr. Bent that to him the song was “doleful rather than plaintive —the sweet expression of a state of utter boredom, as if the bird were saying, ‘Oh, well, what’s the use?”‘

Harrison F. Lewis told Arthur A. Allen that he rendered the song as “Oh gee: it was the whiz-whiskey,” to which the fox sparrow, so often a neighbor in the northland, would answer, “Well, my dear, why did you take it?”

The open, windy nature of the semibarrens that the white-crowns occupy in summer diminishes the impression their song makes on human visitors; so many songs are wafted away on the wind that only dominant phrases force themselves on the traveler’s attention. The word renditions given above are the subjective efforts of pre-technological field ornithology. Today’s field students use tape recorders and analyze their input from the visual record of a sound spectrograph that allows direct reading and comparison of duration and pitch. Donald J. Borror (1961) has analyzed a number of white-crown song recordings made by W. W. H. Gunn in northern Ontario and has provided the following objective description:

The song usually begins with one to three clear whistled notes that are steady in pitch, and ends with three buzzy notes, the last lower in pitch than the two preceding. The first note is about 0.5 sec in length; if there are two or three similar introductory notes the second and third are a little shorter. The final buzzy notes are uttered at three to four per sec. Sometimes there are short clear notes or two-note phrases in the middle of the song and sometimes the second or third note of the song is slurred. One or two of the final buzzes (except the last) may begin with a short sharp note or be slightly up-slurred or both. Some songs end in a low trill rather than a low buzzy note. The songs of a given bird are usually very similar, hut those of different birds often vary slightly.

The pitch of the white-crown’s song is between 2,600 and 7,200 cycles per second.

Morning and evening song periods usually involve 15 to 20 minutes of uninterrupted song, and as each song is of 2-second duration and the interval between songs is 9 to 10 seconds, the total output during such a burst may be 100 or more songs. I have counted 194 consecutive songs.

Francis Harper (1958) writes the principal call note as “a tsit, which, when heard near at hand, seems to have a slight metallic rasp.” Charles W. Townsend and Glover M. Allen (1907) distinguished a metallic chink call note from a sharper chip alarm note. In my field notes I described the call note as pete, identical by both sexes, and the alarm notes as higher pitched than the ordinary scold note.

Amelia R. Laskey (in litt.) mentions a ventrioquial song of the immature as follows: “At first the songs, coming at intervals, seemed to emanate from shrubs some 15 feet behind the bird, but as it came closer I could see its bill open and close. It was a lengthier song than the adults give in spring, and the bird erected its crown feathers as it sang.”

Mortality: It seems to me better to get away from the connotations of the word “enemy” and simply to point out that the white-crown is subject to the usual factors that cause attrition in animal populations, whether disease, the complex of factors engendering winter mortality, or direct predation by accipitrine hawks, shrikes, weasels, and the like.

It has its normal share of parasites, both external and internal. Oscar M. Root has kindly furnished a note on the identification of Hippoboscid louse-flies, Ornithomyia fringillina, found on immature birds by Gary C. Kuyava in Minnesota; Francis Harper (1958) has taken a mite of the genus Lealaps from a juvenile specimen in Quebec; and Robert A. Norris (1954) found biting lice (Mallophaga) on dried skins and also found that four out of nine specimens examined in Georgia had protozoan infections of the blood (Leucocytozoon), and one of these, a smallish individual which had not begun its prenuptial molt on March 17, was doubly infected with the malarial parasite, Plasmodium. One adult was heavily infected with abdominal helininths, the filarid nematode Diplotriaena. The individual infected with Plasmodium also had foot tumors caused by the virus Epithelioma cordagiosm. Alfred 0. Gross (1937) reports the mallophagan Philopterus subflavescens (Geof.) from young on the Labrador coast, and Herbert Friedmann (1938) reports parasitism by the cowbird at Okotoks, Alberta.

Of greater population significance, probably, is the loss of young birds during the first migration. For the Quebec-Labrador segment, especially, this must be a significant decimating factor because the young of the year are often wind-drifted out to sea, where they perish unless they are fortunate enough to reach an island from whence they can return. I have been particularly impressed with this problem in their lives at Block Island, R.I., where hundreds of white-crowns appear in autumn, when cold fronts pass out to sea, all of them immatures.

Fall and winter: Young were on the wing as early as July 11, 1945, at Indian House Lake in northern Labrador, and Austin (1932) saw young flying on July 16, 1928 on the coast, although Late July is a more normal date there. In August at Indian House Lake they were in loose family groups, feeding and playing in the alder strand that fringes the George River, and by mid-September when they leave the region, they are usually restricted to dwarf birch thickets in timberline areas on the slopes. On July 31 and again on August 3, 1957, at Scheffeville, Henri Ouellet noted a goodly number of both adults and young in the open, “feeding very little, and seemingly on the move.” E. P. Wheeler’s last date for Kutsertakh on the Atlantic slope of Labrador is Oct. 5, 1934.

At Buckeye Lake, Ohio, Milton B. Trautman (1940) records first arrival as October 1 to 8, with the peak of migration October 10 to 27, and the last departure November 3. White-crowns first wintered there in 1953. The ratio of immatures to adults, M. B. Trautman reports, was usually 2 to 1, though in some years there were 97 immatures to 3 adults. As earlier mentioned, on Block Island, 10 miles off the Rhode Island shore, immature birds are almost or quite alone, which suggests that adults are too experienced to allow themselves to be wind-drifted out to sea during migration. Robert A. Norris (1954) reports a wintering flock of 30 immatures to 1 adult in Georgia; he considered the sex ratio balanced.

Concerning autumn habitats, Milton B. Trautman (1940) writes, “As in spring, the birds were found in brushy situations, but many were also present in dense patches of high weeds, and in weedy uncut cornfields. Autumn sparrows were somewhat more secretive than were spring birds, and it was only by remaining quiet in a dense weed patch or brushy thicket and giving a Screech Owl whistle that a true indication of numbers could be obtained.”

Robert A. Norris (1954) writes that white-crowns do not flock with other species and that on the Georgia wintering grounds he studied, the birds are “found in more open country with less cover and also farther from water than is typically the case with White-throated Sparrows.” The March-April weights of his Georgia birds ranged from 23.7 grams for the smallest female to 31.2 grams for the largest male, the 13-bird sample averaging 30.05 grams. In a larger sample, however, Paul A. Stewart (1937) found that 21 adults ranged from 19.9 to 37.1 grams, with a mean of 31.26 grams; 31 immature birds ranged from 23.5 to 32 grams, with a mean of 27.56 grams.

Cortopassi and Mewaldt’s (1965) plot of the distribution of this species from the Christmas Bird Counts of Audubon Field Notes shows that the wintering population has two centers of concentration, one m western Texas and southeastern New Mexico and the other in the Appalachian plateau and its western extension, the interior low plateaus and Ozark plateaus. In these two broad belts the bird-watcher may expect to see from 1 to 10 birds per hour afield during a full day’s quest. They warn, however, that the Texas-New Mexico area of concentration may be the result of the oasis-like nature of suitable habitat, and the special attention this receives from the bird-counters. According to their analysis of banded bird data, only the eastern race of the white-crown is regular east of the 90° parallel. Between 90° and 105° (the Great Plains region) the eastern and Gambel’s races migrate and winter together. Intergrades from the west aide of Hudson Bay winter mostly in the Dakotas.

The eastern white-crown has apparently been extending its winter range both eastward and northward since about 1950. A very few now winter fairly regularly as far northeast as the New York City region (John Bull, 1964), where winter reports were hardly credible 20 years earlier (Cruickshank, 1942), and quite unknown when Ludlow Griscom (1923) summed up the status of that region’s bird life. The climatic amelioration of the first half of the century may have facilitated this range expansion, but such new land-use practices as the planting of multiflora (Rosa multiflora) hedges and the great number of bird-feeding stations that have come into vogue during this period have unquestionably helped tide over individual birds.

DISTRIBUTION

Eastern White-crowned Sparrow

Range: Montana, northern Ontario, and Labrador south to central Mexico.

Breeding range: The eastern white-crowned sparrow breeds from north central and northeastern Manitoba (Churchill, Cape Tatnam, intergrades with Z. 1. gambelji), northern Ontario (Fort Severn), northern Quebec (Richmond Gulf, Fort Chimo), and northern Labrador (Port Burwell) south to central northern Ontario (Fort Albany), central and southeastern Quebec (Lake Mistassini, Godbout, Blanc Sablon), and northern Newfoundland (Flower Cove, St. Anthony).

Winter range: Winters from Kansas, central Missouri (Kansas City, St. Charles County), central Kentucky (Louisville), West Virginia (Charleston), and western North Carolina (Asheville) south to Sinaloa (Elota), Aguascalientes, Nuevo León (Monterrey), northern Tamaulipas (Matamoros), Louisiana (Natchitoches, Houina), and south central Georgia (Tif ton, Savannah); casually north to southern Michigan (Jackson) and southern Ontario (Toronto); south rarely to southern Mississippi (Saucier), northwestern Florida (Pensacola), and Cuba.

Casual records: Casual in north central Alaska (Togulak Lake), northern Franklin (Fletcher’s Ice Island at 82°37’ N., 99°50’ W.), Baffin Island (Taverner Bay, Lake Harbour) Greenland (Godthaab, Fiskenaes), and in Bermuda.

Migration: The data deal with the species as a whole. Early dates of spring arrival are: North Carolina: Raleigh, April 14. West Virginia: Morgantown, May 3. District of Columbia: March 26 (average of 14 years, April 26). Maryland: Anne Arundel County, April 10; Montgomery County, April 20; Laurel, May 2. Pennsylvania: Renovo, April 22; Meadville, April 23; Beaver, average of 14 years, May 7. New York: Brooklyn, April 10; Yates County, April 23. Connecticut: West Hartford, April 15; New Haven, May 4. Rhode Island: Jamestown, May 7. Massachusetts: Martha’s Vineyard, April 2. Vermont: Peacham, April 5; St. Johnsbury, April 14. New Hampshire: Tamworth, April 18; Monroe, April 21. Maine: Portland, May 9; Lake Umbagog, May 12. Quebec: Quebec City, April21; Montreal area, April 30 (median of 7 years, May 6). New Brunswick: Andover, May 8. Nova Scotia: Shelburne, May 6; Wolfville, May 13. Prince Edward Island: Canoe Cove, May 27. Newfoundland: Stephenvile Crossing, May21; St. Anthony, June 1. Illinois: Urbana, April 1 (median of 20 years, April 30); Chicago, April 26 (average of 16 years, April 29). Indiana: Lafayette and Indianapolis, March 31. Ohio: Oberlin, April 21 (median of 19 years, May 2); Buckeye Lake, April 23 (median of 40 years for central Ohio, April 28). Michigan: Detroit area, April 20 (mean of 10 years, April 25); Battle Creek, April 28. Ontario: Leeds County, April 22; Ottawa, April 30 (average of 24 years, May 7). Iowa: Sac County, April 26; Sioux City, April 30 (average of 32 years, May 5). Wisconsin: Grantsburg, April 6. Minnesota: Pipestone, March 30 (average of 34 years for southern Minnesota, May 4). Oklahoma: Payne County, March 3. Nebraska: Hastings, March 31; Holstein, April 10; Red Cloud, April 19 (average of 20 years, April 30). South Dakota: Faulkton, April 3; Sioux Falls, April 13 (average of 6 years, May 4). North Dakota: Lower Souris Refuge, April 23; Cass County, April 29 (average, May 4). Manitoba: Treesbank, April 29 (average of 17 years, May 8). Mackenzie: Great ‘Slave Lake, May 13. New Mexico: Los Alamos, March 15 (median of 5 years, March 24). Colorado: Derby, March 30. Utah: St. George, April 28. Wyoming: Careyhurst, March 11; Laramie, March29 (average of 7 years, April 20) ; Cheyenne, April 18 (average of 9 years, April 27). Montana: Libby, March 24 (median of 10 years, April 24); Terry, April 21 (average of 5 years, May 5). Alberta: Warner, April 21. Oregon: Rickreall, March 9. Washington: Shelton, March 20; Potholes, April 10. British Columbia: Milner, March 25. Yukon: Macmillan River, April 28. Alaska: College, May 4; Kenai, May 14; Kobuk, May 21.

Late dates of spring departure are: Florida: southern peninsula, April 27. Alabama: Marysville, May 14; Montgomery, April 28. Georgia: Macon, May 5. South Carolina: Charleston, May 5. North Carolina: Asheville, May 10; Wilkes County, May 3. Virginia: Blacksburg, May 17; Cape Henry, May 15. District of Columbia: May 22 (average of 16 years, May 15). Maryland: Baltimore County, May 26; Prince Georges County, May 22. Pennsylvania: Wilkinsburg, June 5; State College, May 25. New Jersey: West Milford and Morristown, May 27; Cape May, May 7. New York: Westchester County, June 14. Connecticut: East Windsor Hills, May 30; Portland, May 22. Massachusette: Nantucket, May 26; Martha’s Vineyard, May 23. Vermont: Topsham, May 24. New Hampshire: Hanover, May 26. Maine: Lisbon, May 23. Quebec: Arvida, May 31. Nova Scotia, May 24. Louisiana: Baton Rouge, April 27. Mississippi: Rosedale, June 2 (mean of 19 years, May 12). Arkansas: Fayetteville, May 16. Tennessee-Knox County, May 21; Nashville, May 20 (median of 20 years, May 11). Kentucky: Bowling Green, May 11. Missouri: St. Louis, May 30 (median of 15 years, May 10). Illinois: Chicago, May 31 (average of 16 years, May 25); Urbana, May29 (median of 20 years, May 17). Indiana: Wayne County, May 24 (median of 16 years, May 11). Ohio: central Ohio, May 29 (median of 40 years, May 20); Oberlin, April 22 (median of 19 years, May 19). Michigan: Detroit area, June 4 (mean of 10 years, May 24). Ontario: Point Pelee and Whitby, May 29. Iowa: Floyd County, May 21; Sioux City, May 19. Wisconsin: Wausau and Viroqua, May 30. Minnesota: Cloquet, May 25 (average of 6 years for southern Minnesota, May 19). Texas: Sinton, May 7 (mean of 5 years, April 22); Austin, April 29. Oklahoma: Oklahoma City, May 26; Tulsa, May 17. Kansas: northeastern Kansas, May 30 (median of 18 years, May 9). Nebraska: Holstein, May 24. South Dakota: Faulkton, June 14; Sioux Falls, May 16. North Dakota: Cass County, May 25 (average, May 22); Jamestown, May 18. Saskatchewan: Kinloch, June 6. New Mexico: Los Alamos, May 29 (median of 6 years, May 13). Arizona: Tucson, June 6. Colorado-Colorado Springs, June 17. Utah: Kanab, May 14. Idaho-Moscow, May 28 (median of 11 years, May 16). Montana: Libby, June 2 (median of 10 years, May 12); Choteau, May 22. California: Orange County, May 22. Nevada: Mercury, April 28. Oregon: Linn County, May 28; Baker County, May 24; Lake County, May 20. Washington-Spokane, May 22.

Early dates of fall arrival are: Washington: Shelton, August 21; Hidden Lakes, August 30; Potholes, September 14. Oregon: Baker County, September 6 Wheeler County, September 13. Nevada: Clark County, September 13; Mercury, September 21. California: Lafayette, September 3. Montana: Libby, August 27 (median of 10 years, September 9). Idaho Moscow, August 27 (median of 11 years, August 29). Utah: Uinta Basin, September 10; La Sal Mountains, September 13. Arizona: Wagner, September 9. New Mexico: Los Alamos, September 5 (median of 6 years, September 10). Saskatchewan: Sovereign, September 5. Manitoba: Treesbank, September 10 (average of 9 years, September 22). North Dakota: Jamestown, September 15; Cass County, September 18 (average, September 22). South Dakota: Rapid City, September 11; Faulkton, September 13. Nebraska: Neligh, September 22; Lincoln, September 28. Kansas: northeastern Kansas, September 28 (median of 10 years, October 12). Oklahoma: Copan, October 6. Texas: Sinton, October 19 (median of 5 years, November 4). Minnesota: Minneapolis-St. Paul, September 18 (average of 14 years for southern Minnesota, September 29). Wisconsin: Chippewa County, September 3; Wausau, September 9. Iowa: Sioux City, September 24 (average of 32 years, October 15). Ontario: North Ray, September 7; Peel County, September 8. Michigan: Detroit area, September 17 (mean of 10 years, September 23). Ohio: central Ohio, September27 (median of 40 years, October 6). Indiana: Chesterton, September 16; Wayne County, September 30 (median of 15 years, October 9). Illinois: Chicago, September 12 (average of 16 years, September 22). Missouri: St. Louis, September 25 (median of 15 years, October 3). Kentucky: Glasgow, October 3. Tennessee: Nashville, September 28 (median of 20 years, October 15); Knox County, October 18, Arkansas: Arkansas County, September 20; Fayetteville, October 8. Mississippi: Rosedale, October 2 (mean of 31 years, October 18); Saucier, October 12. Louisiana: Baton Rouge, October 28. Nova Scotia: Louisbourg, September 22. Prince Edward Island: Ellerslie, September 22. Quebec: Highmore, September 11. Maine: Lake Umbagog, September 23; Cumberland Mills, September 26; Portland, October 1. New Hampshire: Tamworth, September 20. Vermon: Hartland, September 10; Wells River, September 26. Massachusetts: Nantucket, September 19. Rhode Island: Newport County, September 27. Connecticut: Hartford, September 24. New York: Essex County, September 20; Long Island, September 23. New Jersey: Cape May, September 30. Pennsylvania: Erie, September 19; Renovo, September 23. Maryland: Baltimore County, September 27; Laurel, October 2 (median of 5 years, October 9). District of Columbia: October 1 (average of 7 years, October 13). Virginia: Blacksburg, September 14; Cape Henry, October 8. North Carolina: Mills River Valley, October 6. South Carolina: Berkeley County, October 9; Charleston, median of 5 years, October 20. Georgia: Athens, October 25. Alabama: Dauphin Island, October 7; Birmingham, October 17. Florida: Tallahassee, October 4; Pensacola, October 9.

Late dates of fall departure are: Alaska: Chena Hot Springs, October 18; Cohoe, October 5. Yukon: Macmillan Pass, September 4. British Columbia: Okanagan Landing, October 30. Washington: Wawawai, November 9. Oregon: Coos Bay, October 18. Alberta: Glenevis, October 10. Montana: Libby, October 20 (median of 10 years, September 30). Wyoming: Laramie, October 30 (average of 8 years, October 22); Cheyenne, October 14 (average of 6 years, October 4). Utah: Spectacle Lake, October 26. Colorado: Colorado Springs, November 12. Arizona: Parker Dam, October 30. New Mexico: Los Alamos, December 1 (median of 6 years October 28). Saskatchewan: Regina, November 5. Manitoba: Treesbank, October 27 (average of 8 years, October 2). North Dakota: Grafton, October 14; Cass County, October 9 (average, October 7). South Dakota: Faulkton, November 10. Nebraska: Neligh, November 1. Oklahoma: Payne County, November 14. Minnesota: MinneapolisSt. Paul, November 2 (average of 6 years for southern Minnesota, October 18). Wisconsin: Milwaukee, October 28. Iowa: Sioux City, October 30. Ontario: Ottawa, November 4 (average of 15 years, October 8). Michigan: Detroit area, mean of 10 years, November 11; Battle Creek, October 26. Ohio: central Ohio, November 30 (median of 40 years, October 29). Indiana: Indianapolis, November 23. Illinois: Chicago, November 9 (average of 16 years, October 23). Mississippi: Saucier, October 31. Newfoundlland: St. Anthony, October 5. Nova Scotia: Shelburne, November 6; Yarmouth County, October 14. New Brunswick: Scotch Lake, October 30; Grand Manan, October 5. Quebec-Montreal area, October 29 (median of 7 years, October 17); Kamouraska, October 27. Maine: Falmouth, October 28; Lake Umbagog, October 13. New Hampshire: Ossipie, November 9; Concord, October 24; New Hampton, median of 21 years, October 22. Vermont: Woodstock, October 29; Wells River, October 26. Rhode Island: Newport County, November 24. Connecticut: Bristol, November 2; New Haven, October 26. New Jersey: West Milford, November 14. New York: Suftolk County, November 17; Saratoga County, October 31. Pennsylvania: Wilkinsburg and Indiana, October 30. Maryland: Baltimore County December 6; Laurel, December 4 (median of 4 years, November 11). District of Columbia: November 28. Virginia: Cape Henry, November 2.

Egg dates: Alaska: 213 records, April 1 to July 11; 110 records, June 6 to June 16.

Labrador: 38 records, June 3 to July 11; 24 records, June 20 to June 30.

Ontario: 5 records, June 17 to June 28.

NUTTALL’S WHITE-CROWNED SPARROW *

ZONOTRICHIA LEUCOPHRYS NUTTALLI Ridgway

Contributed by BARBARA BLANCHARD DEWOLFE **

HABITS

White-crowned sparrows are among the best known and most widely distributed passerine birds of North America. They live close to man, and their original range may well have been extended as a result of man’s activities, for the combination of bare ground, grass, and dense shrubbery they prefer is often concomitant to road-building, lumbering, farming, or burning. White-crowned sparrows are easily trapped in migration and on the wintering grounds, and large numbers are banded each year.

The breeding range of the species’ western populations extends from sea level to over 11,000 feet and spans some 3,000 miles both north to south and east to west, from latitude 70°N. on the Arctic slope of the Brooks Range to latitude 34°N. at Gaviota, Calif. and from longitude 105°W. in central Colorado to longitude 1680W. on the Seward Peninsula in Alaska. Within this large geographic area the species is divided into four races which differ only slightly in color pattern, shade of plumage, and other morphological characters, but which vary markedly as to strength of migratory instinct and the timing of the breeding cycle. Nuttall’s sparrow (Z. l. nuttalli) is permanently resident in the fog belt of California, and begins to nest in March or April, depending on the year. The Puget Sound sparrow (Z. l. pugeIensis) comprises populations with every degree of migratory instinct, from those in northern California that forsake their territories in winter to flock only a few hundred yards away, to those of the Canadian border that fly a thousand miles each year between wintering and breeding grounds. This race begins nesting in early to late April, depending upon the latitude of the breeding population in question. Gambel’s sparrow (Z. l. gambelii) is a strongly migratory race. It starts to nest in late May and early June in Alaska, and in the provinces and the Northwest Territories of Canada. The mountain white-crowned sparrow (Z. l. oriantha) is also migratory, and begins to breed from late May to early July in Alberta and the western United States, depending upon when the high mountain meadows become sufficiently free of snow to permit nesting.

Beneath this variability in migratory instinct and in the timing of the breeding cycle lie patterns of behavior common to all four races: the themes on which the variations are based. During territory establishment and defense, courtship, nest-building, incubation, and care of the young, the behavior is similar in all races. Therefore the accounts in the section on “Habits” based on my observations of Nuttall’s sparrow at Berkeley apply also to the Puget Sound sparrow and Gambel’s sparrow during comparable phases, and also to the mountain white-crowned sparrow insofar as my limited field work on this race permits comparisons. In fact, so similar is the behavior in all populations of white-crowned sparrows I have studied during the breeding season, that as I watched them I felt as if I were seeing the same birds, whether in central California, at the Canadian border, or in Alaska. Such racial variations as do exist lie not in the behavior patterns themselves but in the time of year the patterns emerge, and in the duration of the successive phases of the nesting cycle. These variations will be described in the sections under each race, which follow the general account for the species.

Where the observations are neither my own nor quoted from published accounts cited in the bibliography, I refer in the text to the observer who, through personal communication, supplied me with the data. If no citation is made, the statements are based on my field work done in the following localities for the periods stated:

Z. l. nuttalli. California: Berkeley (five years).

Z. l. pugetersis. California: Berkeley (five winter seasons); Eureka (one winter and one spring).
Oregon: Tillamook (one spring).
Washington: Friday Harbor (one spring and summer).

Z. l. gambelii. California: Davis (one winter season); Santa Barbara, (ten winter seasons).
Alaska: Mountain Village (one spring and summer); College (one spring, sum mer, and fall).

Z. l. oriantha. California: Sierra Nevadas and Lassen National Park (one summer).

Spring: Spring is the time of transition from flocking to paired isolation on mutually exclusive areas. Territorial jealousy and sexual interest, which have been at low ebb during winter, emerge and rise to maximum intensity. The male sings with increasing frequency and vigor; he pursues and fights rival males, and the female trills and postures. At first these elements of territoriality and sexual interest may appear separately, unorganized into any integrated behavior. Later they are woven into a characteristic pattern. The end result is that pairs space out in orderly fashion and stay on their territories at least until the young of the last brood become independent.

In early, mid, or late January, depending on the weather, the male Nuttall’s sparrow begins to sing more forcefully than in winter, more frequently, and from a more conspicuous perch. The adult male becomes less and less tolerant of the immature or mateless birds that have spent the winter in his area. When their song also increases in force and frequency, he pursues and attacks them until, about 3 weeks after the beginning of this phase, he regains sole possession of all or part of the territory he patrolled the year before. With this achievement, if he is already mated, he stops singing except for rare, weak songs. Boundary disputes or pursuits are also rare. Settlement is now complete, some 7½ weeks before incubation starts.

The immature or mateless males usually find mates during the period of singing, pursuits, and fighting. If they do not, they continue to sing after the mated males are silent. If a male loses his mate, he resumes loud singing and continues it until another female joins him.

Color-banding reveals that, except in instances of polygamy, territorial jealousy is not expressed by the female Nuttall’s sparrow. Her weak song, which continues until nesting time, is not used for advertisement or warning. I have never seen her join in a fight and only rarely in a chase in which her mate was involved. In polygamy, on the other hand, all elements of territorial jealousy appear. The development of the territorial sense in the female independently of the male is illustrated in the behavior, in 1935, of females I and III, simultaneous mates of male I. The description is taken from Barbara D. Blanchard (1941):

With the approach of reproduction * * * each female created for herself a subdivision of the main territory which she defended against the other female by loud singing and fighting, and in which she finally chose her nest site. From February 1 until late March, by which time both had nests, each female sang frequently from a favorite perch within her section. * * * Twice, when female III followed the male toward the section which belonged to female I, a fight ensued between the two females. They locked feet and jabbed each other on the breast. * ** Had they not been banded, I should have thought I was watching a boundary dispute between two males.

Territory: Just as the behavior involved in territory establishment is nearly identical in the four races, so is the nature of the site chosen for the territory. Despite the variety of climatic and vegetational zones in which the four races nest, all the territories I have observed are strikingly similar in their appearance. So marked is the similarity that I think it must reflect deep seated preferences common to the western portion of the species: a conservative psychological theme with minor racial variations.

The similarity in appearance of the white-crowned sparrow territories at different latitudes and altitudes lies not in the presence of any one common element: such as a particular species of plant-or in any one topographic feature, but in a subtle yet definite combination of three elements: grass, bare ground, and shrubbery. These invariably exist in each male’s territory. Two other elements are usually present near each territory but not necessarily within it: water, either salt or fresh, and tall trees.

The nature of the grass, hare ground, and shrubbery vary with climate and topography. The grass may be pure or mixed with lichens or mosses or sedges and small flowering plants. The bare ground may be a sandy beach or the rocky shore of a lake or river. It may be a road or a pack trail, a ploughed field or a clearing in the forest. The shrubbery may consist of any species of plant, native or exotic, that grows thick enough to shelter a nest or to provide a roost. Bracken fern, scrub conifers and alders and willows, salal, Labrador tea, lupine or rose or Artemesia bushes, and many exotic ornamental shrubs are examples of plants that make up this common element.

To list the three elements and the forms they take in different climates is not, however, to define a white-crowned sparrow’s territory. Rather it must be understood in terms of the bird’s temperament and habits. It seems to me that the essential feature is the mixture of grass, bare ground, and shrubbery in just the right proportions to permit ground foraging with quick escape to shelter. The areas of bare ground and grass must be large enough to facilitate foraging, but not so large as to require the birds to move more than a few wing beats away from cover. The shrubbery must be extensive enough to conceal a nest and the devious route the female takes to and from it, but not so unbroken in extent as to require a long flight to reach open ground. Extensive areas of grass, bare ground, and shrubbery existing side by side in solid masses with abrupt linear interfaces do not constitute typical white-crowned sparrow country. What is most typical of a territory is its patchy appearance, because of the interstitial invasion of the open ground by shrubbery.

Such subtle characteristics of a landscape are easy to sense, but hard to describe. Yet, owing to the presence of the same characteristic elements with, of course, almost infinite local variations, the territories of white-crowned sparrows throughout western North America look much alike. An example of this is the similarity in appearance, or perhaps in “mood,” of the landscapes at Desolation Lake in the high Sierras and at Point Lobos on the Monterey peninsula, both of which support breeding populations of white-crowned sparrows. In June of 1960 I found male white-crowned sparrows of the race oriantha spaced out and singing regularly along the edge of Desolation Lake in the Wilderness Area above and west of Lake Tahoe. I assume they were either nesting or about to nest there. At Point Lobos on the shore of the Pacific Ocean near Monterey, Calif., Nuttall’s sparrows nest abundantly.

In spite of the obvious differences in the details of the two landscapes, the elements characteristic of white-crowned sparrow territories are present in both and confer on the two localities a common stamp. The high mountain lake is bordered by masses of bare granite with gnarled juniper trees and dwarfed pines. Pack trails meander along the edge of the lake, and tufts of grass and patches of alpine flowers grow in the lee of granite boulders. The same elements of a large body of water, bare ground, shrubbery, and grass comprise the landscape at Point Lobos. Here a similar scene is produced by the juxtaposition of ocean, steep granite cliffs, and wind-racked Monterey cypress. The grass is more extensive and the flowers more conspicuous, but the impression conveyed to me by both landscapes is essentially the same, that of a wind-swept barren land with warped trees, and a lee with sheltered spots where grass and flowers grow.

A second example of the similarity of white-crowned sparrow territories in widely separated areas is found in Nuttall’s sparrow country near coastal farm lands of California and in Gambel’s sparrow country on the outskirts of Fairbanks, Alaska. In the summer of 1956 1 found Nuttall’s sparrows breeding on the seaward edge of a farm in San Luis Obispo County. Their territories included both cultivated and wild land. They foraged in a pasture, used the fence posts for singing perches, and the native shrubbery beyond the pasture for shelter and nest sites. In the same pasture cattle grazed, dogs dug for pocket gophers, and domestic ducks waddled at the edge of the swampy patches.

In the summer of 1957 I found a similar landscape in Alaska near the cultivated fields of the University of Alaska’s Agricultural Experiment Station. Gambel’s sparrows had incorporated the cultivated land into their nesting territories. They foraged in the fields, roosted in the willow and rose tangles at the edge, and nested in the tall thick grass. Even the details of the scene reminded me forcibly of Nuttall’s sparrow country 3,000 miles to the south: I found tracks of cloven-hoofed mammals at the edge of the field, in the distance I saw a pair of small carnivores hunting mice, and ducks moved along in the stubble at the far edge of the cultivated plots. It took considerable effort to realize that this was an Alaskan scene, not a California one, that the hoof-prints were those of moose, not cattle, that the carnivores were native red foxes, and that the ducks were wild.

This second example brings out another characteristic of many, if not most, white-crowned sparrow territories: their close connection with human habitation or with country altered by man. This is a feature not only of territories in the densely populated areas of California, but also of those in many parts of Alaska as well. In 1950 I went to Mountain Village, an Eskimo trading post on the lower Yukon, to watch the arrival of Gambel’s sparrows. One boundary of the village is the river. Above the village scrub alder and willow merged into upland tundra. There were no roads or cultivated fields. I found arriving Gambel’s sparrows carving out territories in the village, beside fish camps, even by the schoolhouse and trading post, and including in their areas the river’s edge, which was used by the villagers as a thoroughfare to and from the trading post and as a place to tie their dogs. The undisturbed ground above the village was apparently also suitable for white-crowned sparrows, as a few pairs settled there, but only after all suitable areas in the village had been preempted.

I persisted in my search for Gambel’s sparrows nesting on land in its natural state, and one day I found what appeared to be just such a spot about 6 miles upriver from the trading post. There I found open grassy knolls with bare ground grooved by tiny streams and surrounded by dense scrub alder. Gambel’s sparrows were breeding there, and I found no trace of human beings, not even a rusty knife or a fragment of sawed wood. Imagine my dismay when, on describing the spot to the trader that evening, I found that this clearing was all that remained of a once-thriving Eskimo village that had been abandoned about 40 years before!

Courtship: Courtship behavior includes both pairing and copulation. The two processes may be separated by several weeks. Like territory establishment, the elements of behavior involved in courtship are the same for all races. Such variations as occur involve the duration of courtship behavior, its relation to the stage of the reproductive cycle, and the length of time elapsing between pairing and copulation. Except in Nuttall’s sparrow, I have little or no evidence of the permanence of the bond between mates.

As with few exceptions the Nuttall’s sparrows I watched at Berkeley remained paired for life, only the immature and bereft adults sought mates in the spring. The rising intensity of the male’s song and of the female’s trilling and posturing thus may serve two different purposes in courtship: either they bring together two unmated birds and result in pairing, or they intensify the permanent bond between members of a pair that bred together the previous year. In either case the elements of behavior are the same. I shall describe them in adults already paired.

In January the female’s interest in her mate gradually intensifies, expressed by low metallic trills and wing fluttering. At first her trilling is sporadic and seems to be called forth only by the loud song of a male (not necessarily her mate) or by a chase or fight in which the mate is involved; it may or may not be accompanied by fluttering of the wings. As the season advances, trilling and posturing are almost invariably linked, and more and more often occur independently of any apparent external stimulus. Both actions increase in intensity and frequency until the peak is reached some 6 to 8 weeks later, just before copulation, in early to late March, depending upon the year. As the female becomes engaged in nesting, both trilling and posturing cease, to be resumed in lesser degree prior to copulation for the second brood.

The intensification of the male’s interest in his mate, insofar as I can judge from his behavior, is much more sudden. During January and February he seems indifferent to her trilling and posturing. He pays no special attention to her other than to forage with her and to utter location notes as he has done throughout the fall and winter. From early March, however, he punctuates long periods of indifference by “attacks” upon the female. Suddenly, with no warning that I can detect, his indifference changes to aggression. He chases the female and jabs her with his beak. Such “attacks” take place as much as 18 days before, and also a few days after, the first observed copulation. Another change that precedes coition by from 1 to 3 weeks is the beginning of evening song. During the hall-hour before dark the male intersperses periods of foraging with faint singing, which seems directed at his mate rather than at neighboring males. Such song gains in force and frequency until on the evenings when copulation occurs he follows his mate about, singing loudly before and after he mounts her.

Copulation occurs most often during the half-hour before dark, but may take place also during the day. The female follows her mate about and trills persistently, fluttering her wings and raising her tail. Suddenly she flies straight away from her mate, and lands either on the ground or in a tree some yards distant. Her mate follows and lands near her. He hops toward her, crown raised and tail lowered and spread, flutters above her a few seconds, then flies to a nearby perch and sings. I have seen the same pair copulate as many as 11 times in one evening. The average for 21 pairs is 5 times. The period during which the pair copulate lasts 3 to 6 days.

Since immature birds begin their courtship at the same time as the adults, they may pair as early as January, and the interval between pairing and copulation may last 6 weeks or more. The long courtship period characteristic of Nuttall’s sparrow is a consequence of the fact that courtship begins when both sexes are in the early phases of the reproductive cycle, with immature gonads only a small fraction of their breeding size. This is not true for the migratory races.

In Nuttall’s sparrow polygamy is not uncommon. In 5 years of observation of color-banded birds at Berkeley, I found three cases of a male with two mates.

Nesting: The typical nest site is similar in the four races. The nest is usually placed either on the ground or a few feet above the ground, under or within dense but not necessarily extensive vegetation of whatever kind affords adequate concealment. Some of the species of plants in which nests have been found are conifers, especially young pines and spruce (Bolander, 1906; Farmer, 1952; Jewett, 1916; Johnston, 1943; Kobbé, 1900; McHugh, 1948; Ray, 1906; Warren, 1912); scrub oak, willow and alder (Farmer, 1958; Grinnell, 1900; Macoun, 1909; Ray, 1912); dwarf birch (Dice, 1920); bushes of wild rose, lupine, sage, thimble berry (Jewett, 1916; Ray, 1906); composite perennials such as Ericameria and Eriophyllum (Grinnell and Linsdale, 1936); many species of ornamental exotic shrubs; and, rarely, an annual plant (Grinnell and Linsdale, 1936, found a Nuttall’s sparrow nest at Point Lobos in a 4-foot radish plant). Nests on the ground may be placed in a tussock of grass, in densely matted perennials such as Labrador tea, or, rarely, in a patch of moss (Grinnell, 1900). They are often built at the base of scrub willows or conifers (Grinnell, 1900; Macoun, 1909; Ray, 1912). Occasionally the nests are placed in exposed situations, as for example the Gambel’s sparrow nest Edward A. Preble (1908) found in a tuft of short grass beside a much-frequented path in a field. Some atypical sites include one reported by Louis Bolander (1906) of a nest built 35 feet above the ground in a cypress, another in the outer drooping branches of a tall acacia tree, and a third in an ivy vine on a building (Blanchard, 1941).

The extremes of height of nests reported in the literature range from flush with the ground to 35 feet above it. The average height above the ground for 31 nuttalli nests at Berkeley was 3.5 feet with a range of 1.5 to 11 feet, for 16 nuttalli nests at Point Lobos 1.8 feet with a range of 1 to 4 feet. H. W. Carriger (pers. comm.) states that he occasionally found nuttalli nests on the ground. The Friday Harbor Puget Sound sparrows tended to nest on the ground with greater frequency than did the Berkeley nuttalli. Of the 45 nests I found at Friday Harbor, 14 were built on the ground in masses of dense scrubby salal (Gaultheria shallon) or in dead bracken fern or grass. I also found nests in native and exotic trees and shrubs. As nest-building started at this locality before the trees and shrubs were fully leafed, many of the nests in process of construction could be seen from a distance of several yards. The foliage grew so rapidly, however, that before the eggs were laid the nests were well concealed.

The eight gambelii nests I found at Mountain Village in 1950 were all built on the ground. When nesting began there, the only vegetation thick enough for concealment was either dead grass recently exposed by the melting snow or dense mats of dwarf perennials. At College, Alaska, in 1957 I found 13 nests on the ground and only one nest in dead twigs a few inches above the sloping ground of a railroad embankment.

The four oriantha nests I found in the Sierras were all on the ground, one at the center of an Artemesia bush, the others under scrub lodgepole pine. Records of oriantha nests in the literature include several on the ground in grass, on slanting willow stems or under scrub conifers, and several a few inches above the ground in spruce or pine. The highest on record is a nest Milton S. Ray (1912) found in the Tahoe region 4 feet above the ground in a lodgepole pine sapling.

In the matter of nest material the birds show great catholicity of taste and use whatever suitable material is available. If the nest is built on the ground, it may lack a platform; if it is placed above the ground it may have a bulky platform of twigs (Bolander, 1906). Materials used in the nest are: fine twigs and rootlets (Barlow, 1901), grasses, both green and dry, dead fern leaves, weed stems, shreds of bark and pine needles (Burleigh, 1930). Materials used for lining include fine grasses, feathers, and whatever mammal hair is available such as deer, cow, horse, and dog (Burleigh, 1930; Grinnell, 1900; Nelson, 1887; Ray, 1912).

In the three races I have seen, nest building is solely by the female. The following description for Nuttall’s sparrow applies to Puget Sound and Gambel’s sparrows as well. Such racial differences as exist involve the length of time the female takes to finish the nest and the interval between fledging the first brood and starting the second nest.

In Nuttall’s sparrow the first hints of nesting are the male’s “sseep” note, which may precede nest building by as much as 8 days, and the interest the female shows in nest material. For a week before she begins to build she may pick up straws and then drop them. During this period she may also utter the characteristic “eep” location note which she subsequently uses during nest building and incubation. She may protest if one approaches the future site of the nest. Several days or a week after these behavior elements have appeared separately, they merge and intensify into coordinated activity, and she begins continuous work on the nest. Her mate perches nearby and utters the “ssseep call note, which may stimulate her to build. The details of a single morning’s work of one Nuttall’s sparrow female are as follows:

On March 11, which was the second or third day she had worked on the nest, I watched her from 8:00 to 11:30 a.m. At 8:15 she carried material to the nest, and from then until 11:30 built almost continuously, making 135 trips with material, stopping only three times and then only for a moment to feed. Almost all the material was gathered within a few yards of the nest; it consisted of dead twigs and leaves, strands of dry grass, pine needles, small green plants, and fresh grass stems. Dead leaves and dry grass were brought most often. She brought only three twigs, all within the first hour. After every few trips she could be heard fluttering in the nest, probably molding the cup. This female spent parts of the next three days in building, but never worked as continuously as on March 11. By March 15 the nest appeared complete.

Nest-building in Nuttall’s sparrow begins a few days before copulation, when trilling and posturing are reaching the climax. The number of days spent in building the first nest of the season was from 7 to 8 or 9 for the five nests found just as the female was starting work. Three were worked on during parts of 7 days, one during parts of 8 days, and one 8 or 9 days. If a clutch or a brood of nestlings is destroyed, the female starts a new nest within a few days and may complete it in 5 or 6 days.

Eggs: The color of the eggs has been described variously as “pale greenish-blue, spotted and splashed with liver-brown” (Davie, 1883) or ”a handsome light green or bluish-green shade, * * * heavily dotted, spotted, blotched or clouded with reddish-brown” (Dawson, 1923) or “ * * * pale greenish blue, varying to brownish, spotted with cinnamon or reddish brown” (Gabrielson and Lincoln, 1959). The first description is for a clutch taken at Alameda, California, and hence nuttalli, the second of one collected in Humboldt County and therefore pugetensis, the third of eggs of Gambel’s sparrow. Davie (1883) comments on a clutch assignable to oriantha collected at Hancock, Colorado:  “ * * * the markings are much heavier and thicker near the larger ends.”

Some egg measurements are as follows: For nuttalli, a clutch of four eggs measured by Davie were .94 x .68 inch, .88 x .68 inch, and two .88 x .64 inch. Dawson gives the average measurements of 70 eggs of pugetensis as 20.7 x 16 millimeters, the extremes being 18.5: 22.4 millimeters by 14.7: 16.5 millimeters. For oriantha W. G. F. Harris gives the average of 40 eggs as 21.1 by 15.8 millimeters, the eggs showing the low extremes measuring 22.9 by 16.8, 19.8 by 16.3, and 20.3 by 14.7 millimeters. For gambelii Harris gives the average for 40 eggs as 21.5 by 15.5 millimeters, the eggs showing the four extremes measuring 24.1 by 17.0, 18.5 by 14.7, and 22.4 by 14.2 millimeters.

The limits of the egg-laying period for the four races are from early March to late July, and the date when the first clutch of the season is laid depends upon race and latitude. The number of eggs per clutch varies from 2 to 6, with two exceptional cases of 7 eggs per clutch. The average clutch size varies with race, latitude, the month in the breeding season the clutch is laid, and the age of the female. The eggs of one clutch are laid one each morning on successive days until the clutch is complete.

The rest of this section presents the data on eggs of Nuttall’s sparrow. The respects in which egg-laying in the other races differ from nuttalli is discussed elsewhere.

The first egg is laid from 1 to 7 days after the nest is completed (an average of 3.6 days for 12 records), and from 4 to 9 days after the first observed copulation. The egg-laying season in nuttalli begins in most years in mid-March and finishes by late June. Extreme dates for clutches are Mar. 3,1936, for a clutch I found at Berkeley, and July 24, 1898, for one M. S. Ray found in the San Francisco Bay Region. Median dates for first egg of first clutch laid at Berkeley for 5 years are as follows: Mar. 22: 23, 1934 for six clutches, Apr. 4, 1935 for six clutches, Mar. 11, 1936 for seven clutches, Apr. 13, 1937 for nine clutches, and Apr. 7, 1938 for five clutches. The contrast in the median dates for the Berkeley population between 1936 and 1937 shows how weather conditions (chiefly mean temperature) can influence the time nesting starts in this race. This marked annual variation for the same locality is characteristic of the resident Nuttall’s sparrows, which must adjust to the capricious winter and spring climate of California. The migratory races that nest in the far north where the country becomes suitable for nesting close to the same date each year do not show such wide annual fluctuations in the date of first egg laid.

The number of eggs per clutch varies from 2 to 5. The average for 215 sets is 3.27 eggs per completed clutch. The most frequent clutch size is 3 eggs. The percentages are as follows: 2 eggs per clutch 4.2 percent; 3 eggs per clutch 65.6 percent, 4 eggs per clutch 29.8 percent, 5 eggs (one clutch only) 0.4 percent. The average number of eggs per clutch increases from March through May, after which it declines: for March the average for 15 clutches was 3.00, for April 3.22 for 122 clutches, for May 3.48 for 50 clutches. In June the average of 21 clutches declined to 3.24, and in July, the average for 7 clutches was only 3.14.

The average for 24 first clutches was 3.04 eggs, whereas 10 second clutches averaged 3.60, 5 third clutches 3.00, and 3 fourth clutches also 3.00. I have records of all the clutches laid by one female for 2 years. In 1934 when she bred for the first time she laid four clutches of 3, 2, 2, and 3 eggs respectively; in 1935 she laid four clutches of 2, 4, 4, and 3 eggs.

Incubation: Incubation is by the female alone. She sits on the eggs with her back flush with or below the nest rim, her tail raised at a sharp angle and her chin resting on the edge. She incubates for periods averaging about 20 minutes each, then leaves the nest to forage, uttering the same strident “eep” note she used while building the nest. Hopping about on the ground hurriedly, she covers a large area in a few minutes: This rapidity of movement is so characteristic that I have often identified an incubating female by this behavior. When the female returns to the nest she again utters a series of “eeps.”

At the start of incubation the male achieves the highest development of territorial behavior. He guards his area by loud, almost continuous singing and patrol. Often, but not invariably, he appears to call the female off the nest. At such times he approaches the nest and sings, whereupon the female leaves and the pair forage together. The male therefore does not guard the nest in the absence of his mate, but he may be involved to some degree in her return. Often he flies to a perch near the nest and sings, whereupon the female gradually moves nearer and nearer and finally resumes incubation. At other times the female is the first to stop foraging and to fly toward the nest, followed by her mate.

In Nuttall’s sparrow the beginning of incubation in relation to egg-laying varies; 5 females began on the day before the last egg was laid, 5 on the day the last egg was laid, and 1 the day after. Of 5 females with sets of three eggs, 3 started on the day the second was laid, 1 on the day the third was laid, and 1 the day after the set was completed. Of 3 females with sets of four eggs, 2 started on the day the third egg was laid, and 1 on the day the set was completed; 3 females with sets of only two eggs all started the day they laid the second.

The average time of hatching for 10 sets of eggs was 12 to 12½ days after the first day of continuous incubation. One set hatched in 11 or 12 days, two in 14 days, the remainder at the average time.

As might be expected from the individual variation in the beginning of incubation, the eggs of one set may hatch all on the same day or on two successive days. The longest intervals between first and last hatchings of eggs in one set were somewhere between 7 and 15 hours for a set of three eggs, and between 16 and 17 hours for a set of four. The eggs in any one set do not necessarily hatch in the order in which they were laid.

Young: The following account of Nuttall’s sparrow applies also to those populations of pugetensis and gambelii I have watched during this phase of the breeding cycle. Where racial differences occur, these are noted under the race in question.

On the day the young hatch, the adults follow essentially the same routine as during incubation. The female now alternates between brooding the young and gathering food for herself and for them. The behavior of the male is definitely affected and reflects awareness of the event. He continues his patrol but spends more time near the nest. He responds at once to the alarm notes of the female by flying straight toward the nest and scolding.

For at least the first 3 or 4 days the female bears almost the whole burden of feeding the nestlings. I have seen her start to gather insects within 2 hours after the young hatched, but have never seen the male visit the nest until the 2nd or 3rd day after, and then relatively rarely. At such times he never went directly to the nest, but carried the food to a nearby perch and held it in his beak for as much as an hour before feeding it to the young.

The 6th day after hatching is the last on which I have found a female brooding her young during the day; only rarely have I found one sitting on the nest in the daytime after her young were 4 days old. One female brooded her nestlings each night until the 8th day after they hatched.

Blanchard (1941) describes the nestling on the day of hatching as follows:

The young bird just after hatching weighs a little over 2 grams. It has down on the head, dorsum, wings and thighs; this dries in about two hours and stands straight out from the body. The remnant of the yolk stalk is still visible, and the viscera can be seen through the transparent skin of the abdomen. The bird breaths spasmodically, the entire body throbbing. It responds to jarring of the nest by raising its head and waving it about unsteadily, the mouth wide open. At first it holds this position only for a moment before its head drops forward and the “egg position” is resumed. I can hear no sound even when the beak is wide open.

The following description of the development of behavior of the nestling is based upon my observations and those of Richard C. Banks (1959): On the day after hatching I could hear the nestling squeak faintly. The earliest the bird can right itself is 2 days after hatching. The eyes begin to open 3 days after hatching and by the 5th day they are wide open. Reflexive grasping of the nest lining by the feet begins about 4 days after hatching. On the 5th day after hatching a new posture, crouching, is assumed. By the 7th day after hatching a large number of nestlings give protest notes when handled. The first concerted escape attempt occurs at 8½ days after hatching.

Banks (1959) describes the growth of the feathers in nestling Nuttall’s sparrows as follows:

At hatching, dark spots representing feather papillae are visible beneath the skin surface on the alar, humeral, dorsal, and coronal tracts of White-crowns. External feather sheaths first appear when the birds are 2½ to 3 days of age. The first rupture of these sheaths occurs at age 5½ or 6 days. At 7 days of age, the tips of the primary sheaths begin to break. Feathers of the anal circlet appear without visible external sheaths.

After an initial slow start, feather growth is rapid and constant. Primaries grew at a maximum rate of 3.9 mm. per day; rectrices grew as much as 9 mm. in 3 days. Although weight increases occurred only during daylight hours, feather growth continued throughout the day.

The young normally leave the nest when about 10 days old. For 23 nestlings the time spent in the nest varied from 9 to 11 days, or an average of 10.1 days. Blanchard (1941) describes the subsequent life of the fledglings as follows:

The first few days after fledging, the young perch in the shrubbery near the nest. They are usually so well concealed and respond so quickly to the warning tit of the parents that it is impossible to see them. The male now shares about equally with his mate the work of feeding the young. When the parents bring food or come near them in foraging the fledglings utter teez and flutter their wings.

By the third to seventh day after fledging, the young of seven broods which I followed had moved to clumps of shrubbery some distance away from the nest. This must have involved crossing several yards of open grass—whether by hopping or flying I do not know. I suspect by the former method, since my earliest records of flight are for birds which had been out of the nest from seven to ten days.

When the young of the first brood are about twenty days old, the male takes over most of the task of feeding them. By this time the female has usually begun to work on her second nest. When from twenty-five to thirty-one days old, the fledglings forage for themselves but still beg food from the parents. The adults continue to feed them a little longer but soon ignore their persistent teez and may even chase or fight them. The oldest fledgling I have seen fed by a female was thirty-two days old, the oldest fed by a male, thirty-five days. Once I saw a female fight with her thirty-five-day-old fledgling; the next year I saw this female chase away another of the same age.

At about this time the young start to wander outside their parents’ territory. At least by the time they are forty-eight days old they leave it forever. None, not even those which spent tin winter only a few hundred yards away, has ever been seen again in its parents’ area.

Food: The food of the Pacific coast races of Zonotrichia leucophrys is known chiefly from stomach content analyses made in the early 1900s by the Bureau of Biological Survey, Washington, D.C. Data were made available through the courtesy of Clarence Cottam, formerly in charge of Food Habits, Division of Wildlife Research, and by John L. Buckley, Director of Patuxent Wildlife Research Center. These and numerous references to food habits in the literature cited here show the catholicity of taste in this species.

Analyses of the stomach contents of birds taken in California and designated as “nuttalli” indicate that the predominant food is vegetable material, mostly seeds. (As most of the data are for specimens collected before 1928, the fall and winter specimens undoubtedly included migratory Puget Sound sparrows as well as resident Nuttall’s sparrows.) For 90 individuals collected, some from every month of the year, the stomach contents of 52, or 58 percent, were reported as containing only dry vegel able matter. Seeds of Amaranthus, Calendrina, Erodium, Polygonum, Stellaria, and various grasses are among those listed. The stomachs of the remaining 38 specimens were classified as follows: 19 had vegetable matter plus insect material making up 10 percent or less of the contents; 13 had only fresh vegetable material (flowers, immature fruit buds, fruit pulp); 3 had partly fresh and partly dry vegetable material but no insects, and 3 had vegetable material plus insect material constituting 11 percent or more of the stomach contents. The small number of specimens with insect matter in the stomachs, only 22 out of 90, or 24 percent, is surprising, especially as most of the specimens (58, or 64 percent) were collected during the nesting season.

References in the literature include many observations on seed eating. Sylvester D. Judd (1898) lists Nuttall’s sparrow as the most important gregarious sparrow that destroys weeds. He reports finding in a Nuttall’s sparrow stomach 300 seeds of amaranth, and in another 300 seeds of lamb’s quarters (Chenopodiurn album). He found the following seeds eaten by Gambel’s sparrows: Polygonum aviculare (knotweed), Alsine media (chickweed), Chaetocloa glauca and C. viridis (pigeon grass). John McB. Robertson (1931) observed Gambel’s sparrows and Puget Sound sparrows eating the small black seeds of the blue gum (Eucalyptus globulus), a tree in which they roost. John & James M. Macoun (1909) record Puget Sound sparrows at Huntingdon, B.C. on Sept. 9, 1901, feeding on thistle seed. Joseph Mallard (1927) also saw Gambel’s sparrows eating the seeds of pigweed (Amaranthus), which they preferred to the bait he used for his traps.

White-crowned sparrows also eat fresh blossoms and leaves. Robert S. Woods (1932) claims that the races of white-crowned sparrow rank next to the linnet in destructiveness to vegetable crops. He states they have a special fondness for young plants of the cabbage family and for young beets and peas. Carrots seem to be immune. Fortunately for vegetable growers, the large winter flocks that forage over the farm lands in California’s inland valleys leave for the north in April.

Unusual items of vegetable matter eaten are reported by several observers. Lyndon L. Hargrave (1939) saw Gambel’s sparrows at Roosevelt Lake, Ariz., feeding daily on exposed seed pulp of fully ripened pomegranates still hanging on the bush. In March 1963 I watched Gambel’s sparrows on the University of California campus at Goleta feeding on ripened olives that had fallen to the ground. Charles G. Danforth (1938) saw a Nuttall’s sparrow fly up and appear to drink the sap overflowing from poplars (Populus nigra), which was extremely sweet and suggested sugar water. Nuttall’s sparrows I collected west of Guadalupe, Calif., in February 1957 had the feathers about the base of the bill bright yellow with willow pollen. At College, Alaska, I saw Gambel’s sparrows eating staminate willow catkins and the new leaf buds of willow, as well as the tiny green stalks of Equisetum and the capsules of mosses.

On several occasions I have seen white-crowned sparrows fly out to catch insects in mid-air. Joseph Mailliard (1919) reports seeing Nuttall’s sparrows take insects by jumping into the air after them. The birds usually catch insects by hopping quickly about on lawns or other grassy areas and either picking up the insects from the grass or catching them as they rise from it. The summer of 1950 I watched Gambel’s sparrows feeding on mosquitoes at the water’s edge on the Lower Yukon at Mountain Village, Alaska. To get an approximate idea of the number of pairs with young, one had only to move slowly up the Yukon in a small boat close to shore and count the pairs gathering mosquitoes and other insects by the water’s edge. Even those Gambel’s sparrows whose territories did not abut on the river flew down there to forage. W. J. Maher (1959) observed young and adult Gambel’s sparrows on the Upper Kaolak River in northern Alaska eating bread scraps and foraging actively for mosquitoes near the base of his tent. They were the only species of bird that fed about the tent.

Stomach analyses of 30 Nuttall’s sparrows taken from September through February reveal a complete absence of insect material. In March and April 17 out of 55 birds had some insect matter in their stomachs. A wide variety of insects has been recorded: “Hymenoptera,” ants, caterpillars, beetles, and weevils. Joseph Grinnell and Tracy I. Storer (1924) state that an adult male mountain white-crowned sparrow taken at Tenaya Lake on July 3, 1915, had nothing but beetles in its stomach. G. F. Knowlton (pers. comm.) collected a female Gambel’s sparrow whose stomach was “well filled with five mud-dauber wasps, two ants, six additional Hymenoptera and insect fragments.”

Data on stomach contents of migratory white-crowned sparrows indicate that a shift to insects in the diet may start on the wintering grounds. Of 80 specimens taken in January, February, and March, only 12, or 15 percent, had any insect matter in the stomach, and in only one bird did the insect matter comprise more than 10 percent of the contents. In April 7 out of 21, or 33 percent of birds taken on the wintering grounds had insect matter in the stomach, and in all seven it made up more than 10 percent of the stomach contents. Of 12 Gambel’s sparrows taken in migration in April on Saturnia Island, B.C., 11 had stomachs containing between 15 percent and 100 percent insect material, with the average percentage of insect matter totalling 73.6 percent. The 12th specimen of this group had only green vegetable matter in its stomach. Thus the shift to insects as a major component of the diet may occur a month or more before nesting begins.

Clarence Cottam (pers. comm.) makes the general statement for all species of birds whose stomachs have been analyzed by the Biological Survey, that races of the same species show little difference in their choice of food unless they occupy distinctly different habitats. The data available on food habits of white-crowned sparrows seem to substantiate this statement, although no detailed study of the food of races occupying the same area in winter has been made. As individuals of two or more races commonly forage together, it seems unlikely that any racial differences in food preference exist.

Voice: The call notes of the Pacific Coast races are identical, and are used under identical circumstances. The location note, “eep,” is used by members of winter flocks and by mated pairs as they forage, move through shrubbery, or fly from one place to another. The same note, uttered more stridently, is given by the female when building her nest and when she leaves or returns to her nest during incubation. A modification of this note, which sounds like “ssseep,” has already been described as the note uttered by the male prior to and during nest-building. A scolding note, which to my ears sounds like “ip” but which William L. Dawson (1923) writes as “zink” or “dzink,” is used to protest intrusion by a person or animal. It is also uttered when there is no obvious cause for alarm or excitement. Lone migrants frequently utter this note over and over again when they arrive on the wintering grounds in early fall. Flock members utter it as they go to roost when the note may be repeated with ever-increasing frequency, followed by a burst of song. A squabbling note that defies syllabification may be used by two birds as they dispute a morsel of food, or by a lone bird as it forages. One other call note is more restricted in its use: a warning “tit” given by parents with nestlings or young fledglings makes the young stop their food cries and stay quiet. As this note is not used until after the young hatch, one can be sure that a pair using it has young rather than eggs.

The habit of this species of singing at night is so characteristic that it is one of the best ways to identify the bird to the layman. Song in Nuttall’s sparrow may express by itself or accompany other expressions of at least six unmistakable purposes or attitudes of mind: defiance or warning to territorial rivals, the search for a mate, sexual excitement, announcement of territorial boundaries (as when used on patrol during the incubation period), interest in the female’s return to her eggs, and fright or physical shock. Examples of the first five have been given. I have two examples of the sixth (fright or shock). In the first, a male foraging with its mate dove into a clump of juniper; when I suddenly shook the bush into which he had disappeared, he burst into song. The second is furnished by Thomas L. Rodgers (pers. comm.). While transporting Nuttall sparrows in darkened cages by car through several miles of city traffic, he noticed that each time he put on the brakes suddenly, the sparrows would just as suddenly burst into song.

The song of the white-crowned sparrow is best described as a theme common to the species, with an almost infinite number of variations: racial, populational, and individual. The musical elements common to the song of the Pacific Coast races consist of one or two initial notes held for nearly half a second each, followed by two or more rapidly sung notes or by a trill or both. On this simple theme are built a wide variety of melodies, from robust simple tunes like those of Berkeley Nuttall’s sparrows to the more delicate and complex melodies of Friday Harbor Puget Sound sparrows. So similar is the quality of voice in all members of the species that a white-crowned sparrow song, once learned, is unmistakable, no matter what the dialect. The racial and populational variations are like different melodies sung by the same voice. The individual variations are like the same melody sung with different accents.

Nuttall’s sparrows and mountain white-crowned sparrows use rhythmic patterns closely related to each other. Gambel’s sparrow uses a markedly different pattern described here in detail, and the Puget Sound sparrow uses patterns reminiscent of the more complex Nuttall’s sparrow patterns, but pitched higher. Some of the racial patterns are diagrammed in Peterson (1941); and Blanchard (1941) shows diagrams for populational variations in Nuttall’s sparrow and the Puget Sound sparrow.

Anne H. Wing, a trained musician with a keen sense of pitch, studied the song of Gambel’s sparrows at Johnson’s Crossing, Yukon Territory, in June and July of 1948 (in a Report on a Study of Arctic Birds and Mammals, by Leonard Wing, 1948, manuscript for the Research Report of the Arctic Institute of North America). Her comments and interpretations are those of a musician rather than a biologist, and I include them along with her excellent musical description for, although I do not necessarily agree with all of them, they represent a fresh and provocative approach to the study of song in this species. Mrs. Wing (pers. comm., 1960) writes, in part:

I found that the Gambel’s sparrow sang melodious, high-pitched songs of tour beats in rapid march time. Stepping upward through the first three beats, downward with the fourth, these decidedly rhythmic songs were clearly whistled on the first two beats, burred on the third and fourth beats. Each song encompassed a range of pitch no greater than a “perfect fifth” interval, e.g., C to G (do-sol).

In its simplest form, the song contained four single notes. Any of the first three beats, and most often the third, might contain not one note but two or three shorter notes on the same pitch. The first beat, whether it contained one note only or two or three notes of shorter duration, was on one pitch. Occasionally in the third beat the short notes were successively a half-step higher. The fourth beat was always a single note, and this was lower in pitch than the last and highest pitch of the third beat. To put it simply, the melody marched quickly up a little musical hill, then stepped down.

In terms of the beats of the song, four beats of song were followed by ten beets of rest or silence. During the rest period, another bird could and often did fit its songs so as to achieve antiphony. Duetting was always in good musical taste, both in rhythm and in melody. If one altered its song in detail, the other was likely to do so as well, keeping the two songs in harmony.

Responsive singing took place most commonly in the hours of singing at dawn, when many birds seemed to congregate at different locations for the purpose of singing. One of my favorite listening places was the shore of the Teslin River. Wakened by the singing soon after midnight or at least by half-past twelve, I would walk down the hank, notebook in hand. Sometimes several birds would be singing the same tune in different keys, the songs overlapping in great confusion. This would soon resolve, perhaps as more birds arrived, into question and answer singing. Around three or three-thirty most of the birds had dropped out, their voices heard, perhaps, on the hillside where there were nests.

Musical replies to musical questions were definitely melodious by the highest standards of human musicians. The duets might be arbitrarily classified as, first, variations on a theme, and, second, mutually complementary songs, one the more obviously a completion of the other. To illustrate, (using letter names of scale tones and recalling that the songs rose through three beats, falling on the last) the singing of the song ‘B flat, C, D—D, B flat’ in alternation with tbe song ‘D, E flat, E—F, B flat’ settles nothing, and Is an endless musical argument variations on a theme. But the singing of the song ‘G, C—C—C, D—D, B’ and Its response ‘B, D, E flat—E natural, C’ completes a musical sentence ending with a cadence (a return to the tonic). Even so, the birds go on singing it over and over again, a characteristic of birds but not of human musicians.

Twilight singing, especially morning twilight, with its greater proportion of duets and of unusual versions of the songs, was very interesting. If any originality of melody or any creative singing was possible, I believe that it occurred then. But if the birds were drawing upon a set of traditional tunes, they made aesthetic choices of rhythmic and melodic details within their songs, particularly when two birds were singing together in antiphonal duet.

Behavior: As all the sections under “Habits” fall properly under the heading of “Behavior” in the broadest sense of the term, only a. few mannerisms that distinguish white-crowned sparrows from other species likely to be found in the same habitat are mentioned here. One of the habits of winter flocks of white-crowns is to forage on the ground by roadsides. When disturbed, they fly or hop up and plunge over the embankment into the nearest cover. If the flock is large, the individuals may move almost as one bird. The essential feature of this escape reaction is the force with which they make a dash for the nearest cover. So characteristic is this behavior that one can use it as a diagnostic field character when traveling at high speed. Grinnell and Storer (1924) describe the ensuing behavior as follows: “If a flock of white-crowns is come upon while it is foraging on the ground, the birds get up quickly and dart into the shelter of some nearby thicket, each pursuing a separate course. There they remain for a short time, silent and motionless, but peering furtively at the intruder. After a short period of quiet, if there be no further cause for fright, they become active again, giving voice to faint seeps and, individually, they begin to hop up in the brush where they can see about before venturing into the open again.

Ralph Hoffman (1927) describes another trait of winter flocks: “When individuals of the flock disagree about a resting place they utter little confused squabbling notes, and just at dusk, when birds settle in thick trees for the night, and at dawn, they repeat for several minutes the alarm note, a metallic pink.”

Field marks: Adult white-crowned sparrows are distinguished by the high, puffy crown, broadly striped with black and white, and by the clear pearly-gray breast. The sexes are alike, but females usually look somewhat duller, that is, the contrast between black and white stripes is not so striking as in the male. Immature birds are buffier than adults, with head stripings of dark red-brown and light buffybrown instead of black and white. Ralph Hoffman (1927) writes:

“No other bird can be confused with an adult sparrow of the White-crowned group, if the markings of the head are seen. Immature birds might be mistaken by a beginner for several other birds with reddish brown on the crown. * * * When there are adults with the immature birds, the resemblance in the pattern of the head markings is obvious.” Hoffman mentions the rufous-crowned and chipping sparrows as species with which immature white-crowned sparrows could be confused. The rufous-crowned sparrow lacks the pale stripe through the center of the crown, and the reddish crown of the chipping sparrow is set off sharply from the sides of the head by a white or whitish stripe and a black line below it.”

Although one can easily identify the species, one cannot always distinguish between races of white-crowned sparrow on the basis of morphological characters alone. With practice one can, however, assign most individuals to one of two groups: nuttalli-pugetensis or gambelii-oriantha-leucophrys. For adults, the differentiating characters are the hue of the feathers on back and rump and the color of the bill. If an adult has brownish back and rump feathers and a yellowish b~, it belongs to the nuttalli-pugetensis group. If it has back feathers with reddish centers and grayish margins, grayish rump feathers and a cinnamon brown or reddish bill, it belongs to the gambelii-oriantha-leucophrys group. To distinguish between immatures of these two groups, bill color, but not color of back or rump feathers, is a reliable criterion. A different grouping, based on the color of the lores in the adult, places those with white or grayish lores in one group (nuttalli, pugetensis and gambelii) and those with a black lores (oriantha and leucophrys) in another. This distinction does not hold in all cases, however. Walter E. C. Todd (1953) reports “white-browed individuals” from the offshore islands and mainland of Hudson and James Bays in the midst of the leucophrys breeding range.

To distinguish between the races within each group is still more difficult. Nuttalli and pugetensis cannot be told apart by morphological characters alone. So similar are the members of these two races that separation requires comparison with large series of study skins, and then only the individuals at the extremes of the range of variation can be identified with certainty. Song pattern, on the other hand, is a reliable criterion for distinguishing between the two races. In localities where both winter together, the best way to distinguish the song patterns is to follow the advice of Peterson (1941) and “learn the song pattern of the local breeding birds thoroughly, and when the migrants* * *  arrive * * * proceed to memorize carefully their respective song patterns.” In the San Francisco Bay Region this system works particularly well, for the local populations of Nuttall’s sparrows sing short simple songs quite different from the longer and more embellished ones of wintering pugetensis. At Carmel and Guadalupe, Calif. the distinction is more difficult to make, for the Nuttall’s sparrows use a more complex pattern. With practice, however, one can use the same system here too. Behavior of the adults also gives valuable clues. Nuttall’s sparrows tend to stay paired on their territories, are bold, and sing with considerable force in fall and winter. Puget Sound sparrows winter in large flocks, are more easily frightened, and sing with less force. In short, the adult Nuttall’s sparrows behave like the permanent landowners they are, whereas the Puget Sound sparrows behave like visitors without attachment to a specific piece of ground. Distinctions of behavior do not apply to the immatures, for those of both races flock in fall and winter. With experience, however, one can usually distinguish between the song fragments uttered by Nuttall’s and Puget Sound young of the year.

Trained observers can make the previous distinctions at close range and in a good light even with the naked eye. An additional character can be used with trapped birds: the color of the bend of the wing. It is bright yellow in adults and immatures of nuttalli-pugetensis, and whitish or only faintly yellowish in gambelii-oriantha-leucophrys. The point to emphasize is that whenever feasible, the individual should be examined for all these characters. In most cases the yellow bill and bend of wing and the brown back and rump feathers will occur together; likewise the cinnamon-brown bill and the reddish and gray back feathers and grayish rump will coexist in the same individual. In exceptional cases, however, one of these characters may be combined with those of the other group. I trapped one adult in early spring at Berkeley that had the bright yellow wing bend characteristic of nuttalli-pugetensis and the cinnamon-brown bill and reddish and gray back feathers and grayish rump of the second group.

As has been said, lore color can be used to separate typical Gambel’s sparrows from the two other races of the second group, but the difference does not hold for all individuals. A number of museum specimens I have examined have loral areas partly white and partly black. Obviously the race to which such individuals belong cannot be determined by this character. As with nuttalli and pugetensis, the morphological distinctions between oriantha and leucophrys are slight, and identification requires comparison with large series of museum skins. Here again song pattern may be a more reliable criterion where two or three races winter together. I have no first-hand knowledge of the song patterns of leucophrys, but Otto Widmann (1911) states: “To one accustomed to the song of the species in the East the song of this Rocky Mountain bird is a great surprise, for it has no resemblance at all, only one note at the beginning to the monotonous ditty reminding one of the much more powerful and melodious song which we hear every May in the Mississippi Valley.”

To my ear the song patterns of the California populations of mountain white-crowned sparrow are quite distinct from those of Gambel’s sparrows wintering in California or breeding in the parts of Alaska where I have studied them. This correlates with Anne H. Wing’s statement (pers. comm.): “The fact that the range of individual Gambel’s sparrow songs heard at Johnson’s Crossing, Yukon Territory, did not exceed a major fifth interval is interesting in the light of the fact that similar songs heard from white-crowned sparrows in the Rocky Mountain National Park (oriantha) containing the same note-names often ranged through more than an octave.”

As one illustration of how color characters normally confined to one group of races may crop up in the other, Austin L. Rand (1948) describes an adult female collected in fall migration at Shoal Lake in southern Manitoba that had a pink bill, pale gray breast and belly, white bend of the wing and black lores combined with an olive brown rump, olive-toned edgings to the feathers of the upper parts, and heavily pigmented flanks. The black lores is referable to Z. l. leucophrys, the general color of the upper-parts and flanks is “very similar to those of pugetensi.s from the British Columbia coast,” and the bill color and bend of wing is referable to either Z. l. leucophrys or Z. l. gambelii.

To sum up, racial differences in morphology are slight, and in many cases cannot be used as sole criteria to differentiate between races. Differences in behavior and song pattern are more marked, and therefore whenever possible should be used in addition to the morphological characters. Physiological differences, such as extent of prenuptial molt, amount of subcutaneous fat, and date of recrudescence of the gonads, are even more reliable criteria, but only in spring. Discussion of these lies outside the scope of this article. The locality of observation or collection, while important, gives only a partial clue to the identity of a given individual, for the winter ranges of several races overlap, and the breeding ranges of some are contiguous. Therefore the more characters one can scrutinize: morphologic, behavioristic, physiologic: the more reliable will be the identification. Intermediate individuals should be given no racial designation.

While the vagueness of the differentiating characters and their fluidity of distribution may be disconcerting to persons accustomed to less plastic species, they are the most fascinating properties of this interesting bird. The lack of trenchant racial differences and the abundance of intermediate individuals indicates the close relationship of all members of the species and, presumably, the recency of the evolutionary development of the present races.

Enemies: I interpret the term “enemies” in its broadest sense, to include both native and introduced predators, parasites, and man in those rare cases where death of white-crowned sparrows results from Ins activities.

There is direct evidence that snakes, hawks, and owls prey on white-crowned sparrows. I have indirect evidence that shrikes, jays, and crows are also enemies. Grinnell and Linsdale (1936) found a 4-foot gopher snake in a bush with a nest of Nuttall’s sparrows and almost touching one of the two remaining young birds. The same authors state that at Point Lobos Reserve sharp-shinned hawks make frequent captures. Their account of one such capture states that a hawk appeared from the pine woods, dashed down into the radish patch where white-crowned sparrows were feeding, and captured a bird apparently of this species. Winsor M. Tyler (1923) observed prairie falcons in the San Joaquin Valley and in the arid hills along the western rim of this valley. “From the time the falcons return to their nest cliffs in early spring through egglaying and incubation periods the Gambel sparrows (Zonotrichia leucophrys gambelii) are very abundant in the regions where falcons abound and a very heavy toll of these sparrows is taken.” F. C. Evans and J. T. Emlen, Jr. (1947) found the remains of crowned sparrows (Z. leucophrys and/or Z. coronata) in barn owl pellets at Davis, Calif. Albert C. Hawbecker (1945) found six white-crowned sparrow remains in barn owl pellets at Struve Ranch near Watsonville, Calif. Herbert Brandt (1951) mentions the northwestern shrike as being a predator on Gambel’s sparrows, and I have occasionally found a shrike perched on a trap where I was banding sparrows. I have seen jays and crows watch me intently as I made trips to nests. At Friday Harbor crows appeared to be chiefly to blame for destruction of the nests I was watching that did not last through to the fledging of young.

At Mountain Village weasels would undoubtedly have constituted a danger to the ground-nesting Gambel’s sparrows had not Eskimo trapping kept their numbers to a minimum. In the 3 months I spent at Mountain Village in 1950 I saw a weasel only once. I watched it walk boldly along a path in full view to a point where two male Gambel’s sparrows were perched near the ground facing each other at a territorial boundary. They were preoccupied with each other and hence unaware of the weasel, which gazed intently at them and stayed ready to spring as long as the birds remained near enough to the ground to be almost within striking distance. As soon as the birds flew off the weasel disappeared. At this locality jaegers that flew over the tundra above the village seemed to watch me as I looked for nests. As most of the Gambel’s sparrows nested close to the village where the jaegers came only rarely, these probably did not constitute a great threat. At College most of the Gambel’s sparrow nests I found were at the edges of farmland, and I never saw an actual raid by a predator. I saw red foxes nearby, however, and noticed the strong scent of fox near one Gambel’s sparrow nest from which nestlings disappeared several days before they were due to be fledged.

As white-crowned sparrows live near human habitation, cats must be considered as enemies. They often interfere with banding operations by developing the habit of visiting the traps. One such case is interesting because of the reaction of the victim. As I was inspecting my traps I came upon a cat crouched beside one of them. It had badly mauled a Gambel’s sparrow inside the trap. The bird was bleeding, some of its feathers were scattered on the ground, and it gave every sign of terror. I chased tho cat away, banded and released the bird, and reset the trap. Less than 20 minutes later the same bird reentered the trap.

There are two records of white-crowned sparrow nests parasitized by cowbirds. Henry J. Rust (1917) found a nest with five eggs, including one egg of a cowbird, along Little Dry Creek in Fremont County, Idaho. Friedmann (1938) describes a nest in the Royal Ontario Museum at Toronto collected at Okotoks, Alberta, that contains one white-crowned sparrow egg and two eggs of the cowbird.

I have frequently found nematode worms in the body cavity of sparrows collected in or near Santa Barbara. Rarely I have found parasitic flies of the genus Ornithoica clinging to the feathers of trapped birds. Otto E. Plath (1919) found 36 full-grown larvae of Protocalliphora azurea (Fallen), (now placed in the genus Phormia) in the nest of a Nuttall’s sparrow in the San Francisco Bay Region. They were determined to be sucking the blood of nestlings; some died as the result, others were retarded in growth. C. M. Herman, H. A. Jankiewicz and R. W. Saarni (1942) found protozoan parasites of the genus Isospora in one Gambel’s sparrow.

In the category of accidents should be mentioned the occasional sparrow that kills itself by flying against a window. Alfred B. Howell (1914) reports seven Gambel’s sparrows destroyed in an orange grove in Covina, Calif., when the trees were fumigated with hydrocyanic acid gas. Frank S. Daggett (1902) reported three Gambel’s sparrows found dead on the surface of cyanide tanks at a gold mining camp in the Cargo Murchacho Mountains west of Yuma on the Colorado desert. On the whole, however, the effects of man’s activities are usually beneficial to white-crowned sparrows. Roadbuilding and clearing allows them to extend their range into areas formerly too densely wooded for them.

The mortality rates for nestlings I followed are highest for Berkeley Nuttall’s sparrows, lower for Friday Harbor Puget Sound sparrows, and lowest for Alaska Gambel’s sparrows. Of 30 broods of Nuttall’s sparrows I watched, only 12, or 40 percent, were successfully fledged.

Of 32 broods of Puget Sound sparrows followed in 1936, 19, or 59 percent were fledged. Of 8 broods of Gambel’s sparrows at Mountain Village and 12 broods at College, 6 and 9 or 75 percent were successfully fledged. This suggests the interesting possibility that the northern breeding grounds may actually be safer than coastal California for raising young. The very fact that Gambel’s sparrows maintain their numbers by raising only one brood is also indicative of the relatively lower nestling mortality.

Fall and Winter: An arbitrary division between fall and winter does not accord either with the climate of coastal California or with the phases of the annual cycle in Nuttall’s sparrow. A more meaningful designation of the time between breeding seasons would be the “base level,” when territorial and sexual behavior decline to a low level but do not quite disappear. This period begins with the postnuptial molt in late July or early August and ends with the surge of territorial and sexual activities the following January.

The quotations that follow from Blanchard (1941) describe the behavior of Nuttall’s sparrows on the University of California campus at Berkeley, Calif.

My banded pairs (of adults) remained on their breeding areas through the fall and winter. Mates foraged and perched together and followed each other about, uttering the eep which serves as the location note. Both sexes sang sporadically, and an occasional territorial dispute involved forceful singing and chasing. The instinct to patrol the area was absent or nearly so, however, and several of the pairs were joined by flocks of immatures and, rarely, by mateless adults. These newcomers were treated with such complete tolerance that only the most careful observation could detect a certain aloofness as well as a certain dominance in the established pair.

When the adults begin their postnuptial molt in late July and early August, they become so secretive and silent that it is nearly impossible to find even those banded birds whose forage routes one knows well. Trilling and posturing, chasing and fighting cease. Singing usually stops also, although I have one record of a loud and complete song from a heavily molting male.

Even though external signs of territorialism are suspended during the molt, a definite interest in the breeding area must persist in both male and female, as the following accounts show:

On August 14, 1935, I trapped an adult male about a mile north of the campus. Its tail was barely an inch long and its wings were still in the process of molting. I put the bird in a darkened cage and took it by car to the campus, where I banded and released it in an area at that time unoccupied by Nuttalls; the area was later taken over by a pair and therefore must have been suitable. On August 31 I found the male back on the spot where I had trapped it. Three days later I retrapped it and found that the tail had not yet reached full length and the feathers of the pileum were still in sheaths.

On September 3 of the same year I tried an identical experiment with an adult female which was just starting to molt. I trapped and released her at the same points as I had the male, and nine days later found her back where I bad trapped her. By this time she was molting heavily on the wings, breast, and belly. Her tail feathers bad been renewed but were just beginning to grow out.

The significant point of these experiments is not that the birds returned to their breeding areas, but that they did so at a time when activity is normally reduced to the minimum and when the pairs into whose area they may have wandered would almost certainly have made no effort to drive them out.

Banding data for the young birds indicate that during most of fall and winter they too are probably limited to a restricted area, within a few hundred yards of which they attempt to establish territories the next spring. Such wandering as they do probably occurs before they undergo the postjuvenile molt. Depending on when they are fledged, therefore, the period of wandering could last from May through most of August for young of a first brood, or about a month for young fledged in July.

On August 26, 1034, I trapped within 400 yards of its birthplace, a young bird beginning its postjuvenile molt; it had been banded as a nestling the previous May. Through the following winter this bird (male xi) stayed with a flock of unbanded immatures within an area of 1.7 acres which included the spot where I trapped him and at the edge of which he settled the next spring. In October and November of 1935 I handed three immatures, two of which stayed near by the rest of the winter and bred the next spring within 50 yards of where trapped. One of these was retaken the following July less than 200 yards away. These histories, together with the fact that I had seen groups of immatures foraging day after day in the same spots, made me suspect that they, like the adults, might limit themselves to small areas in fall and winter.

To substantiate this, I color-banded fourteen immatures in August and September of 1936. All but one were seen again that winter: twelve, from one to fifteen times within 200 yards of where trapped, and one, once, within 400 yards. Eight settled the next spring on territories from 100 to 400 yards from the trap sites. None was ever found farther than 400 yards away.

* * * The young birds begin to sing, though rarely, in mid-July, while still in juvenile plumage. This song may take the form of either a weak or abbreviated version of the immature song or a prolonged trill. As in most passerine birds, the immature song is much more variable than that of the adult. The young bird sings weakly, hesitatingly, often repeating notes of the first phrase before uttering the trill, which is sung so slowly that the separate notes can he heard. Every conceivable variation is used: the first three notes may be sung and the trill omitted, or one or more of the first three notes may be left out, and there are innumerable variations in the trill itself.

By mid-September —when most individuals have finished the molt —song and other evidences of renewed territorial interest may recur. One male I watched in 1936 on the Berkeley campus sang from a conspicuous perch in early September and throughout October. Twice in the first week of October he sang loudly every few seconds; his song was indistinguishable in force and frequency from that of breeding time. On rare occasions I even found two birds perched conspicuously, alternating in loud singing. Although females do not usually show signs of territorial jealousy, I have one case of a banded adult female which not only sang forcefully but engaged in a dispute with another adult. The immediate cause seemed to be the desire of both birds to feed on the same pyracantha berries.

During early September female III had frequented a certain bush on the edge of her territory until, we may imagine, habit verged upon necessity and the spot was unseasonably invested with the power to stimulate possessive jealousy. On September 9 and 15 I saw her sing there several times. Her song was complete but weaker than that of her mate. On the morning of September 17 I found her there on the ground, singing as forcefully as any male in breeding time. Then an unbanded adult in the same patch of shrubbery sang, somewhat less vigorously. This was male VIII, the neighbor on the east. They sang at one another for four or five minutes, shifting over the ground about six feet apart. Then the male chased female III around the edge of the shrubbery. She lit on the ground and both sang again. Another chase took place, then more singing. The next morning this was all repeated, but female III sang less forcefully. The same afternoon I found both feeding silently on the same berries two feet apart, and so for the next four mornings. Both sang occasionally, but I saw no more pursuits.

Territorial and sexual displays by adults in the fall are known for many species of birds (Marshall 1952).

“Pursuit is common among immatures in fall and winter. It is usually momentary, involving flock mates which may resume peaceful foraging a minute or so later. Sometimes, however, pursuit has been accompanied by loud singing and fighting, suggestive of a territorial dispute.”

The data cited point to ” * * * a more or less passive limitation of pairs to their breeding areas in winter, with almost complete tolerance of sojourning strangers and only rare flare-ups of jealousy between established neighbors. Restriction in winter is obviously not the product of conflicts between neighboring pairs; chasing, fighting, and patrol are almost completely absent. The attitude of the male toward immature and mateless birds gives us little reason to suppose that he would resent invasions of adjoining pairs if such occurred.”

Banding: White-crowned sparrows live close to man, trap easily, and forage and perch with the lower part of the leg exposed so that color bands are plainly visible. Many banded birds re-enter the same traps again and again. Each year thousands of white-crowned sparrows are banded, and hundreds previously banded are recaptured. In the April issue of the “Western Bird-Bander,” the Western Bird-Banding Association publishes annually the numbers of birds reported banded by its members in western North America. In 1959, 6,616 white-crowned sparrows were banded. This is 5.9 percent of all birds reported banded that year, and is the largest total for any non-game species. Corresponding figures for 1960 are 5,338, or 4.3 percent of all birds banded, and for 1961, 5,418 or 4.2 percent of all birds banded. The number of banders reporting to this Association increased from 167 in 1959 to 185 in 1961 (Stoner et al, 1960, 1961 and 1962).

Because so many white-crowned sparrows are trapped each year, the data from banding would fill a large volume. No attempt is made here to summarize all the facts. Instead, I have chosen two long term studies to discuss in detail: one, a project conducted for 18 years under conditions as stable and as close to primitive as are ever likely to be achieved today in California, and the other, an experimental study begun in 1961 and still in progress as this article goes to press. Both projects illustrate the unique contributions of banding to our knowledge of Zonotrichia leucophrys.

The first is a study of survival of banded birds conducted by Jean and Mary Linsdale and co-workers in the winter seasons of 1937 through 1955 at the Hastings Reservation in the Santa Lucia Mountains, Monterey County, Calif. During the 18 seasons the study was made, the Linsdales banded [3,366 individuals of 46 species of native birds. Of these, 2,299 were wintering white-crowned sparrows, probably chiefly pugetensis, although the race is not specified. The results of the study are reported by Linsdale (1949) and by Linsdale and Linsdale (1956). Of special interest are the age profiles for each year for the population (Table 4, p. 94 of Linsdale, 1949). In 1942-43, for example, a total of 189 individuals were trapped. Of these, 113 were birds with immature plumage which were therefore less than 1 year old, and 16 were adults trapped for the first time. The rest were recaptures of birds banded in previous years; 51 were banded as immatures and hence their approximate age was known. Of these, 25 were between 1 and 2 years old, 9 were between 2 and 3 years old, 8 were between 3 and 4 years old, 7 were between 4 and 5 years old, and 2 were between 5 a.nd 6 years old. The remaining 9 of the 189 birds trapped that season were banded as adults, and 2 of them were at least 6 to 7 years old.

Table 4 also gives data on the yearly decline in percentage of recaptures for a given batch of birds banded as immatures. Of the 122 birds banded as immatures in 1941-42, 25 or 20.5 percent were re-trapped in 1942-43; 10 of these, or 8.2 percent of the original 122 birds, were recaptured in 1943-44, 6, or 4.9 percent returned in 1944-45, 5, or 4.1 percent were taken in 1945-46, and 2, or 1.6 percent of the original total survived to re-enter the traps in 1946-47. As would be expected, the corresponding percentages of survivors are not quite the same if a different year is used as the starting point, but all the curves are similar in showing that the heaviest drop-off in captures of individuals surviving to their first winter comes between then and the next winter season. From then on, the decline in retrapped survivors is much more gradual.

The Linsdales’ study reveals wide annual variations in the ratio of immatures to adults trapped. In 5 out of 10 seasons the ratio of immatures to adults was about 1 to 1. As a corollary, the average percentage of immatures per year for the 10-year period from 1938-39 through 1947-48 was 52.6. Yet in 1943-44, the ratio was about one immature to three adults, whereas the next season the ratio was reversed. This indicates the false impression that can be gained from sampling of atypical years.

The work just discussed is one of the most exhaustive banding operations involving white-crowned sparrows ever carried out. Yet the investigators state:

In reviewing these observations, attention is directed to the small size of the area concerned in these winter studies. A long strip of 20 acres would include every trapping station as well as the ranges on the Reservation of nearly all the individual birds concerned. When this is compared with the long line of travel, over a thousand miles for the migrant species, the remarkable effectiveness of the controls over the birds which come and stay and return repeatedly between the two homes is given special emphasis. The rigidity of these controls is further demonstrated when we realize how small a part of the whole population of each species is represented on our minute area. If 4,000 crowned sparrows [here Linsdale refers to both golden-crowned and white-crowned sparrows] have wintered on our 20 acres in the last 11 years, how many have come to the 100 million acres of California? How was each of them able to find and stay in its particular home area? Before we make guesses pertaining to these questions we need to know more about what the birds really do.

One answer to some of these questions may be forthcoming from an experimental study of homing in Zonotrichia leucophrys and atricapilla begun in 1961 at San Jose, Calif., by Richard Mewaldt and his students. Mewaldt is attempting to find out, not what the birds do under natural conditions, but what they can do under extraordinary conditions they would not meet in nature (Mewaldt, 1962a, 1962b, and 1963; Roadcap, 1962). The results to date on displacement of white-crowned sparrows are briefly summarized here. He and his co-workers displaced 65 white-crowned sparrows of the races pugetensis and gambelii distances of between 9 and 164 miles from the banding station at San Jose. The most significant return was made from Visalia, Calif., 164 miles away, by a first-year pugetensis in only 7% days. This bird had to cross the interior coast ranges to make its return.

Experiments with releases at much greater distances from the banding station are in progress. During the 1961-62 winter season, Mewaldt shipped 233 Puget Sound and 79 Gambel’s sparrows by commercial aircraft from San Jose to Baton Rouge, La.; 67 of the displaced pugetensis and 2 of the gambelii individuals were proved “homers,” for they had already been trapped at San Jose in at least 2 successive winter seasons. Releases at Baton Rouge were made on Oct. 28 and Dec. 5, 1961, and on Feb. 2 and Apr. 14, 1962. On June 24, 1962, one Puget Sound sparrow of the April 14 release was trapped at the banding station at San Jose. This individual, a male, had been banded as aii immature at San Jose in March of 1957, and recaptured each season for the five intervening winters until he was shipped to Baton Rouge in 1961, so he was 6 years old when he made the trip back from Louisiana.

This was the first Zonotrichia to appear in June at the station in its eight summers of operation * * * He was not seen after 24 June until again retrapped on 27 October, 1962. * ** Because be appeared to be in migratory condition in June, and because he was not detected between 24 June and 27 October, I suspect he spent July and August in his nesting territory somewhere in the Pacific Northwest.

It is most reasonable to assume that this bird returned directly to San Jose, his winter home, from the release area at Baton Rouge, Louisiana, an airline distance of approximately 1,800 miles. The effective rate of return was at more than 25 miles per day over terrain not frequented by the race pugetensis. It seems inescable that this sparrow “homed” to his winter range from a remote and unknown release point. Bi-coordinate navigation of a very effective nature seems possessed by this bird. * * * As of 31 December 1962, 21 of the Baton Rouge releases have been retrapped at the banding station in San Jose. Although the numbers are not large, certain trends are appearing in the data. Thus far 7 pugetensis, 8 gambelii and 6 atricapilla have returned. It is perhaps significant that a greater percent (10%) of the strongly migratory race gambeiji have returned than of either pugetensis (3%) or atricapilla (6%). Also of importance is the observation that 8% of adults have returned compared to but 3% of birds less than one year old at the time of displacement.

A brief review of earlier experiments with displacement of Zonotrichia winter visitants in California is given in R. Roadcap (1962).

Other important data contributed by banding are recoveries in the north of white-crowns banded on the wintering grounds, longevity records, and descriptions of the rate and sequence of prenuptial molt. In spite of the thousands of white-crowned sparrows banded each year, records of recoveries on the migration route and within the breeding range are still rare. Especially valuable, therefore, are the recoveries of Puget Sound sparrows mentioned by Mewaldt (1962b). In a personal communication he furnishes the following details of these recaptures:

25-11892 banded Nov. 23, 1957 as Im at San Jose; returned Oct. 25, 1958; returned Oct. 9, 1959; recovered Apr. 6, 1960 at Blame, Wash. by Mrs.

Lucille Kline . . . returned Oct. 15, 1960 to San Jose; returned Dec. 2, 1961.

22-136419 banded Aug. 26, 1959 at Vancouver, B.C., taken at San Jose Dec. 31, 1961.

27-169514 banded Sept. 18, 1959 as Im at Blame, Wash, by Mrs. Kline; taken at San Jose Nov. 7, 1959 at our banding station and released 27: 104107 banded Nov. 11, 1959 as Im at San Jose; found dead at Cassidy, Vancouver Island, B.C., letter Aug. 22, 1960.

31-138666 banded Jan. 28, 1962 as Ad at Alviso (5 miles from banding station at San Jose) was killed by a cat on August 19, 1962 at Bellingham, Wash.

31-188552 banded March 26, 1962 as Ad at Coyote (19 miles from band station) was shot May 12, 1962 at Port Angeles, Washington.

Most of these birds were recovered in the north on dates marginal to the breeding season, so we cannot say with certainty that they had bred or been hatched where they were taken. The last individual listed is an especially valuable record, for May 12 falls within the nesting season. Hence we can say that this bird wintered at least as far south as latitude 37° N. and bred at latitude 48° N., an air distance of about 750 miles. This checks with the record of the Puget Sound sparrow banded at Berkeley, Calif. that was found the next summer at Victoria, B. C., also an air distance of about 750 miles (Clabaugh, 1929).

A wealth of data on longevity in white-crowned sparrows is accumulating through banding records. Linsdale (1949) gives the following records for white-crowned sparrows banded at the Hastings Reservation: “18 white-crowned sparrows have returned after 5 years, while 5 have been captured after 7 years, and 2 lived at’ least 8 years.” C. G. Thompson (1960) reports capturing a Gambel’s sparrow known to be 7 years old, a Nuttall’s sparrow 6 years old, and four Puget Sound sparrows known to be 5, 6, 7, and 8 years old, respectively. The most complete individual record I know of is that for a Gambel’s sparrow banded as an immature by Franklin G. Crawford in Altadena, Calif. Nov. 22, 1942. It was recaptured 11 times, at least once each year, through March 1950. On March 5 of that year it was taken for the last time when it was nearly 8 years old. Crawford (1950) states that this is the only individual, out of 848 birds he banded between 1941 and 1946, with a proved life or more than 5 years.

No list of the types of information yielded by banding would be complete without mention of the study conducted at Pasadena, Calif. by Harold and Josephine R. Michener (1943) on the rate and sequence of prenuptial molt in Z. l. gambelii. Repeated observations of the same individuals obviously played an important role in this thorough analysis. Banding and retrapping made this possible, and at the same time permitted the subjects of the study to live under natural conditions.

DISTRIBUTION

Range: NuttaIl’s white-crowned sparrow is resident along a narrow coastal strip of central California from Mendocino County south to Gaviota Beach, Santa Barbara County. (Two breeding records at Goleta Flat, Santa Barbara County.)

Egg dates: California: 215 records, March 3 to July 24; 112 records, April 4 to April 27; 28 records, May 2 to May 15; 7 records, June 12 to July 24.

*This account also contains material on Zonotrichia leucophrys gambelii, Z. l. oriantha, and Z. 1. Pugetensis.

**The author acknowledges with thanks Grant number 2804 from the Penrose Fund of the American Philosophical Society, which made possible the literature search for data on range, migration, and nesting cycle. Thanks are due F. S. L. Williamson, who contributed unpublished data on clutch size in Z. l. gambelii from the Terrestrial Avifauna Study, Project Chariot of the Atomic Energy Commission, and L. H. Walkinshaw, who contributed data on behavior, nesting, and clutch size in this race. Richard C. Banks contributed data on distribution.

The author also wishes to thank Barbara Lilley Mooney, who carried Out the literature search, Priscilla Phillips, whose banding records and observations made possible the writing of the sections on fall and winter for Z. l. gambelii, and Anne Hinshaw Wing, whose analysis of Gambel’s sparrow song appears in the text. Many ornithologists contributed unpublished data through personal communications, which are acknowledged in the text.

The original field data supplied by the author were obtained under grants from the National Science Foundation, Society of the Sigma Xi, American Philosophical Society, and from the Committee on Research, University of California, Santa Barbara.

GAMBELS WHITE-CROWNED SPARROW

ZONOTRICHIA LEUCOPHRYS GAMBELII (Nuttall)

Additional material on this race is included in the account of Nuttall’s white-crowned sparrow.

Contributed by BARBARA BLANCHARD DEWOLFE

HABITS

The populations of Gambel’s white-crowned sparrow discussed in this section are those wintering at Davis, Calif., and those breeding at Mountain Village and College, Alaska. As none of the thousands of Gambel’s sparrows banded on the wintering grounds has been recaptured on the breeding grounds, we do not know where the birds of a given wintering population breed or vice versa.

Gambel’s sparrows nesting in Alaska compress their breeding cycle into an even shorter period than the Puget Sound sparrows at Friday Harbor. They establish territories, nest, and care for the single brood of young until they become independent in about 2½ months. The greater part of the time saved lies in the fledging of only one brood, but compression of several other intervals is necessary to make fullest use of the short time between arrival on the breeding grounds and gonad regression. Males arrive first and in almost full breeding condition. They immediately establish their territories, so that no time is lost after the females arrive either in achieving physiological readiness to mate or in finding and defending nesting areas. The females are ready to mate end to begin work on the nest within a few days. The interval between completion of the nest and laying the first egg also averages shorter than in pugetensis.

Spring: In spring the members of flocks wintering at Davis showed some slight hostility toward one another about 10 days before the first individuals departed. They squabbled over food and, less frequently, fought or pursued one another. Otherwise the behavior prior to migration was identical to that described for the Puget Sound sparrow. In 1943 the Gambel’s sparrow flocks at Davis began to decrease in numbers on April 13, the last banded bird was seen April 20, the last unbanded one on April 30.

My observations of Gambel’s sparrows in migration were chiefly from April 21 to 26, 1936, when I watched a few transients at Friday Harbor. In marked contrast to the noisy breeding Puget Sound sparrows, the Gambel’s sparrows sang so softly that it was hard to locate them even when they perched no more than 10 feet away. The few I saw were either solitary or in the company of migrating golden-crowned sparrows. Twice I found an individual perched in semidarkness high on a rafter inside a barn, with some golden-crowns. T. T. McCabe (personal communication) states that migrants may arrive m mountainous country in British Columbia long before they can live on the breeding grounds and may spend a month in valleys directly below suitable breeding areas.

The arrival of Gambel’s sparrows on breeding grounds at Mountain Village and College, Alaska, differed from that of the Friday Harbor sparrows in two respects: The Gambel’s sparrow males arrived more than a week ahead of the females, and they did not come in a body, but filtered in gradually over periods of 2 or 3 weeks. The influx of Gambel’s sparrows destined to breed at Mountain Village in 1950 lasted at least 19 days. Two males came on May 9. I color-banded one and followed it through the nesting cycle. From then until May 25 males. continued to come in. The average arrival date for 17 males was May 15. The females arrived between May 17 and 28, and the average date for 11 females was May 23. These data accord closely with those of Henry C. Kyllingstad (personal communication) for the same locality from 1942 to 1948. I assume his records of early arrivals are for males. His earliest record is for May 7, 1943, and his average date for “first Gambel’s sparrow seen” is May 10. His average date when the species becomes common falls between May 14 and 15.

At College, Alaska, in 1957 the influx of males lasted at least 15 days. Gambel’s sparrows were first reported in the area May 4, and migrating flocks were seen May 6. My observation provide the average arrival date of May 8 for 19 males I saw in places where the day before there had been none. By May 11 the numbers of males in the areas I was watching began to stabilize. The last male I suspected of being newly arrived came May 19. On May 16 I first saw a female. The average date of arrival for 12 females was May 17.

The Gambel’s sparrows at both Mountain Village and College were less vigorous in their territorial displays than were the Friday Harbor Puget Sound sparrows, in spite of the fact that the Gambel’s sparrows arrived with gonads much larger and closer to breeding condition than did the Puget Sound birds. On the margin of the range at Mountain Village territorial behavior was so weak as to scarcely deserve the name. At College the males displayed more forcefully, but still not so vigorously as had the Puget Sound males. The sparseness of the breeding population and the cold and inclement weather at Mountain Village could conceivably explain the lack of territorial display there, but at College, where Gambel’s sparrows were abundant and the weather more favorable, other factors must have been responsible, the most likely one being the absence of females during this phase.

At Mountain Village newly arrived Gambel’s sparrows sang weakly and showed no attachment to any piece of ground. If I approached them they flew straight away for distances up to a mile. Unlike the Nuttall’s or Puget Sound sparrows in comparable stages of development, they showed no tendency to face me and sing, or even to seek shelter nearby. Some days after arrival, however, each male had established a headquarters where he spent the greater part of each day. The area still did not merit the term “territory” in the strict sense, for the males sometimes left their headquarters for hours. I saw some pursuits, but also cases of complete tolerance of one male by another. When the females arrived, each pair restricted their activities to an area, including the male’s headquarters, but, with one notable exception discussed in the next section, neighboring males never had violent disputes such as I saw commonly at Friday Harbor.

One unique feature of behavior I observed at Mountain Village was that a few pairs nested where only one element of the typical breeding territory, dense shrubbery, was present. To reach the other two essential elements, namely grass and open ground, these pairs regularly flew from their nesting ares s above the village a mile or so down to the river’s edge. There they foraged, and if they had young they carried food back to the nest. To make these trips they had to pass directly through the nesting areas of other Gambel’s sparrows, which did not appear to resent the intrusion. This is the only case I know where the three essential elements of the breeding territory were not contained in an area inhabited exclusively by a pair.

At College the settlement of the ground followed a more conventional pattern. Except for the fact that the males arrived first, territory establishment contained all the behavior elements of the Friday Harbor-Puget Sound sparrow population. Each male became attached soon after arrival to en area where he sang loudly and regularly. I saw frequent pursuits and fights, but they were neither so common nor so intense as those at Friday Harbor.

Courtship: Courtship in Gambel’s sparrow begins just as the males are achieving full breeding condition. Therefore the courtship period is shorter than those in the Nuttall’s and Puget Sound sparrows, which begin this phase in a less advanced stage of gonad development. As in the Puget Sound sparrows wintering at Berkeley, there was no evidence of sexual interest between flockmates in the Gambel’s sparrows wintering at Davis or at Santa Barbara. Indirect evidence from collecting records suggests that members of a given breeding pair may winter not only in separate flocks but at different latitudes. John T. Emlen, Jr. (1943) found from examination of Gambel’s sparrow specimens at the Museum of Vertebrate Zoology and from his own collecting records that the sex ratios are unequal. At Davis, males outnumber females by more than 5 to 1. Emlen’s findings are in accord with my own less-exact figures for the proportion of males to females in gambelii collected at Davis and in pugetensis taken at Berkeley. I, too, recorded the preponderance of males to females by about 5 to 1. That this is not entirely due to accidents of collecting the bolder and more conspicuous males is indicated by the fact that Emlen found Gambel’s sparrow females outnumber males by almost 2 to 1 in the collections from the southern and eastern portion of the winter range. As we have no evidence of disparity in numbers of males and females on the breeding grounds of either race, at least some of the breeding pairs must break up and repair to different localities in winter.

At Mountain Village in 1950 the females must have gone directly to the males’ territories on arrival. I never found any wandering free. Hence the day of arrival must also have been the day the bond between the pair started to form or to re-form between mates of the previous year. The interval between the day of arrival of the female and the first observed copulation varied from 1 to 10 days for three pairs. The same behavorial elements of singing, trilling, and posturing were present as in the other two races.

At College, on the other hand, the first females I found were not followed by or following a male, and did not appear to be attached to any particular area. They flew long distances, trilled, and postured without any external stimulus that I could detect, and as they foraged they poked into crevices and peered under tufts of dry grass. A few days later all females I saw were paired. Each was followed closely by a male, and she trilled and postured vigorously when he sang or came near her. She continued to peer into crannies and under grass tufts. Four days after I first saw a female I saw two pairs copulate and watched others go through all the characteristic behavior preliminary to coition. Also at this time the females were building nests.

I have no evidence of polygamy in the Gambel’s sparrow, but color-banding adjacent pairs at Mountain Village revealed one case where a male tried unsuccessfully for 3 days to steal the mate of a neighboring male while his own female was completing her first clutch and starting to incubate. The facts were as follows (Oakeson, 1954):

On May 23, Male 1 acquired a mate. She built one nest, deserted it, and was finishing another when on May 28 copulation occurred. Female 1 must have begun incubation May 31, judging by the date her eggs hatched. Meanwhile the neighbor on the east, Male 7, had acquired a mate. She started her nest May 30. That same evening, Male 1 ‘attacked’ her. * * * Male 7, which until then had been singing regularly but not forcefully, chased Male 1, then began to sing loudly from conspicious perches, patrolling his area as do males during incubation. The next evening, Female 7 was attacked again by Male 1. Then each time she flew, she was pursued by both males. Finally Male 7 began to chase the intruder, but could not drive him hack to his own area. At 9:10 p.m., over one and one-half hours after the dispute had begun that evening, Male 7 began to patrol his territory, which now, owing to his intensified behavior, merited the term in all its connotations, and incorporated into his patrol a high aerial pole, from which he sang loudly. (Since there are no tall trees at Mountain Village the aerial poles near the Trading Post were favorite singing perches.) There followed a ‘singing contest’ between the two males. This lasted until 9:45 p.m., some time after other birds had quieted down for the night. During the contest, Female 7 trilled and postured vigorously hut her mate was too preoccupied to pay any attention to her. The next evening, June 1, the performance was repeated, and Male 1 flew at Pair 7 whenever they attempted copulation, preventing the mating not only psychologically but also physically, at least as long as I watched. By the next day the situation had returned to normal: each male sang from his respective territory. Pair 7 must have succeeded in copulating, for the female finished her nest June 1, laid her first egg June 3, and all eggs hatched June 19.

Nesting: At Mountain Village I first saw nest building 11 days after the average date for arrival of the males and only 3 days after the average day date for arrival of the females. The intervals between arrival and nest building at College are closely comparable: I first saw a female building on May20, 12 days after the average date of arrival for males and 3 days after average arrival date of the females. At both localities the interval between arrival of the female and the start of nest building lasted only 2 or 3 days, as contrasted with the corresponding interval of 1 to 2 weeks for Puget Sound sparrows at Friday Harbor. The time spent in building is also shorter. At Mountain Village four Gambel’s sparrow females spent 2 to 4 days in building. At College one female was first seen building on May 21, and again on May 24. She laid her first egg May 26. It will be remembered that both Nuttall’s and Puget Sound sparrows spend from 7 to 9 days building the first nest of the season.

Eggs: Gambel’s sparrows lay the first egg from 0 to 3 days after the nest is finished. I have two records of females that worked on their nests the same day they laid the first egg of their clutch. For seven females seen while building, the average interval between the date they were last seen building and the date they laid their first egg was 1 day. The corresponding interval for nuttalli is 3.6 days, for pugetensis, 2.6 days.

Egg laying extends from late May through mid-July in this race. The earliest date for eggs is May 21 for two clutches I found at College in 1957. The median date for the same locality for 13 clutches I found was May 23. At Mountain Village in 1950 the median date was later, between June 2 and 3 for six clutches.

Clutch size averages greater in the Gambel’s sparrow than in either nuttalli or pugetensis. The number of eggs per clutch ranges from 3 to 6, with one record of 7 eggs in one nest. The average of 76 clutches is 4.58. The most common number of eggs per clutch is 5. The percentages are as follows: three eggs per clutch 9.21 percent, four eggs per clutch 32.9 percent, five eggs per clutch 50.0 percent, six eggs per clutch 6.6 percent, and seven eggs per clutch (one record only) 1.3 percent. The average clutch size by month is: May 4.71, June 4.65, July 3.40 for 14, 57, and 5 clutches, respectively. I have no evidence that Gambel’s sparrows lay more than one clutch per season if they successfully fledge one brood.

As in pugetensis, clutch size in the Gambel’s sparrow appears to increase with latitude. Five sets of eggs taken between latitudes 51° and 55° N. average 4.20 eggs; 4 clutches collected between 56° N. and 60° N. average 4.50; 32 clutches between 61° N. and 66° N. average 4.56 eggs, and 27 clutches taken between 67° N. and 69° N. average 4.74 eggs per clutch.

Incubation: I know the length of the incubation period for three females of this race. Two at Mountain Village hatched their eggs 12 days after they started incubating. One at College hatched her eggs 11 days after she began incubating. This is the only case in any of the races I found where the incubation period was less than 12 days. One of these females began sitting the day she laid the fifth and last egg of the clutch. The other two began sitting the day they laid the fourth of five eggs.

Young: The nestling period in gambelii averages shorter than in nuttalli or pugetensis. For 13 nestlings watched at Mountain Village, the time spent in the nest ranged from 8 to 11 days, or an average of 9.6 days. For 15 nestlings watched at College, the time spent in the nest ranged from 7 to 9 days, or an average of 8.4 days.

Gambel’s sparrow young become independent of their parents at least by the time they are 28 days old. I trapped three young of a pair at this age. In this, they do not differ significantly from the Puget Sound sparrow young at Friday Harbor.

Fall: In 1957 I watched Gambel’s sparrows before and during fall migration at College, Alaska (lat. 64°49’ N.). That same year, Mrs. Priscilla Phillips (personal communication) watched the arrival of Gambel’s sparrows at Santa Barbara, Calif., some 2,400 air miles away. Though observations at these two localities complement each other in several respects, no identity of the two populations is implied, for despite the thousands of Gambel’s sparrows banded each year, none has been taken on both breeding and wintering grounds, and we do not know how far a given individual travels. By analogy with the Puget Sound sparrow, we should expect birds nesting in the northern part of the breeding range to migrate to the southern part of the wintering range. Thus it is possible that Gambel’s sparrows breeding at the latitude of College may winter in southern California. In this connection it is interesting that a singing male in breeding condition collected at College on May 6 was using a song pattern identical with one of the song patterns heard in winter at Santa Barbara.

Excerpts from my field notes outline the start and progress of migration as I recorded it at College in 1957.

“August 6. Kallio, Superintendent of the University of Alaska Experiment Station, reports Gambel’s sparrows gathering in the strawberry patch at the Experiment Station. He interprets this as a sign the birds are getting ready to migrate.

“August 10. Gambel’s sparrows appear to be definitely in migration. They utter eep’s rapidly, and occasionally sing weak, fragmentary songs.

“August 13. After hunting along roadsides where breeding birds had been common I observed that the birds behave as a winter flock. They appear to be migrating. Two adults were seen in a flock of about ten birds. All were furtive, shy, keeping foliage between them and me. Other birds suspected of being in migration: Myrtle warblers, Ruby-crowned kinglets, Savannah sparrows.

“August 14. For the past week or so I have noticed that trapped birds utter call notes when I hold them in a darkened cage. They fight to escape and squirm more vigorously in my hand than they did before.

“August 15. Mr. Kallio reports that the numbers of Gambel’s are now markedly fewer. About a week or ten days ago he saw the greatest number.

“August 17. There are definitely fewer Gambel’s sparrows than on August 6 and 8. Other species flocking this a.m.: Juncos, Savannah sparrows.

“August 18. Heard only a few Gambel’s sparrows.

“August 19. At meadow between 8- and 9-mile on road to Nenana, saw a flock of 18 Gambel’s sparrows. No singing or uttering of location or alarm notes. They perched close together.

“August 25. 8:30 a.m. For the first time I do not see a sizable group of Gambel’s at the trap sites.

“August 26. At meadow between 8- and 9-mile, found only three immatures. Heard no song. Saw no adults. I listened and looked at spots where found Gambel’s sparrows last week. None there. Did not see any large migrating flocks anywhere this a.m.

“August 27. At junction of railroad tracks and road to Nenana, no Gambel’s sparrows seen or heard. Saw one immature Gambel’s in meadow between 8- and 9-mile. Heard two Gambel’s songs in rapid succession.

“August 30. Drove slowly along Chena Pump Road to North Star Ranch. Stopped to look and listen. No birds of any species except fox sparrows seen or heard. Stopped back of trap site 3 and listened. Heard geese and a fox sparrow but no Gambel’s sparrows.”

To fill in the details of the picture of fall activity in the Gambel’s sparrow at College, I operated 9 Potter traps at four stations around a field near the territories of about 11 nesting pairs, banded and released 256 adults and young, and collected specimens for analysis of molt, fat, gonad size, thyroid activity, and body weight. The data from banding and collecting during the period from June 26 to August 27 substantiate the impression, given by the field notes on behavior just quoted, that by early August migration was in full swing. The local breeding birds probably left the trapping area at least by late July, and from then on until late August successive waves of Gambel’s sparrows passed through the area, staying on the average only a few days before moving on.

The data reveal a major difference between fall departure at College and spring departure at Davis and Santa Barbara: in the north the residents at the trapping area left very early in the migration period, and were replaced by successive influxes of birds, presumably from farther north, whereas in the south the winter residents gradually decreased in numbers until all had gone, and they were not replaced by others. One similarity between spring and fall departure has been noted: in both fall and spring, the birds may start their migration before the molt is finished, and while still accumulating fat.

Mrs. Priscilla Phillips (personal communication) made daily observations on the arrival of white-crowned sparrows at her home in Hope Ranch for five consecutive years, from 1957 through 1961. In addition, during the first three years she trapped and banded 399 Gambel’s sparrows at her feeding station, where the combination of a nearby lemon orchard, brushy hillsides, and a well-watered lawn provided optimum conditions for attracting and observing the flocks, some adults of which had doubtless fed there the previous winter.

One point of particular interest is the near identity of dates for the first white-crowned sparrow seen by Mrs. Phillips at Hope Ranch in five consecutive years. Another is that the arrival consists of a series of discrete influxes, often several days apart. The numbers observed may even decrease between successive influxes, suggesting that not all the birds coming to Santa Barbara in the fall actually winter there. A third point is that the arrival period lasts at least 6 to 7 weeks. A fourth is that in the early part of the arrival period the birds come either singly or in small inconspicuous groups and are trap-shy, so it would be easy to miss the first arrivals. Fifth, the earliest Gambel’s sparrows to appear are accompanied by Puget Sound sparrows, which do not tarry long in the area. By early to mid-October of 1957, 1958, and 1959, all of the birds entering the traps were Gambel’s sparrows. All these observations are substantiated by Mrs. Phillips’ banding records. One point on which observation and banding are not entirely in agreement is the proportion of young-of-the-year to adults. On this point, the data from observation tend to be more variable than those from banding.

Except for the sight record of one white-crowned sparrow on September 3, 1957, the dates for the first individual seen by Mrs. Phillips at hope Ranch (either gambelii or pugetensis, race not determined) are Sept. 20, 1957, 1958, and 1959, Sept. 18, 1960, and Sept. 19, 1961. The earliest date when birds identified with certainty as gambelii were seen at Hope Ranch are also almost identical: Sept. 20, 1957, 1958, and 1960, Sept. 21, 1961 and Sept. 22, 1959.

Both observations and banding indicate that the arrival consists of separate influxes, with many or most members of the early ones probably continuing their flight south. Each influx is accompanied by much commotion: choruses of song, often weak and fragmentary, the uttering of call notes and squabbling notes, restlessness characterized by almost continuous movement in the trees or brush, and the frequent sudden flight or dive for cover of the whole foraging flock. On the basis of these characteristics of behavior, Mrs. Phillips estimates that in 1957 at least eight influxes occurred between September 20 and October 31. Others may have occurred in November, but by then the numbers coming to the feeding station were too large and too stable to reveal with certainty any new arrivals. In 195S she again observed at least eight influxes, between September 20 and November 7, and in 1959 at least six influxes between September 20 and October 16, when she ended her daily observations.

Banding records indirectly substantiate Mrs. Phillips’ impression that many members of each influx move on. Only 3 out of 60 Gambel’s sparrows she banded between Sept. 20 and Oct. 17, 1957, were I retaken the following winter.

The intervals between observed influxes for the 3 years at Hope Ranch varied from 2 to 15 days, with the shorter intervals tending to be in the early part of the arrival period. The average of 19 intervals between influxes is 6.0 days. The most commonly occurring interval (6 of the 19) is 5 days.

The period elapsing between the date the first white-crowned sparrow identified as a Gambel’s sparrow was seen in Hope Ranch in 1957 (September 20) and the end of the arrival period, judged by stability of numbers trapped and observed and the general quieting down of the flocks (November 1 at the earliest) is at least 42 days. The comparable period for 1958 is at least 48 days. The fall arrival at Santa Barbara over a period of 6 to 7 weeks agrees with the data gained from banding and watching the birds at College, Alaska, in the fall of 1957, which showed the birds continued to pass through the trapping area from at least late July through August 27 or later.

To the question of whether immatures or adults arrive first in the Santa Barbara region, the data available give no consistent answer. Mrs. Phillips’ observations at Hope Ranch agree with those of Henry Kyllingstad at Mountain Village as to the varying proportions of immatures and adults in successive influxes of migrants. The ratios also varied during the years Mrs. Phillips watched.

Although, as was stated at the beginning of this section, no identity of populations is implied, the observations and banding data from College and from Santa Barbara agree in the following respects. First, the periods of time the birds are on the move, passing through College and arriving at and passing through Santa Barbara, are comparable. Each is 6 to 7 weeks or more. Second, both at College and at Santa Barbara, the migration consists of a series of influxes. There is strong evidence that the members of a given influx did not stay long in the College area, and only slightly less conclusive evidence that many of the arrivals at Santa Barbara did not stay long there either. Third, neither at College nor at Santa Barbara was there a consistent pattern of arrival according to age group. Fourth, the ratios of immatures to adults trapped at College from mid-July through late August varied between approximately the same limits as did the sight estimates made of relative numbers of immatures and adults for the fall arrival at Hope Ranch the same year, and as did the trapping records at Hope Ranch for November of two years. Whatever the hazards of the flight south, the immatures appear to fare about as well as the adults.

The observations from College, Alaska, and from Santa Barbara, Calif., furnish data for estimates of the length of time spent in the southward migration. Assuming that birds of the latitude of College may fly as far south as Santa Barbara to winter, we know that the interval between the time the local breeding population left College (at least by July 22) and the earliest data of arrival of Gambel’s sparrows that same year at Hope Ranch (September 20) is at least 60 days. This estimated migration period is over 1½ times as long as that estimated for the flight north in spring. Qakeson (1954) estimated that it requires approximately 35 days for the flight north from the latitudes of southern California to those of Mountain Village, Alaska, an air distance of about 2,700 miles.

An individual bird flying in about 60 days from the latitude of College to that of Santa Barbara, a distance of about 2,450 air miles, would average about 41 miles per day, which a white-crown flying at 20 miles per hour (Pearson 1961) could cover in about 2 hours. The birds should be able to fly twice as long, and twice as far daily, and still have ample time each (lay to forage and rest.

Winter: In winter the Gambel’s sparrow, like the Puget Sound sparrow, forms flocks that are homogeneous assemblages, with no hint of the tendency of flock members to pair off. Also like pugetensis, the gambelii flocks observed at Davis were approximately stable as to size and tended to restrict their daily movements to a limited area. The following quotation from Blanchard and Erickson (1949) refers to flocks whose members were color-handed at Davis in 1942-1943.

Throughout the winter the birds flock in groups of 30 to 50. Each flock is restricted to a given area, though there may be a substantial overlapping of areas. Of the 48 individuals either retrapped or identified by sight at Davis, 29 were found at the same spot where they had been banded, 12 were found not more than 500 yards away, which is not over the normal maximum distance traveled by any one flock in the course of a day. The remaining few were seen up to a maximum of 976 yards away. General localization of each flock to a specific area is further indicated by many hours of observation of individual flocks throughout the winter. Each flock had its headquarters, within a few yards of which it spent the greater part of each day.

Once the birds were established, behavior during the ensuing five months until mid-March showed little change. No fights and few pursuits were seen. Singing continued sporadically but was lacking in force, and the individuals which sang always did so as integral parts of the flock.

Not only may individuals stay in a restricted area one winter, but they may also return to the same place year after year. Bird-banders have furnished a wealth of data on returns of white-crowned sparrows Blanchard and Erickson (1949) note: “In Santa Barbara between 1943 and 1947, 282 Gambel sparrows were banded at stations where traps were operated at least occasionally throughout the months from September to May. One hundred, or 35 percent of the 282, were taken again. Of these 100, 47 were retaken in two succeeding winters, two in three, and one in four winters.” Of 223 Gambel’s sparrows Mrs. Phillips banded at Hope Ranch in 1957-1958, 14 returned the following fall. Of 129 birds banded there in 1958-1959, 8 were recaptured in the fall of 1959. F. G. Crawford (personal communication) states that “in preparing return records for Gambel sparrows, the similarity of calendar dates for original capture and return seemed to occur too frequently to be purely coincidental.” Of 121 banded Gambel’s sparrows recaptured by Crawford in succeeding years, 50 percent were taken within 30 days of calendar date of original capture.

The waves of migrants arriving at a given place on the wintering grounds do not stop with the onset of winter. Elliott McClure (personal communication), who banded nearly 3,000 white-crowned sparrows (presumably all or nearly all gambelii) at Bakersfield during four winters, states: “A wave of birds appeared to enter the area in October, increase in November, and decrease in December. This was followed by another increased flow through January and February which then decreased in March and April.” After presenting several possible explanations for the wave-like movements of white-crowned sparrows suggested by the banding records, McClure concludes, “The wave action of these birds seems inescapable, both going south and returning north, and the mass of birds moving was great enough that unmarked ones were constantly entering the vicinity of the traps. Jean M. Linsdale (1949), working with White-crowns at hastings Reservation about 150 miles north of Bakersfield and in the coastal range, found a similar wave action but with different distribution during the winter months.” Although, as Mewaldt’s data discussed in the following paragraph indicate, one cannot assume that the mid-winter waves observed by McClure came from any particular compass direction, the interesting point is that influxes of birds appeared during months when migration proper does not occur.

That white-crowned sparrows are potentially mobile in winter is indicated by the trapping records of L. R. Mewaldt (1963) who trapped and banded white-crowned sparrows (Z. l. pugetensis and Z. l. gambelii) at San Jose, Calif., for 8 consecutive seasons. From 1954-55 through 1960-61, the total numbers of both races trapped ranged from 181. to 292 per season, or an average of 227.1 for these 7 seasons. In 1961-1962 Mewaldt removed large numbers of banded white-crowns and shipped them by air to Baton Rouge, La. One of the many interesting results of this work was the increase in numbers of white-crowns trapped at San Jose that season—a total of 670 unbanded individuals. Mewaldt (1962) removed 430 birds for his homing experiment. He states: “Repeated removal of the ‘hard core’ of dominant birds coming to bait at the banding station probably permitted population pressure to fill the vacuum thus created.” While the situation created by removal of many wintering whitecrowns is obviously an unnatural one, the fact that so many birds came to Mewaldt’s station shows the inherent mobility of winter flocks, and also that no assumption can be made as to the direction from which mid-winter influxes come, such as those reported by McClure.

In summary, the data presented in this section indicate flexibility, as well as stability, of the location of winter flocks. Apparently movement may occur during mid-winter as well as during the migration periods proper. We must think of wintering Gambel’s sparrows as geographically stable, in that some individuals return to the same place winter after winter, but also as flexible, in that some may move about during mid-winter. Perhaps the relatively low recapture percentage of banded white-crowned sparrows in any wintering population reflects something more than mortality rates, or the low statistical probability of retrapping a given banded individual. It is conceivable that, during winter, some individuals may stay close to the continuous food supply that a banding station provides, while others may move about. In a species that shows high individual and racial variability in every physiological character so far studied, there is no reason to exclude the possibility of differences in individual behavior in winter. As an example of an individual that moved about during winter, T. E. Balch (personal communication) reports banding a Gambel’s sparrow at Glenn, Calif., Jan. 24, 1960; on March 1 the same year the bird was taken 25 miles away at Chico, Calif.

Gambel’s White-crowned Sparrow

DISTRIBUTION

Range: Central and western Canada and Alaska to Baja California and central Mexico.

Breeding range: This race intergrades with Z. l. leucophrys on the south shore of Hudson Bay from Churchill to James Bay. The breeding range of the Gambel’s white-crowned sparrow extends from northern Ontario (Weenusk, Fort Severn) to northern Manitoba (York Factory, Ilford, Bershmer, Churchill) into the Northwest Territories, (South Henick Lake, Artillery Lake, Coronation Gulf, Fort Good Hope, Mackenzie River Delta) west to the Yukon Territory (Lapierre House, Old Crow River) west to Alaska (Sheenjek River, Colville River Delta, Koalak River, Pitmegea River, Cape Lisburne) south in western Alaska (Kotzebue Sound, Wales, Nome, Norton Sound, Mountain Village) to southwestern Alaska and the Alaskan peninsula (Kanakanak, Nushagak, Egekik River, Port Moller, Izembek Bay) northeast to the Kenai Peninsula, the Knik Arm of Cook Inlet, to Copper Center, the Chitina River, into southern Yukon Territory (Burwash Landing, Slims River mouth) into British Columbia (Mile 85 on Haines Road, Bennett, Atlin, Telegraph Creek, Chezacut Lake) to northern Washington (Hart’s Pass) east to Field, British Columbia, intergrades with Z. l. oriantha near the United States-Canada border; north in Alberta (Banff Park, Jasper and Grimshaw) northeast to Saskatchewan (Fond-du-lac and Reindeer Lake).

Winter range: Winters from southern British Columbia (Comox, Okanagan Landing), southeastern Washington (Cheney, Pullman, Snake Rixer Canyon), southern Idaho (Heyburn), central Wyoming (Thermopolis), north central and southern Colorado (Mesa County, Boulder County, Pueblo County), northern and eastern Kansas (Wallace, Manhattan, Lawrence), western Oklahoma (Roger Mills County, Comanche County); west to the coast; south to southern Baja California (San Jose del Cabo), to the islands of Las Tres Marias, east of Nayarit (Las Varas), Aquascalientes (6 miles southwest of Aquascalientes), San Luis Potosi (San Luis Potosi), north to northern Tamaulipas (Nuevo Laredo, Matamoros), and to southern and central Texas (Brownsville, Boerne, Concho County).

Casual records: Casual in migration or in winter in southern Ontario (Toronto), southern Wisconsin (Madison), Michigan (Jackson County; Huron Mountain, Marquette County; Whitefish Point), New York (Ithaca), Illinois (Waukegan; Newton, Jasper County), Ohio (Leetonia, Columbiana County; Waterville Township, Lucas County; Fairfield County), Maryland (Patuxent Research Refuge near Bowie), Virginia (Blacks; Lexington) Tennessee (Nashville), South Carolina (Mount Pleasant), Georgia (Tipton), and Alaska (Barrow), and British Columbia (Okanagan Landing).

Accidental in Alaska on Priblof Islands (St. Paul Island), and Little Diomede Island (Ignalook Village); on Bank’s Island (Sach’s harbour); District of Franklin; and in Japan (Honshu).

Migration: Early dates of spring arrival are: Alberta: GO miles north of Edmonton, May 4. British Columbia: Pentincton, Lake Okanagan, April 23. Manitoba: Churchill, June 8. Mackenzie: Fort Enterprise, approximately 150 miles north of Great Slave Lake, May 26: Fort Providence on Great Slave Lake, May 9. Yukon: Forty-mile (lat. 64), May 10; La Pierre House, May 25. Alaska: Kodiak Island, before May 23; Mountain Village, May 7; Nome, May 29; Kowak delta, May 21; College, May 4 (2 years record).

Late dates of spring departure are: Aguascalientes: 6 miles southwest of Aguascalientes, March 1. Durango: 12 miles west of Lerdo, one specimen, March 1. Baja California: San Fernando; islands offshore: Los Coronados, Cedros, San Benito, April 29. Sonora: Guaymas, April 24. Coahuila: Boquillas wash in the Sierra del Carmen, April 27. Texas: Marathon, Brewster County, May 6; Pine Springs Camp in the Guadalupe Mountains, May 3. Oklahoma: Cheyenne, May 13; Indiahoma, May 8. Arizona: Tucson valley area, May 13; Baboquivari Mountains, May 12; Fort Verde, May 11. Colorado: Mack, Mesa County, May 29; Clear Creek district, June 12. California: Twenty-nine Palms, May 6; Polvadero Gap, Kettleman Hills area, May 17. Nevada: Meadow Valley, Lincoln County, May 24; Eldorado Mountains, May 3. Oregon: Lake County, May 20. Washington: Cheney, April 27.

Early dates of fall arrival are: Washington: Prescott, September 2. Oregon: Wheeler County, September 13; Warner Valley, August 30. Nevada: Indian Springs, September 10; Boulder Beach, September 17. California: Eagleville, Modoc County, September 2; Los Angeles, September 20. Utah: -St. George area, numerous by October 1. Colorado: Denver area, mid-September; Colorado Springs, September 8. Arizona: Pima County, September 23. Kansas: Manhattan, October 9; Wallace, October 12. Texas: Guadalupe Peak (10 miles east), October 6; Chisos Mountains, Green Gulch, November 14. Sonora: Cajon Bonito Creek, Septembe~ 8. Baja California and offshore islands Los Coronados, Todos Santos, San Benito, Cedros, October 7. Tamaulipas: -Matamoros, November 25. Las Tres Marias: M aris Madre, December 28.

Late dates of fall departure are: Alaska: Kowak River (now Kobuk) opposite the mouth of Hunt River, September 2; Nushagak, September 18; Fairbanks (70 miles southeast), October 20; Juneau, October 1. Mackenzie: Great Bear Lake, September 5. Ontario: Shagarnu River, August 10. British Columbia: Alberta: Edmonton, September 26. Washington: Monument 83 on U.S. and Canada boundary at long. 120°38°½’, September 3; Harts Pass, during the second week of August.

Egg Dates: Alaska: 52 records, May 21 to July 13; 42 records, May 23 to June 15. Alberta: 3 records, June 13, June 21, June 28.

British Columbia: 6 records, June 7 to June 22. Mackenzie: 3 records, June 15, June 30, July 15.

Yukon Territory: 11 records, June 1 to June 17.

MOUNTAIN WHITE-CROWNED SPARROW

ZONOTRICHIA LEUCOPHRYS ORIANTHA Oberholser

Additional material on this race is included in the account of Nuttall’s whitecrowned sparrow.

Contributed by BARBARA BLANCHARD DEWOLFE

HABITS

This race consists of a group of western populations of whitecrowned sparrows that breed chiefly at high altitudes in the Canadian and Hudsonian life zones and winter in the lowlands. Until they were designated as a separate race in 1930 by Harry C. Oberholser, they were included in the race leucophrys, although their breeding range is separated from the main portion of the leucophrys range by several hundred miles. In this article I include under oriantha those records for leucophrys prior to 1930 that pertain to the western populations.

One of the most interesting aspects of this race in California, where I observed it, is the fact that the breeding population consists of a series of isolated colonies highly variable in the time they begin nesting. Because the date when the snow melts enough to permit birds to nest on or near the ground varies considerably from one breeding locality to another, even in the same year, some populations of this race begin nesting in early June, whereas others have to wait until July. In fact many oriantha populations begin nesting later than do Gambel’s sparrows in northern Alaska.

Another characteristic of oriantha in California is the small size of many of the breeding populations. Some of the high mountain meadows it inhabits are too small to support more than four or five nesting pairs. Also in marginal areas minor vegetative changes, such as those brought about by a series of unusually dry or wet years or by the interference of man, may render a suitable locality unsuitable for nesting white-crowns, or vice versa. This may explain discrepancies in the literature regarding the occurrence of this race, such as the early reports (Merrill, 1888, E. A. Preble, MS) of whitecrowned sparrows breeding in the Mount Shasta and Crater Lake regions, where recent attempts to find them (Farner, 1952; Banks, personal communication; DeWolfe and DeWolfe, 1962) have failed.

A third interesting fact is that, although the latitudinal range of oriantha in California is approximately the same as that of nuttalli, its average clutch is significantly larger than that of the coastal form.

Data on clutch size and egg dates for oriantha are plentiful, but the literature contains almost nothing on its behavior. My own observations of this race in the Sierras and Cascades during the summer of 1960 include brief glimpses of territory establishment, incubation, and care of nestlings, none of which show obvious differences from those of the other races. I have no firsthand knowledge of behavior in this form during fall and winter. Owing to the paucity of information on the annual cycle, I present the data on habits in a continuous account rather than under the subheadings used for the other races.

Records of its movements in spring and fall suggest that not only does oriantha leave its wintering and breeding grounds later than does gambelii, but also that, depending on local weather conditions, oriantha may tarry en route between the two. Stephen S. Visher (1910) states that leucophrys (now oriantha) is an abundant winter visitor in Pima County7 Arizona and that “they remain several weeks longer than Z. gambeli feeding on the blackberry-like fruits of the mulberry. Last seen June 8.” Laurence M. Huey (1926) observed a flock of fifty white-crowns (which I assume were oriantha) on May 22 at El Rosario in Baja California. The following day only two were seen and one collected, indicating that most had departed the previous night. Joseph Grinnell and T. I. Storer (1924) state that what is now called the mountain white-crowned sparrow is a summer visitant to the Hudsonian Zone in the Yosemite region and that it passes through the lower levels on both sides of the mountains during spring migration.

The earliest definite records of the arrival of the Hudsonian white-crown in the Yosemite region are for May 10 (1916) * * * and for May 8 (1917) * * Migration was still in progress on May 22 (1919), as a male bird in Yosemite Valley on that date tarried only a short time before moving on. * * * Some individuals continue in their summer haunts until the end of September, several having been noted by us at Tuolumne Meadows on Sept. 29, 1915, but none anywhere later than that date.

Our highest record for the Hudsonian white-crowned sparrow was close to 11,000 feet altitude, in a patch of stunted willows in a draw between Mount Gibbs and Mount Dana, July 29, 1915.

Joseph Grinnell, Joseph Dixon and Jean M. Linsdale (1930) collected white-crowned sparrows thought to be still on their breeding grounds at Warner Creek in the Lassen Peak Region on Sept. 16, 1923 (elevation 8,000 feet).

Eustace L. Sumner and Joseph S. Dixon (1953) give many records of oriantha in Sequoia National Park. In 1942 the first spring arrival in the park was on May 12 after a snowstorm at Lewis Creek. In 1934 three white-crowned sparrows were seen (one was taken) in a willow thicket at 9,700 feet near Gallats Lake on the Kern-Kaweab River, June 7. That same year six were seen at Kaweab Gap, at 10,700 feet elevation, on June 16; one of these (a female parent, no. 9090 in the Dixon collection) and a nest with four fresh eggs were collected. The fall records are as follows: On Sept. 20, 1933, when the willows were turning yellow at heather Lake, just one bird, an immature, was present; the adults apparently had left on their fall migration. In 1940, one was found in Deadman Canyon on September 22 (they were no longer common in the vicinity); on September 24 one was seen at Scaffold Meadow. Two white-crowned sparrows noted at Giant Forest on Oct. 3, 1934, were evidently fall migrants from higher regions, as this species had not been there during the summer; two immatures were seen just below Tharp’s Rock on October 5; one adult and one immature at Willow Meadow and three adults at Cahoon Meadow on October 13.

For other states we have the following records: Percy M. Silloway (1907) states that the first white-crowned sparrow was heard singing at Flathead Woods, Mont. on June 5, 1906. Elliott Coues (1874) quotes Trippe as saying that this race appears in the lower valleys of Clear Creek County, Colo., the first or second week of May. As the snow disappears, the race ascends the mountain, reaching timberline by the middle of June. It commences building in July and young are hatched about the 20th of this month. In September it begins to descend; by October it is abundant at Idaho, and by November has disappeared. William L. Slater (1912) states that white-crowns arrive at Colorado Springs the last week of April, gradually move up the mountains, reaching timberline the middle of June. After nesting they descend in September and October to lower levels and linger until early in November. The species is common on the western slopes during migration.

At Mineral, Calif., a few miles from the south entrance to Lassen Volcanic National Park, Ranger Naturalist and Mrs. Merle Stitt (pers. comm.) reported seeing white-crowned sparrows in migration, both in spring and fall. The last date for spring of 1960 was May 24, two days after the last snowstorm of the season. On June 25 of that same year my husband and I found oriantha males starting to establish territories at King’s Creek Meadows inside the Park, (elevation 7,400 feet) but no females had yet arrived. By July 4, Park Superintendent and Mrs. Dixon Freeland reported pairs of white-crowned sparrows common at that location. Where the birds had been between May and July is a mystery, but if they kept a schedule comparable to those described above by Trippe and Slater they probably stayed in the vicinity below the nesting-grounds and moved up as the snow melted and the high country became habitable.

When we arrived at King’s Creek Meadows on June 25, 1960, we found several male mountain white-crowned sparrows already spaced out, singing frequently but weakly. We saw no females. The males showed about the same level of territorial jealousy as did newly arrived Gambel’s sparrow males at College, Alaska. They perched conspicuously, alternating in song with their neighbors, and occasionally pursued each other, but we saw no noisy conflicts, such as I have described for the newly arrived Puget Sound sparrows at Friday Harbor. By the next day the volume and frequency of song had increased, but there were still no females. On July 4 other observers (Beatrice Freeland, pers. comm.) found these meadows “alive with song,” with white-crowns of both sexes present in numbers. Whether the males arrive ahead of the females in all localities is not certain.

The eggs are laid from late May through early August. William L. Dawson (1923) states that this race may raise one or two broods. I have found no records, however, of a pair with young out of the nest and with a second nest of eggs or young such as was the rule in the Puget Sound sparrows at Friday Harbor. When all the available egg dates are arranged chronologically, they show no hint of a gap in the series that would suggest a second brood. My observations in 1960 provide one illustration of the wide variation between populations as to the time nesting may start. On June 19, 1960, my husband and I found a pair of mountain white-crowned sparrows at Tuolumne Meadows with young about four days old. If we assume that the incubation period was the same length as in the Nuttall’s sparrow, the female would have laid the first egg of this clutch on June 1. At King’s Creek Meadows, on the other hand, the females had not even arrived by June 26. Even assuming they came the next day and began work on their nests at once, they could hardly have started laying before June 29 at the very earliest, some four weeks later than the pair at Tuolumne Meadows.

The average number of eggs per clutch for 164 clutches is 4.03. If only those clutches of oriantha from localities within the same latitudinal range as nuttalli are averaged, the mean is almost identical, —4.04 for 145 clutches. This is significantly larger than the average clutch size for nuttalli (3.27 for 215 clutches). The number of eggs per clutch in oriantha is usually four whereas in nuttalli it is usually three. Percentages for oriantha are as follows: 3 eggs 5.0 percent; 4 eggs 87.6 percent; 5 eggs 6.8 percent; 6 eggs (one record only) 0.6 percent. Two extreme records of two and seven eggs each were not included in the averages; it is not certain that the set of two was complete and we have no evidence that the clutch of seven was laid by a single female.

As clutch size varies with month of laying in nuttalli, I calculated the average clutch size for oriantha for June and July separately: for June 122 oriantha sets average 4.08 eggs per clutch, 21 nuttalli sets average 3.24; for July 37 oriantha clutches average 3.89 eggs per clutch, 7 nuttalli sets average 3.14 per set.

These comparisons show that some factor or factors other than latitude and month of laying must be responsible for the difference in average clutch size between oriantha and nuttalii. The most obvious environmental difference is that of altitude and the resulting differences in the climates of the breeding grounds. A scatter diagram we constructed of the number of recorded oriantha eggs per clutch in relation to elevation, which ranges from 4,000 feet to 11,000 feet, shows no obvious correlation, either positive or negative, with altitude.

As the sets of oriantha eggs were taken over many years at many different latitudes, other variables may mask the altitudinal effect.

DISTRIBUTION

Range: Southern Saskatchewan and Alberta to Baja California and central Mexico.

Breeding range: This race intergrades with gambelii from northwestern Alberta in the Canadian Rockies to Glacier Park, Montana. An isolated population breeds in the Cypress Hills region of southwestern Saskatchewan and southeastern Alberta. The Mountain White-Crowned Sparrow breeds from western Alberta (Gorge Creek, Bragg’s Creek, Banff National Park) to northern Idaho (Continental Mountain) south to eastern Oregon (Wallowa Mountains), southwest through central and southern Oregon (Haney, Hart Mountain, Adel) to northeastern California (Warner Mountains). Isolated populations occur in the Cascades of west central Oregon (Three Sisters, Tumalo Creek, Crescent Lake) and of California (Bray, King’s Creek Meadows in Lassen National Park). From the Warner Mountains the range in California extends southward through the Sierra Nevadas (Mohawk, Plunias County to Horse Meadow, Tulare County). Isolated colonies breed in southern California (San Bernardino Mountains) and in eastern California in the White and Sweetwater Mountains, and on peaks in Nevada (Ruby Mountains and Jarbridge Mountains). The boundary of the range continues through southern Utah (Cedar City, Kanab area, Bluff) to northern New Mexico (Pecos Baldy, Wheeler Peak) through central Colorado (Mount Ptarmigan, Colorado Springs, Boulder County, and Northgate) to Wyoming (Sierra Madre Mountains, Albany County, Fort Steele, Muddy Creek, Beartooth Plateau) and north through central Montana (Shriver) and on isolated mountain ranges (Crazy Mountains, Big Snowy Mountains, Little Belt Mountains, Big Belt Mountains) to Saint Mary’s Lake.

Winter range: Winters from southern California (casually to Los Angeles area, and Twenty-nine Palms; Coachella, Holtville), east to southern Arizona (Gila Bend; Pinal County), east to southern New Mexico (Silver City), and western and Central Texas (Frijole in Culberson County; Concho County; Kerr County); south to southern Baja California (San J05~ del Cabo); south and east through Sonora, Sinaloa, Coahuila (Sabinas; Hipolito), Nuevo Leon (Monterrey), Aguascalientes (Aguascalientes), San Luis Potosi (Rio Verde), Guanajuato, Quer6taro (San Juan del Rio); south to Jalisco (Atoyac; OcotIan), and Michoacan (P4tzcuaro).

Migration: Early dates of spring arrival are: Colorado: Salida, April 19; Sulphur Springs, May 2. Montana: Gallatin County near Bozeman, early in May; Bitterroot Mountains 7000 ft., June 11; Flathead Woods, June 5. Saskatchewan: Southern edge of Cypress Hills, May 21. Alberta: Waterton Lakes, May 21. California: Florence Lake, near Fresno in Sierra Nevada, April 20; Kings Canyon Park, May 12; Yosemite Valley, April 17. Oregon: Fort Klamath, April 26.

Late dates of spring departure are: Michoacan: Pátzcuaro, April 26. Baja California: Cape district, May 23. Sonora: Magdalena, May 15. Sinaloa: El Fuerte, May 13. Jalisco: Ojuelos, May 15. Texas: Tom Green and Concho Counties, until May. New Mexico: Apache, southwestern Grant County, April 30. Anzona: Pima County, June 8; Baboquivari Mountains, 3500 ft., May 22; Huachuca Mountains, May 13. California: Buena Park, May 1; Twenty-nine Palms, May 14; Palm Springs, April 26.

Early dates of fall arrival are: Arizona: San Francisco Mountain region, September 21. Texas: Concho County, mid October. Baja California: La Grulla, September 28. Sonora: Cajon Bonito Creek, September 8.

Late dates of fall departure are: Oregon: Southern Willamette Valley, early October. California: Eagleville, September 18; Panther Creek and Shasta River, late September; Florence Lake, September 1; Charlotte and Bullfrog Lakes, Kings Canyon, September 5. Saskatchewan: Cypress Hills, July 11. Montana: Upper Holland Lake, Missoula County, September 10; Gallatin County near Bozeman, mid October. Idaho: St. Joe National Forest, October. Utah: Uinta Basin, (October 5: 25, date not specified) ; southern Utah, October. Colorado: descends to lower levels September: October, lingers until early November. Rocky Mountain National Park, October 26.

Egg Dates: California: 155 records, May 30: August 7; 125 records, June 10: July 5. Nevada: 7 records for June 22. Oregon: 1 record for July 13. Alberta: 2 records for July 4 and July 6.

PUGET SOUND WHITE-CROWNED SPARROW

ZONOTRICHIA LEUCOPHRYS PUGETENSIS Grinnell

Additional material on this race will be found in the account of Nuttall’s white-crowned sparrow.

Contributed by BARBARA BLANCHARD DEWOLFE

HABITS

The Puget Sound sparrows that compose the subject of this section are those that winter at Berkeley on the same ground with the resident Nuttall’s sparrows and those that breed at Friday Harbor, Wash. Identity of these populations is not implied, although Ernest D Clabaugh (1930) reports the capture in summer at Victoria, B.C., of a Puget Sound sparrow he banded in Berkeley the previous winter.

In addition to these variations, Puget Sound sparrows differ from Nuttall’s sparrows in the timing and duration of the phases of the breeding cycie, in the average clutch size, and in the age when the young become independent of their parents. The strongly migratory populations of pugetensis nesting at the Canadian border compress the active part of their reproductive cycle into less than two-thirds the time consumed by the Nuttall’s sparrows of central California. In 1936 the Puget Sound sparrows at Friday Harbor paired, established territories and fledged three broods in less than 4 months, whereas from 1935 through 1938 the Berkeley Nuttall’s sparrows consumed from 6 to 6½ months each year to achieve the same fraction of the cycle. The compression is accomplished by the omission of one and the shortening of three other more or less sharply delimited phases. In pugetensis, territorial establishment requires about three weeks and is not complete until the first day of incubation. In nuttalli the same process is begun much earlier, and also requires about three weeks, so that territory establishment is finished some six and one-hall to eight and one-hall weeks before the first day of incubation. The ensuing period of relatively settled conditions, when chasing and fighting have either decreased or disappeared, does not occur in pugetensis. The three phases that are shortened are: the period of temporary abeyance of song (14 days in pugetensis in contrast to 46 to 59 days in nuttaili, depending upon the year) the interval between completion of the first nest and laying of the first egg (an average of 2.6 days for seven records in pugetensis compared with 3.6 days for 12 records in nuttalli) and the interval between fledging of one brood and laying of the first egg for the next (an average of 8.9 days for five records of pugetensis in contrast to 20 days for six records in nuttalli).

Spring: The same behavior elements as those described for nuttalli appear in spring in pugetensis: increased force and frequency of singing, pursuits and fights, and trilling and posturing. Only the first element is manifested on the wintering grounds. The others are delayed, either until migration or more probably until arrival on the breeding grounds. Then both sexes of Puget Sound sparrows regularly engage in chasing and fighting and in trilling and posturing, with much greater intensity than occurs in nuttalli.

In late Feburary, about four weeks before migration, a few Puget Sound sparrows begin to sing with greater vigor and more frequently than in winter. From then on until late March and early April an increasing number of birds in a given flock participate in the singing until, on the eve of departure, one hears a chorus of song from each flock headquarters. This group singing is not an act of defiance, and provokes no hostile response either from other Puget Sound sparrows or from the by now highly territorial Nuttall’s sparrows in whose areas the migrants live. I never saw either chasing, fighting, or trilling and posturing in the flocks of Puget Sound sparrows wintering at Berkeley.

Daily trapping of banded birds reveals that it takes about as long for the individuals of any one flock to depart as it does the entire local wintering population. The departure is spread over about two weeks. Adults tend to depart earlier than young of the year. The last members of a given flock to disappear are usually first-year females. From departure and arrival dates for this race at the extremes of the range, I believe that the longest spring migration is accomplished in not more than two weeks. I know nothing of the behavior during this period.

In the spring of 1936 I went to Friday Harbor, Wash., to watch the arrival of birds destined to breed there. The largest influx occurred between the evening of April 9 and the next morning. On April 10 I found almost every piece of suitable ground, which only the day before had been empty and silent, occupied by a male perched conspicuously and singing with full volume every few seconds. Close to him I often saw another, silent bird, the female. The shift from the unassorted gregariousness of the wintering grounds to paired isolation must have taken place either during migration or immediately upon arrival, for no flocks or fragments of flocks were in evidence. Observations on color-banded birds by Mrs. Forrest Fuller at Friday Harbor indicate that on the day of arrival the birds go directly to the areas where they subsequently breed, and that they return to the same spot in successive years.

From the day after arrival both chasing and fighting were common and continued unabated until the beginning of incubation about three weeks later. Territorial boundaries were so fluid as to scarcely deserve the name. Both sexes seemed more excited and restless than Nuttall’s sparrows in the corresponding stage. During a boundary dispute neighboring males trilled vigorously and fluttered one wing, and females frequently took part in pursuits. The whole scene was one of confusion and unsettledness, in sharp contrast to the orderly and law-abiding Nuttall’s on their mutually exclusive areas during the seven weeks before the start of incubation. Eventually, however, the same result was achieved. By tbe first day of incubation in late April and early May the fighting subsided and the pairs settled down to a routine identical with that of Nuttall’s sparrow.

The behavior, then, of pugetensis in spring differs from that of nuttalli, in the interpolation of a migratory flight between the beginning and the end of the transition from flocking to paired isolation, in the time the three behavior elements appear relative to the start of breeding, and in the intensity with which they are manifested. The relative disorder among the migrants during the “shake-down” period is not unnatural and may represent merely the result of circumstances rather than an intrinsic variation in habit or temperament. The migrants must establish themselves on ground unseen for seven months and perhaps wholly unfamiliar to some of the adult birds. The first-year birds must seek and seize at once what the young Nuttalls drift into or seek at leisure. The sudden change from a gregarious to an isolated habit perhaps creates a deeper psychological disturbance and contributes to the violence of the process of spatial arrangement.

Courtship: The Puget Sound sparrows arrived at Friday Harbor apparently already paired. On Apr. 12, 1936, two days after the main influx, I saw a female trill and posture. From this time on, both actions increased in force and frequency until April 21, when I first saw a pair copulate. By April 27 trilling and posturing ceased temporarily, to be resumed prior to the start of the second brood.

We have evidence that, as in nuttalli, the members of a breeding pair of Puget Sound sparrows may remate the following season. Mrs. Forrest Fuller (pers. comm.) observed the same color-banded pair nesting in the same hedge in her garden in 1936 and 1937.

Nesting: At Friday Harbor in 1936, the females started to build the nest one or two weeks after their arrival. Behavior during nestbuilding and the time it took to finish the nest were the same as in the Nuttall’s sparrow, but the interval between fledging the first brood and starting work on the second nest was shorter than in nuttalli. Six Puget Sound sparrow females that I followed started work on their second nests only two to four days after the first brood was fledged.

The female alternates between feeding her young and building the new nest. That this may result in confusion is illustrated by the behavior of one female I observed on the morning of May 28 (Blanchard, 1941):

* * * I saw her disappear with material into the hedge a few feet to one side of the first nest site. There I subsequently found a new nest with the walls partly formed. * * * Between half-past eight and half-past nine she made seven trips to the hedge to feed the fledglings, then stayed away for half an hour. She returned at ten, carrying straws in her beak, flew to the abandoned nest, hesitated a little, flew to the hedge where a nestling was perched, and emerged without the straw. I sought out the fledgling, which was crying for food, end found she had laid the straws at its feet!

About fifteen minutes later, female X brought food to the fledgling * * Five minutes later she again flew to the hedge with straws in her beak and this time went straight to the new site. When I looked at the nest that evening, its walls were complete, and a few pine needles and a little fine grass were heaped in the cup. On May 31, just six days after the first brood had left the nest, this female laid the first egg of her second clutch.

Eggs: The interval between completion of the nest and the date the first egg is laid is shorter for pugetensis than for nuttalli. For seven records the interval ranged from one to four days and averaged 2.6 days, as compared with 3.6 days with a range of one to seven days for 12 records for nuttalli.

In pugetensis the eggs are laid between late April and late July. The earliest date recorded is April 23 at Eureka, Calif.; the latest date, July 23 at the same locality. (One clutch in the Museum of Comparative Oology collection at Santa Barbara is labeled “Oct. 31, 1908” but the condition of the eggs is not given.) At Friday Harbor in 1936 the earliest date for the first egg of a clutch was April 24, the latest, July 4. For first clutches ~of the season, the median date for 22 females at this locality was Apr. 29: 30, 1936. That year one color-banded female laid her first egg on May 1, only one or two days from the median date for the population. The next year she laid her first egg on May 7. If she again came as close to the median date for the population, then in 1937 the median date would have fallen between May 5 and 9. This annual difference, if accurately estimated, would be less than a third as great as the difference in median dates for Nuttall’s sparrows the same two years at Berkeley.

The clutch size in pugetensis averages greater than in nuttalli. rrhe number ranges from three to five, and the average of 90 clutches is 3.87, compared with the average of 3.27 for 215 nuttalli clutches. The most common number of eggs per clutch is four, as compared with three in nuttalli. About two-thirds of the clutches in pugetensis had four eggs each, whereas in nuttalli about two-thirds of the clutches had three eggs each. The percentages for pugetensis are as follows: 3 eggs per clutch 23.3 percent, 4 eggs per clutch 66.7 percent, 5 eggs per clutch 10.0 percent. The average clutch size for all records, by month, is as follows: April 3.77, May 3.98, June 3.82, and July 3.25 for 22, 47, 17 and 4 clutches, respectively.

The second clutch averages larger than the first. Among 22 first clutches of eggs found at Friday Harbor in 1936, 3 had three eggs each, and 19 had four eggs. The average was 3.8. Of 7 second clutches, 4 had four eggs each and 3 had five eggs. The average was 4.4. A single third clutch had five eggs.

Clutch size varies with latitude in pugetensis. 32 clutches collected at latitude 41°N. average 3.56 eggs per clutch, whereas 48 clutches taken at latitude 49°N. average 4.04 eggs per clutch.

Incubation: The start of incubation in relation to egg laying shows about the same variation as in nuttalli. The length of the incubation period is the same, and the eggs of any one set show about the same variability in hatching time as do those of nuttalli. I marked the eggs of three sets at Friday Harbor. They did not hatch in the order laid. One set of three hatched in the sequence 2, 1, 3, another 3, 2, 1, and a set of four in the order 2, 1, 3, 4.

Young: The behavior of the parents up to the time the young leave the nest is identical with that of Nuttall’s. For about the first week after the young are fledged the female alternates between feeding them and building the second nest. Thereafter the male takes over all care of the fledglings. On an average of 11 days after one brood has been fledged, the female has begun to sit on her second clutch. The male then performs the duties of feeding the fledglings and guarding the territory and the second nest. He makes short excursions with his young outside the territory, but never for more than a few minutes at a time. Once I found three males from contiguous areas foraging with their young in the same 50-yard-square raspberry patch. None appeared to resent the presence of the others.

As the time for the hatching of the second brood drew near, the males spent more and more time near the nest and less in feeding and watching the fledglings. The young birds left their parents’ territories at an earlier age than did the young Nuttall’s. One young bird was found alone across the border from its parents’ territory when only 25 days old. Out of seven broods that I watched almost daily, the oldest fledgling fed by the male was 27 days old.

Owing to the compression of several phases of the cycle, particularly of the interval between broods, the Puget Sound sparrows, although they began to breed more than 6 weeks later than the Nuttall’s, had time to raise three broods before the regression of the testes in late July. Unfortunately I was obliged to leave Friday Harbor before recording the fledging of the third brood. Two females that had fledged two broods each were sitting on their third clutches when I left. All but 2 of the remaining 11 pairs of the group I watched would have had ample time for a third brood before gonad regression as recorded at other stations of like latitude. The general average of success with the broods was not high. Before I left, only 3 out of 13 pairs raised two broods successfully, whereas each of the remaining 10 had raised one brood and made one or more further unsuccessful attempts.

Fall and Winter: Most of the Puget Sound sparrow populations leave their nesting grounds in the fall and form large flocks which persist for 6 or 7 months. I have no first hand knowledge of their behavior in early fall before migration, nor during the flight south. This account begins with the arrival in California of the populations that winter at Berkeley on the same ground with nuttalli. The quotations below are from Blanchard (1041).

In late September and early October flocks of from twenty to fifty birds reach Berkeley, announcing their arrival by a chorus of song which is unmistakably distinct from that of the Berkeley Nuttalls.

Arrival dates are not identical for successive years, nor do all birds come on the same day in any one year. From 1934 to 1937 I saw the first Puget Sound sparrows on the campus on September 27, 30, 13, and 23, respectively. In 1935 a few birds came September 30, but it was not until October 5 that the first large flock arrived. In 1936 I heard a Puget Sound sparrow sing on September 13, and another observer heard one on the sixteenth, but I saw no large flocks until the twenty-sixth. In 1937 the period of arrival extended from September 23 to October 5.

In spite of the fact that one pair, at least, remated for two or more years on the same territory at Friday Harbor, I have detected no tendency to remain paired on the wintering grounds, no hint of such persistent mutual awareness as might reveal, within the winter flock, mates or nest mates of the past season. The flock is a homogeneous assemblage, without visible subdivision. It is more coherent than the smaller groups of immature Nuttalls. The latter are loose aggregates—individuals foraging on common ground, unified only by fright. The flocks of Puget Sound sparrows show a continuous simultaneity, a common responsiveness, producing true flock reactions.

Once on the wintering grounds the migrants show almost as strong a tendency to localization as do the residents. Year after year four flocks of Puget Sound sparrows have settled, each on a different area on the campus, and from the day of their arrival to that of their departure six months later have never left it but have foraged and roosted regularly at the same spots. Restriction to mutually exclusive areas is the rule, as has been proven at Palo Alto by the banding records of Price (1931). In February, 1928, by banding and by painting the tail feathers of Puget Sound sparrows on the Stanford campus, Price worked out the areas of three flocks and found almost complete mutual exclusiveness.

The adult Nuttalls are perfectly tolerant of the Puget Sound sparrows for the whole winter season. Even in spring, when the resident males are ready to fight and drive out other male Nuttalls, they allow the pugetensis flocks to wander unmolested through their territories and may even forage peacefully in the midst of the flock, set off from the rest by distinctions which can hardly be based on physical appearance. Differences in individual and flock behavior and in degree of aggressiveness undoubtedly account for the perfect cleavage between the groups. The larger flock of silent birds, obviously with no interest in the ground besides the food they are seeking, suggest no threat, rouse no jealousy. When a Nuttall landowner chances to hop or to fly straight at them, the Puget Sound sparrows always give ground, even to the extent of relinquishing a morsel of food too large to carry off.

* * * Up to the day of departure, the behavior of the wintering flocks suffers no radical change except for the increased frequency of song, beginning some four weeks before migration.

* * * Neither chasing nor fighting, other than occasional momentary disputes over food, occurs on the wintering grounds. There is no stimulation of trilling and posturing in the female. Neither territorial jealousy nor sexual excitement have yet come to the surface. The coherence of the flock is maintained up to the moment of departure.

DISTRIBUTION

Range: Southwestern British Columbia to extreme southwestern California.

Breeding Range: The Puget Sound White-crowned sparrow breeds from the east coast of Vancouver Island (Quinsam River) and from the southwest mainland of British Columbia (Vancouver, Mission City) south, west of the Cascade Range through Washington (Nooksack Valley, North Bend, Chehalis, Cathlamet) and Oregon (Logan, 50 miles inland along the North Santiam River, Corvallis) southwest into extreme northwestern California (Smith River, east of Eureka) south to Weott and west to Cape Mendocino.

Winter range: Winters from southwestern British Columbia (casually to Comox and Victoria); but more commonly from Oregon (southern Willamette Valley; Coquille) west to the coast; east to Dudley, Mariposa County; south to southwestern California (San Nicolas Island; La Jolla).

Casual records: Casual in winter in Oregon (Crater Lake National Park: east entrance), Washington (Puyallup Valley in Pierce County) and California (Dudley, Mariposa County).

Migration: Early dates of spring arrival are: Oregon: Tillamook, March 18. Washington: Seattle area, March 24; Friday Harbor, April 4; Bellingham, March 26. British Columbia: Colwood, Vancouver Island, last week of March; Vancouver, April 2.

Late dates of spring departure are: California: Sebastopol, May 1; Berkeley, April 21.

Early dates of fall arrival are: California: Sebastopol, September 3; Woodacre, September 21; San Nicolas Island, September 24.

Late dates of fall departure are: British Columbia: Alta Lake region, September 17; Colwood, Vancouver Island, first week of October; Vancouver, October 15. Washington: Friday Harbor, September 21 (2 years record); Seattle area, October 3. Oregon: northwestern Oregon, October 1; Netarts, September 10.

Egg dates: British Columbia: 6 records, May 4: June 26.

California: 31 records, April 23: July 23; 9 records, April 28: May 8; 12 records, May 11: May 31; 8 records, June 6: June 26.

Oregon: 5 records, May 5: June 6.

Washington: 46 records, April 24: July 22; 22 records, April 27: May 6; 7 records, May 27: June 9; 3 records, June 24: July 22.