Spotted Towhee

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

This and the following 11 subspecies, plus some 8 more races resident extralimitally in Mexico and Guatemala, form a complex commonly called the spotted-backed towhees. All show varying amounts of white streaks or spots on the scapulars and wing coverts, which occur very rarely in the four eastern races treated in the preceding account. In addition, females of these western races have the black of the male replaced by dark gray instead of the reddish brown of eastern females. Until recently they were regarded as specifically distinct from the eastern forms and grouped together in the species Pipio maculeztu.s. In suggesting the conspecificity of the mdculatus and erythrophtkolmus groups, Charles G. Sibley (1950) notes that “many specimens of otherwise typical P. e. erythrophthalmws have varying amounts of white spotting on the same feathers which are normally spotted in the races inhabiting Mexico and the western United States” and adds:

1. In all plumage characters other than the dorsal spotting, the two are identical except for normal geographic variation.

2. Eggs and nests are no more different than is to be expected between subspecies.

3. Ecological requirements vary more within the spotted races than between the spotted and the eastern ones.

In a later paper, Sibley (1959) suggests:

The geographical distribution of the dorsal color pattern suggests that it has adaptive significance in some way correlated with climate. The races which are spotted dorsally (mecutntus group) tend to live in areas which are more arid than those occupied by the unspotted erythropthalmus group. The vegetation occupied by the spotted races is usually a “chaparral” formation of woody shrubs without an arboreal cover. The unspotted races tend to occupy the understory shrubbery of eastern deciduous woodlands, a formation of more humid climates. Common observation indicates that the amount of sunlight reaching the ground and producing a sun-dappled pattern will be greater in a chaparral habitat than in a woodland habitat where the canopy will intercept more of the light. Hence we suggest that the dorsal spotting is a cryptic pattern induced by selection through predation and correlated indirectly with climate through the effects described.

In appearance arcticus is closest to P. e. montanus, from which it differs chiefly in having heavier white dorsal spotting and a more olivaceous back. In his original description of the form Swainson (1831) states he observed it “only on the plains of the Saskatchewan,” where it frequented shady and moist clumps of wood, being generally seen on the ground. He says “It feeds on grubs, and is a solitary and retired, but not a distrustful bird.” P. A. Taverner (1926) remarks:

“Like the Eastern Towhee, the Spotted is a bird of the brush and almost identical with it in general habits. To those familiar with the former, the latter presents nothing strikingly new. The notes are similar enough to be recognized as a Towhee’s, but with a sufficiently different tone and accent to attract attention. On the whole, the Spotted Towhee’s voice is hoarser, and its song less clearly musical than that of its eastern relative.”

E. S. Cameron (1908) states that in Custer and Dawson Counties, Mont., the average date of the towhee’s arrival is the second week in May, and its departure for winter at the end of September. He mentions sage brush as the common nesting site; on June 20, 1898, he found a nest containing five eggs of the towhee and two of a cowbird. A. A. Saunders (1921) adds that in Montana this race breeds in June in the Transition Zone in thickets of willow, wild rose, and other shrubs. J. C. Merrill (1881) found it abundant in Montana wherever a stream with bordering underbrush afforded the needed shelter. He writes: “There is a great diversity in the time of laying, or rather in the contents of nests found on about the same dates from the middle of May until late in July, which I attribute more to the great number of nests that must be destroyed by snakes, birds, and small mammals, and to the attempts of the parents to raise another brood, than to any other cause.” He found nests on the ground usually under a bush, often a cherry bush. He describes them as strongly built, with an internal diameter of about 2% inches; the rim is flush with the ground, the birds scratching a hollow large enough to contain it; externally it is made of dead leaves and broad strips of bark, then a wall of finer strips of bark and blades of dry grass, and finally a lining usually of yellow straws.

Wilbur C. Knight (1902) mentions two nests at Newcastle in extreme northeastern Wyoming which were built on the sloping sides of a canon about six feet from the bottom beside small rocks. The nests were made of pine needles and lined with fine grass.

Eggs: The Arctic Towhee lays from two to five eggs with three or four being the most common. They are short ovate or ovate, only slightly glossy, and practically indistinguishable from those of the eastern races. The ground is grayish or creamy white, sometimes very pale greenish white, and generously speckled all over with “russet,” “Mars brown,” “chestnut brown,” or “Carob brown,” with undermarkings of “light purplish gray” or “light neutral gray.” The markings on most eggs tend to become concentrated toward the larger end, where they are often so thick as to obscure the ground. Although some eggs are blotched, in most instances the markings are numerous, well-defined, small spots or speckles.

The measurements of 50 eggs average 24.1 by 18.0 millimeters; the eggs showing the four extremes measure 9~7.0 by 1~.9, 24.9 by 19.3, ~2l.1 by 17.3, and 23.4 by 17.0 millimeters.

DISTRIBUTION
Range: Great plains from southern portion of the prairie provinces south to central northern Mexico.

Breeding range: The Arctic towhee breeds from central Alberta (Fort Saskatchewan), central Saskatchewan (Canton), and central northern North Dakota (Turtle Mountains) south, east of the Rocky Mountains, to southeastern Wyoming (Laramie), northeastern Colorado (Wray), and central northern Nebraska (Long Pine).

Winter range: Winters from Colorado (Boulder) and Kansas (St. John, Lawrence) south to southwestern New Mexico (Deming), central Chihuahua (Chihuahua), central Nuevo Le6n (Monterrey), and southern Texas (Laredo, 15 miles west of Bastrop).

Casual records: Casual west to Utah (Provo) and Arizona (Camp Verde) and east to Minnesota (Madison), Iowa (Woodbury and Plymouth counties), Illinois (North Evanston), New York (Bronx Park, Jones Beach), New Jersey (Metuchen), and North Carolina (Fayetteville). Sight records (presumably this race) southeast to Tennessee, Mississippi, Louisiana, and Arkansas.

Egg dates: Alberta: 5 records, June 10 to June 22. Montana: 2 records, June 14 and June 19.

SPURRED TOWHEE
PIPILO ERYTHROPHTHALMUS MONTANUS Swarth
HABITS

Contributed by TRAVIS G. HAWS and C. LYNN HAYWARD

The spurred towhee is a common and characteristic bird in central Utah, particularly along the western bases of the Wasatch Mountains. Its most favored habitat seems to be large clumps of Gambel’s oaks, Quercus gambel~i, that grow in scattered patches on the hot, dry slopes. Here the birds may be seen the year round, although in winter they may drift downward into the valleys or more protected canyons where they can scratch for food among the ground debris.

Broadly speaking, Utah marks the center of the east: west breeding range of this race, which the 1957 Check-List defines as extending from central eastern Califronia, southern and central eastern Nevada, northernUtah, and northern Colorado south to southeastern California, southern Arizona, northeastern Sonora, northwestern Chihuahua, and central New Mexico. In southern Arizoi~ia and adjacent Mexico, Joe T. Marshall, Jr. (1956) notes that while the towhee spans several vegetation zones, it is partial to bush growth, particularly chaparral and brush within woods or forests at altitudes above the deserts.

More recently Marshall writes us from Arizona that this race “is a common breeding bird in the mountains here wherever any brush or bushes grow under the trees or beside creeks, but only in the zones well above the desert. It breeds in patches of manzanita among the oaks, in manzanita or ceanothus under the pines or among the pinons, and in snowberry bushes and under aspens in the high forests. It is most abundant where true chaparral grows, as on the Pinal Mountains near Globe in central Arizona. In the Rincon and other southem mountains it is plentiful only where heavy brush growth has replaced the forests after fires or logging operations.”

In the common with other Pipilos, the spurred towhees are inseparable from dense cover throughout their range. They are rarely seen in the open except as they may be flushed from one bush and fly for the closest thicket, usually close to the ground. In spring and summer the casual observer will most likely see the male as he sings from the top of a favored clump of oakbrush or similar shrub. When too closely approached, the male invariably darts downward into the thickest cover available. The female is less often seen, for she usually stays on or near the ground where she rustles among the dried fallen leaves in the dense underbrush, or goes quietly about the tasks of nest-building, incubating, and brooding. Rarely is either sex seen in long, sustained flight.

Spring: About the first of April the winter flocks of towhees break up and the birds pair and establish their nesting territories. By mid-April the males sing vigorously. At this time the leaves have not yet appeared on the oakbrush or other deciduous shrubs, and the males are often conspicuous as they sing from vantage poi.nts on the tops of shrubs where they can overlook their domains. By the first of May when most of the nesting activity is well under way, leaves have usually appeared on the shrubs and the birds have ample protective cover.

Courtship: J. T. Marshall, Jr. (1957) comments that the territories the towhees battle over in spring seem rather small. He has sent us in a letter the following excerpts from his field notes ddscribing a territorial conflict he witnessed in an abandoned orchard within the pine forest on El Tigre Mountain in Sonora on Apr. 4′, 1953: “Two males battling over a female. All three birds squalling loudly. Real fight in clump of locust, banging wings, flying at each other, tumbling down, calling. Then out to middle of flat. Ticking call and mew call both used. Female joins them and both males begin to sing. Intruder male moves off to north and owner follows him, both still singing from high up in the trees. Female squalling from middle of orchard. Intruder male sings farther off to north for quite a while. Original male returns and seen later scratching in oak leaves around house. No more singing.”

Nesting: In Utah nesting begins in April. The female builds her nest in such great secrecy that we have never been able to witness the entire process. We have seen females carrying nesting material, and observed one just starting to build a nest under a small sagebrush Apr. 21, 1938. All the nests we have found were on the ground under low bushes, usually sagebrush, and not in the dense, taller thickets of scrub oak from which the males do most of their singing. The nest is placed in a depression with the rim level with the ground surface. It has an outer coarse shell of dried oak leaves and bark and is lined with finer grasses. The cup ranges in diameter from 7 to 9 centimeters and in depth from 5 to 6 centimeters.

A summary of nesting dates for the vicinity of Provo, Utah, is as follows: May 5, nest and four fresh eggs found under a sagebrush; May 6, nest and four fresh eggs under a sagebrush; May 29, nest with four eggs well advanced in incubation under a sagebrush, these hatched June 2. For this same general area R. G. Bee and J. Hutchings (1942) give nesting dates of May 20, May 25, June 1, June 10, and June 26.

Eggs usually number four, but we have seen as many as five and as few as three apparently forming a full clutch. Their color, using Ridgway’s Color Standards, is near pale olive gray but lighter, and finely speckled or spotted with army brown and sometimes blotched with light mouse gray. Most of the eggs are heavily pigmented at the large end.

The measurements of 40 eggs average 23.6 by 17.8 millimeters; the eggs showing the four extremes measure ~5.9 by 18.5, ~2O.8 by 18.3, and 23.9 by 16.8 millimeters.

Incubation is apparently carried out entirely by the female. She sits very closely during that period and may be approached within a few feet or even a few inches before she will flush. She leaves the nest by slinking rapidly over the ground to the nearest cover, and her movements are so swift she may easily be mistaken for a small mammal. She returns to the nest the same way, never by direct flight to the site. Insofar as we have been able to observe, the male does not approach the nest during the incubation period. We have not been able to determine the length of the incubation periods in P. e. montanu.s accurately, but our meagre information suggests that it agrees with that of the eastern subspecies, P. e. erytkrophthalmus, recoi~ded by F. L. Burns (1915) as 12 to 13 days.

Young: On June 2, 1956, we were able to observe the hatching of the young and their development through the first week. At 8:00 a.m. on June 2, two eggs had hatched and the third was pipped. When the nest was revisited about six hours later, the young of the third egg was dead half out of the shell and the fourth egg was pipped. By 7:30 p.m. the fourth egg had hatched successfully. The empty shells are removed apparently soon after hatching, though we were unable to observe the process.

The young at hatching were naked except for a few down feathers along the spinal and capital pterylae. The day after hatching the feather papillae had darkened and enlarged noticeably. By the third day the sheaths were emerging, especially on the wings, and by the sixth day they had begun to open. In common with most passerines, the growth of the young towhees is very rapid. The weight of these nestlings increased about fourfold during the six days of observation.

Brooding is seemingly entirely by the female. At first her periods off the nest are brief, but they become more sustained by the fourth or fifth day. Feeding the young is apparently largely the responsibility of the male. On June 6 a male was observed to feed the young six times during a 160 minute period. Each time he approached the nest he gave the pshew call, softly and muffled by the food in his beak. The female left the nest immediately and he fed the young. After feeding them he usually returned to his perch to sing for awhile before setting out to scratch for food and feed the young again.

Voice: Tn addition to the spring song of the male, which shows considerable variation from individual to individual, the spurred towhee has two distinct calls which vary but little and are used by both sexes. The first of these we designate as the tseep note. The female gives it as she approaches the nest containing young, and it is the first sound the young make. It is not very loud, and can be heard only at distances of a few feet. It is heard most commonly during the winter as the birds move about through the shrubbery and underbrush.

The p.s&w call is uttered commly by both sexes at all seasons. This is a scolding call, and usually indicates disturbance of some sort, but it may also be heard when the birds are feeding and apparently unexcited. At the close of a period of vigorous singing, the male will often fly down into his shrubby habitat and call [pshew steadily for a considerable period. Frequently, and especially during the mating season, the pahew call is answered by several nearby males.

The spring or mating song, given by the male only, consists of one to five introductory notes followed by a trill usually higher than the introductory tones, sometimes at the same pitch, more rarely at ~lower one. Singing starts in Utah late in February or early in March, and continues until after the young leave the nest, but is most intense in April and May. Song is most pronounced on clear, warm days. Males start to sing at the first indication of daylight and continue singing intermittently throughout the day until dark, with two apparent periods of intensity, one just before, and the other just after sunrise.

The longest sustained period of singing by a single bird, heard soon after sunup on Apr. 14, 1955, lasted 42 minutes. In this time the bird sang 260 consecutive songs involving 6 variations. The males continue to sing while the females are brooding, while the young are in the nest, and between their own periods of feeding the young.

J. T. Marshall, Jr., writes us from Arizona: “There are differences in the songs in different areas, and each male has several different songs. When a male uses a particular song, those near him often respond with the same song. The most frequent pattern consists of two sharp notes followed by a trill, such as clip-clip-c&eee. One bird in northern Sonora changed to a song exceedingly like a brown towhee’s for several minutes, and then changed back to his own again. A common variation you hear in Sonora is the trill preceded by a whistle like the call of the evening grosbeak, which is very confusing.”

Summer: Our observations in Utah indicate that after the young leave the nest the family stays together near the nesting site all summer. Young of the year in full juvenal plumage are seen regularly in July. As the singing of the males diminishes after the nesting season, the birds become relatively inconspicuous as they forage quietly in the dense cover of their preferred habitat.

In the southern part of its range this towhee is apparently doublebrooded. J. T. Marshall, Jr., writes us from Arizona that “in mid-July in the Pinal Mountains independent juveniles were everywhere in the brush within the aspens and firs. The males were singing all day, and were often seen chasing the juveniles as if to get them out of their territories.” He sends us the following notes from northern Mexico: “Northern Sonora mountains, July 8: female carrying nesting material and another feeding young just out of the nest; July 16: juveniles fully grown; July 28: a pair hovering in alarm around juveniles just out. Both adults and juveniles had the same call note, a piercing chip so ventrioquial you could not locate the birds by the sound. Chihuahua, August 22: independent juveniles in complete juvenal plumage; August 25: juvenile male half molted into the rich adult plumage.”

Food: Herbert II. Frost (1947) studied the food habits of the spurred towhees along the Wasatch foothills east of Provo, Utah. lie found that their principal food in winter was vegetable, with the fruits of the hackberry (Celtis) most commonly identified. In summer, animal matter increases in the diet markedly. Remains of coleoptera and orthoptera were found in many stomachs. It must be recognized, however, that examination of stomach contents alone may not give a true picture of the bird’s food habits, as die hard fruits of the hackberry add the chitinous exoskeletons of the insects are more likely to remain identifiable than any softer material ingested.

The towhee obtains most of its food by scratching in the leaves and other ground litter beneath the shrubbery in which the birds live. A. M. Woodbury (1933) describes its method of feeding as follows:

The Spurred Towhee (Pipilo maculatus montanus) is a perching bird that has entered the field of scratching to earn a living. In Zion Canyon in Utah it is an inhabitant of the dense thickets of oak, sarvis-berry, squawbush and streamside deciduous trees. It is primarily a ground-dwelling bird, nesting among the thickets and hunting its food chiefly among the trash and leaves, but does not hesitate to ascend the trees and brush at other times.

If the visible food supply on the surface is not sufficient for its needs, the towhee takes to turning over the leaves and scratching among the trash with its feet. This is a complex operation that it is fitted admirably to perform. * *

Scratching birds like chickens stand on one leg and scratch with the other, but not so with the towhee. Being a small bird, it would have a difficult time turning over a leaf with one foot while standing on it with the other. Such difficulties are solved by using both feet in scratching. In order to use both feet, the body must be balanced in the air during the scratching operation.

This is accomplished by jumping into the air and drawing the feet backward while the upward momentum lasts, Drawing the feet backward and raking trash or leaves at the same time tends to overbalance the body forward. The bird uses several methods to hold its balance, either singly or in combination. Nearly always, the scratching motion of the feet is accompanied by an upward and forward jerk of the tail. Sometimes the wings flutter forward, and always after the scratchtag stroke the feet are brought forward quickly to catch the body and keep it from falling. Sometimes a backward movement of the body is made in jumping and the feet rake the trash while the momentum lasts. This is accompanied by a downward movement of the tail. All of these movements are carried on automatically and seemingly with the greatest of ease.

Sometimes, when the jump is made, the feet are thrust forward and trash in front of the bird is caught and pulled backward. Other times material underneath is moved, while occasionally material just behind the feet will be kicked out of the way by vigorous backward strokes.

Sometimes the trash is kept flying by quick successive strokes, but if insects, spiders, or other interesting food items are exposed to the eye of the bird, it suddenly stops and picks up such items one by one. And thus it taps a food supply not available to its competitors in Zion Caflon. On one occasion, I saw a Woodhouse Jay make a dart at a towhee. The smaller bird merely flitted away a few feet and stopped. The jay did not pursue any farther. At another time a gray rock squirrel came nosing around very close to the towhee, evidently paying no attention to the bird. The bird, however, flitted quietly out of the way a few feet and went on scratching.

Winter: In Utah the spurred towhecs begin to gather into small flocks in September. However, the flocks are loosely organized, and many of the birds remain more or less solitary. Flocking seems to result from a tendency to gather in favored localities rather than true gregariousness, and the closeness of the flock depends largely on the size and density of the cover. By about September 21 the postjuvenal molt seems to have been completed, and from then on adults and juveniles cannot be distinguished readily from one another in the field.

Their winter distribution and activities here seem to be governed to a large degree by the amount and duration of snowfall. Usually the lower west: and south-facing slopes of the Wasatch Mountains are free of snow except for short intervals. However, when the ground is covered for longer periods in midwinter and food becomes difficult to find, the birds tend to drift downward to streamside thickets and other suitable cover along ditchbanks and elsewhere in the lower valleys. Ordinarily no marked wholesale exodus occurs from the summer habitats to the valleys, and some birds may be seen regularly in the foothill brushlands throughout the year.

From Arizona, J. T. Marshall, Jr., writes us: “We find a few towhees here in winter high in the mountains, and we also have it as an uncommon winter visitor in the lowlands around Tucson, where it is confined to mesquite woods. On a mountain near Hermosillo, Sonora, where we have never found the bird in summer, we have taken specimens in winter up in the oak zone. The most abundant winter population I have seen was in manzanita scattered among oaks at the east base of the Rincon Mountains, where I recorded about 25 in about % mile on January 21, 1951.”

DISTRIBUTION
Range: Eastern California, Utah, and Colorado south to northwestern Mexico.

Breeding range: The spurred towhee breeds from central eastern California (Benton), southern and central eastern Nevada (Grapevine Mountians, Lehman Creek), northern Utah (Stanshury Island, Unita Mountains), and northwestern and central northern Colorado (Boulder) south to southeastern California (Providence Mountains), central western and central southern Arizona (Harquahala Mountains, Baboquivari Mountains), northeastern Sonora (San Jose and San Luis mountains), northwestern Chihuahau (Sierra Madre, south to lat. 290 N.), and central southern and northeastern New Mexico (Mesilla Park, Sierra Grande).

Winter range: Winters from southern Utah (Beaverdam Mountains), central Colorado (Golden), and western Texas (Palo Duro Canyon) south to northern Sonora (Sierra Carrizal, Nacozari) central Chihuahua (Chihuahua), and central Texas (Del Rio, Kendall County) ; casually farther southeast in Texas (Victoria, Eagle Lake).

Casual records: Casual in Nebraska (North Platte) and Kansas (Morton County).

Egg dates: California: 7 records, April 24 to June 7.

OREGON TOWHEE
PIPILO ERYTHROPHTHALMUS OREGONUS Bell
HABITS

Contributed by OLIVER L. AUSTIN, JR.

The Oregon towhee is the darkest and least spotted of the western complex of spotted-backed Pipilos. Ira N. Gabrielson and Stanley G. Jewett (1940) consider it “one of the most common permanent resldent birds in western Oregon, where every rose thicket and evergreen blackberry patch has its pair of Oregon Towhees. They are present throughout the year so commonly that it is unusual to walk along the bottom lands at any season without seeing a handsome black and white and reddish fellow flirting his tail nervously as he glides to a landing around a clump of bushes or hops about in the thickets.”

In Washington State, Stanley G. Jewett, Walter P. Taylor, William T. Shaw, and John W. Aldrich (1953) say it “spends most of its time in the shelter of the shrubbery of its humid environment. It is found in the brush of abandoned clearings, along roadsides, in burns, coastal gulches, and in swamps. Log heaps in timbered areas often shelter it; and on Mt. Rainier towbees presumed to be of this subspecies were observed in the azalea, mountain-ash, and huckleberry brush.”

The same authors continue:

Brown found a nest and 4 eggs in Kings County, June 29, 1909, and a nest and 3 eggs in Seattle on July 13, 1908. Dates for full sets of fresh eggs are as follows: early, May 4, mean, May 17, late, June 20 * * * Bowels reports a pair feeding young in the nest at Gravelly Lake, Pierce County, August 21, 1933 * *

Burleigh says nests found by him were invariably sunk flush with the ground, at times at the base of an old stub or small sapling, and were quite well concealed by Oregon grape and clusters of ferns. Of 9 nests examined 7 held 3 eggs, two 4 eggs. Ratlibun flushed a female from a nest on the ground in an open spot among scattered tall firs, and alongside a path running through a dense growth of salal. This nest was unusual in that it contained 5 eggs. On another occasion Rathbun found a nest containing 2 eggs only, both heavily incubated. Immature birds are commonly observed into September.

A little patience enables the observer to watch the towhee as it forages among the leaves in the shelter of the brush. It hops industriously over the ground, its whole attention, apparently, given to the work in hand. * * * On catching sight of the observer the bird may retreat to Lhe inner fastnesses, its spotted tail becoming conspicuous as it flies. It sometimes becomes quite familiar, and is a characteristic bird visitor to food tables provided in fall and winter. The trill of the male Oregon spotted towhee is much like that of the Nevada, though Dawson * * * says “the damps of ten thousand winters have reduced his song to a pitiful wheeze.” The call note is a mewing jo-ree.

In the fall the species becomes scattered. Individuals are often found associ ated in flocks with other species such as song sparrows and juncos. Since the Oregon spotted towhee is observably scarcer in winter than at other seasons it seems a fair assumption that some of the birds migrate, but the fact that a large proportion of individuals stay very close to the same area throughout the year is attested by the record of 12 Seattle banded towbees which were trapped and recovered in both winter and breeding season at the same station. * * *

Gordon W. Gullion wrote Mr. Bent of his experiences with this ï race in the Willamette Valley of Oregon, where the birds inhabit “most of the brush piles and blackberry tangles throughout the valley. ï They always seem to occur in pairs, and very seldom is a single bird encountered holding a territory. In fact it rather seems that they pair for life. In the fall of 1946 I banded a pair of towhees at my home.

As they were the only banded towhees in the vicinity, it was quite easy to keep tab on their activities. Whenever one was seen, the other would be found close by. They were seen almost daily from the time ï of banding until late December 1947. Then the female vanished.

The male remained around about another two weeks, then he too disappeared. Within a few days another and unbanded pair came in and took possession of the territory. Now they are also banded, and they maintain the same sort of constant companionship the first pair exhibited.”

Gabrielson and Jewett (1940) state that in Oregon “The eggs are usually laid in May. Our nesting dates extend from May 3 to June 25, although young of the year are always on the wing before the latter date.”

The measurements of 17 eggs average 23.3 by 18.3 millimeters; the eggs showing the four extremes measure £5.5 by 19.5, 24.5 by £0.0, ££.5 by 19.1, and 24.5 by 17.0 millimeters.

DISTRIBUTION
Range: Pacific Coast from British Columbia to California.

Breeding range: T he Oregon towhee breeds, and is largely resident, from southwestern British Columbia (Comox, Chilliwack) south through western Washington to southwestern Oregon (Roseburg).

Winter range: Winters south to northwestern California (Trinidad, Willow Creek); casually south to central (Colusa) and southern California (San Clemente Island).

Egg dates: California: 1 record, June 9. Oregon: 4 records, May 10 to May 14. Washington: 5 records, April 13 to July 26.

SACRAMENTO TOWHEE
PIPILO ERYTHROPHTHALMUS FALCINELLUS Swarth
HABITS

Contributed by OLIVER L. AUSTIN, JR.

Harry S. Swarth (1913) described this race as “Most nearly similar to Pipilo maeulatus megalonyz Baird, from which it differs in weaker foot, with noticeably short, weak, hind claw, in somewhat greater extent of white markings, and olivaceous or grayish rump.”

Joseph Grinnell and Alden H. Miller (1944) point out that within its range “Ihere is some altitudinal movement up mountain slopes after nesting and descent from higher parts of breeding range in winter, but no migration is known that carries birds outside the limits of breeding range.” They describe its habitat as “Chaparral, river bottom thickets, and brush patches in open forests. Among the widespread plant formations of such types available in the range of this race, the Spotted Towhees are to be found especially where there is a good accumulation of leaf litter and humus. For this reason partly dead or dying brush, ravine and river bottoms, and bases of cliffs or of steep slopes are favored situations. If the ground forage beat is thus supplied, sufficient screening plant cover is usually present and nest sites on the ground or in we]l supported vine tangles are also available. Some common plant associates are ceanothus bushes of several species, poison oak, willows, blackberry, cascara, and manzanitas.”

William B. Davis (1933) gives May 1 as his earliest nesting date in Butte County, Calif. Near Fyife in the central Sierra Nevadas on June 8, 1897, Chester Barlow (1901) “found a nest containing three unfeathered young and one egg on a hillside under a bush. By far the prettiest nest found was on June 11 of the same year. The situation was a small clearing in the forest grown up to cedar saplings about two feet high. Beneath one of these reposed the nest and its three eggs, the lining of light grasses setting them off to good advantage. As in the valley this towhee does not nest on the ground entirely, for Mr. Taylor found a nest on June 12, 1897 containing two eggs, placed six feet up in a bush beside a ditch. It was composed of pine and spruce bark and lined with light yellowish grass.”

From the small portion of this subspecies~ range that penetrates into south-central Oregon, Gabrielson and Jewett (1940) state “Patterson (MS.) furnished dates of numerous nests at Ashland between May 2 and June 14 * * * ” The measurements of 20 eggs average 24.2 by 17.9 millimeters; the eggs showing the four extremes measure ~5.4 by 17.8, 23.9 by 18.8, !23.1 by 17.8, and 23.5 by 17.1 millimeters.

DISTRIBUTION
Range: The Sacramento towhee is resident from the interior of southwestern Oregon (Grants Pass, Medford) south through the northern interior coast ranges, the western and southeastern slopes of the Sierra Nevadas, and the Great Valley of California (Hoopa and Mount Shasta to Vacav-ille, and Kings and Tulare counties; Laws, Olancha).

Egg dates: California: 4 records, May 1 to June 12.

SAN FRANCISCO TOWHEE
PIPILO ERYTHROPHTHALMUS FALCIFER McGregor
HABITS

Contributed by OLIVER L. AUSTIN. JR.

Harry S. Swarth (1913) characterizes this subspecies as “Coloration dark; white markings more restricted than in megalonyx but much more extensive than in oregor&u~. Hind claw smaller and weaker than in mega~onyx.” Within its narrow range along the northwest California coast, Grinnell and Miller (1944) consider it a “Permanent resident. Common generally, although sparse in extreme northwestern part of range.” They describe its habitat as:

“Chaparral and forest undergrowth as in other races of Spotted Towhees * * ‘~. Apparently avoids the dense brushlands of the fogswept coastal slopes of Humboldt and Del Norte counties, although P. m. ore gonu~ finds suitable wintering grounds therein some of the less compact tracts of plant growth. Elsewhere Jalcijer occupies heavy chaparral on shaded canyon slopes, as also streamside tangles, low second growths of forest trees and the understory of oak and madrone woodlands. Blackberry vines, willow thickets, baccharis and poison oak brush, and ceanothus and manzanita bushes commonly constitute the essential plant cover.”

Milton S. Ray (1906) notes this race “nests in low bushes, scrub oaks or willows, or among overhanging blackberry vines. I have never found a nest placed on the ground, except once.” Emerson A. Stoner (1931) reports a case of its multiple nesting and reuse of an old nest in Palo Alto, Calif., as follows:

“The first nest was constructed early in May among some geranium bushes growing against a private garage. After the young had been successfully reared and had left the first nest, a second nest was built closer to the house in a hedge of cherry oak some thirty feet from the first one. The young left this second nest about July 10. Within a week after the departure of the young from the second nest, the parents returned to the first nest and successfully raised a third brood therein, the young of the third brood leaving the nest on August 9.”

The measurements of 25 eggs average 23.6 by 17.8 millimeters; the eggs showing the four extremes measure 25.9 by 18.3, 25.4 by 18.5, 22.7 by 18.1, and 23.6 by 16.2 millimeters.

Alden H. Miller (1942) presents the following observations on the bathing habits of this subspecies:

In the long dry summers of coastal central California, chaparral-dwelling species may find water locally scarce except as it collects on foliage from the nightly fogs that blow in from the ocean. Use of this supply for drinking is probably widespread, but its availability for bathing had not been appreciated by me.

The morning of July 29, 1942, was cool and foggy in Berkeley, and on the hillside at my home * * * the trees and bushes were dripping with water. An adult Spotted Towhee * * * came to the feeding tray at 7:15 and ate some of the cracked grain offered there. It was a dejected looking individual, with bare patches of skin showing around the head, for it was in the middle of its annual molt; indeed it left a spotted tail feather behind on the tray. It flew but a short distance, stopping on top of a tangle of baccharis bushes and poison oak. At once it began scuttling about under and over the wet foliage, rubbing against it and shaking down drops from overhead. The wings were half spread and were fluttered in the fashion customary in bathing; also the bird bent the legs, crouching down rather than standing normally erect. It moved about within a radius of about two feet, always in the crowns of the bushes, three to four feet above the ground. After approximately a minute of this the towhee moved on, but it was detected at a distance, perched, fluttering its wings and preening. The bath was not by my standards especially effective, as the bird was only slightly wet, but it had apparently satisfied an instinct at least. All this time there had been a pan of water on the feeding tray, but it was small and fairly deep and evidently was not so stimulating of the bathing reaction as the natural supply of water.

DISTRIBUTION
Range: The San Francisco towhee is resident along the coasts of northwestern and central western California (Smith River south through Santa Cruz and San Benito counties).

Egg dates: California: 2 records, May 7 and May 13.

SAN DIEGO TOWHEE
PIPILO ERYTHROPHTHALMUS MEGALONYX (Baird)
HABITS

Contributed by JAMES E. CROUCH

The rufous-sided towhee of southwestern California was described by Baird in 1858 under the name Pipio megalonyx. The type locality iS Fort Tejon, Kern County, Calif. The distinguishing characteristics of P. e. megalonyz, according to Swarth (1913), are “Coloration very dark, and white markings restricted. Adult male (and sometimes the immature male as well) with the entire back uniformly deep black (except for the usual white markings), the rump being deep black instead of more or less grayish or olivaceous. Hind claw longer than in any other California race of Pipilo maculatu.s.”

The behavior of P. e. megalonyx differs in no important way from that of other populations of the rufous-sided towhee. The bird seldom forages in the open or on bare ground. It shows a strong preference for situations which give a good overhead cover, some lateral cover, and a ground surface well supplied with humus and litter. This provides protection and desirable foraging. These conditions are provided by the coast live oaks (Quei~cus agmjolia), said by Davis (1957) to be “optimal undertree foraging sites for Spotted Towhees.” Other oaks (Q. doiiglassi and Q. lobata) are commonly used, as are willow (Salix sp.) and other streamside species of plants. Rufous-sided towhees are found in chaparral, but mostly where there are clumps of larger shrubs. The chamise (Adenostoma Jasciculatum) and California sagebrush (Artemisia californica) provide good cover, but little litter. Where there are large isolated shrubs of Toyon (Photinia arbutifolia), blue elderberry (Sambucu.s eoerulea), or lemonade berry (Rhus integrijolia), the towhees are likely to be found. Other plants forming good cover and forage areas are coffeeberry (Rhamnus californica), redherry (Rhamnus crocea var. ilicijolia), poison oak (Rhus diversioba), California blackberry (Rubus ursinus), California wild rose (Rosa califor-nica), and coast ceanothus (Ceanothus ramulosus).

In the forested mountain areas such as at Palomar and the Laguna Mountains of San Diego County the rufous-sided towhees are seen mostly in the clumps of brush at the forest edge or isolated brushy areas out in the meadows. Where the forests give way at the abrupt eastern sides of the mountains these towhees are commonly seen in the heavy brush of the upper slopes. Here they are found in the company of black-chinned sparrows (Spizeild atrogularis).

This towhee is reported occasionally from high altitudes. Grinnell (1908), writing of the San Bernardino Mountains, states: “It was found on the south side of the ranges as high as 7,000 feet. A few were seen as far up theSanta Ana as the mouth of Fish Creek, 6,800 feet.” Again Grinnell and Swarth (1913) writing about the birds and mammals of the San Jacinto area of Southern California report towhees at 8,000 feet on Toro Peak on July 1 and at 9,000 feet at Round Valley on July 10. They state that these birds, full-grown juvenals, were “probably far above normal breeding range.”

Cardiff (1956) collected an adult male megalonyx below sea level on Oct. 8,1949, from a growth of arrow-weeds and salt brush along the New River northwest of Westmoreland, Imperial County, Calif. This specimen was identified by Alden H. Miller. Gilman (1903) reports these towhees about a half mile from Palm Springs, Calif.

Eggs: The measurements of 32 eggs average 23.4 by 17.9 millimeters; the eggs showing the four extremes measure 25.1 by 18.9, 23.5 by 17.5 and 23.8 by 16.8 millimeters.

Food: The rufous-sided towhee gets most of its food by scratching in the litter under shrubs and trees. Sometimes it feeds up in trees or shrubs. Seeds, seed capsules, and bracts of miner’s lettuce (Montia perJoliata) are eaten by these birds between April and June, according to Davis (1957). Also, he says that they eat various fruits such as elderberries between July and September, coffeeberries between August 22 and December 24, and that acorns constitute an important part of their diet in the winter. I have seen them occasionally “hawking” for insects and they quite often pick up insects as they forage for seeds.

Voice: In his lengthy study of the song and breeding of the rufous-sided towhee made at the Hastings Reservation in northern Monterey County, Calif., John Davis (1958) states: “Singing usually starts between mid-January and the first week of February, and it comes to an end in early August. in September and October, there is a slight but regular appearance of singing, involving only a few males.

“Singing is widespread by mid-March, but in early April there is a noticeable decrease in the amount of song. By the last week in April, at the time that nesting gets under way, singing is again at a high level. Nesting males sing a higher percentage of the time during incubation than after the young have been hatched. Unmated males are the most persistent singers of all.”

For description and analyses of the various types of song in this race, the manner of its delivery, and its correlation with season, time of day, and the reproductive cycle, the reader is referred to this soundly detailed report.

DISTRIBUTION
Range: The San Diego towhee is resident in southwestern Califor.. ma (Monterey and west slope of Walkers Pass south to Santa Cruz Island, Little San Bernardino Mountains, and San Diego County) and northwestern Baja California (south to about lat 320 N.).

Casual record: Casual on San Miguel Island, Calif., and in southeastern California (Westmorland).

Egg dates: California: 5 records, April 19 to May 22.

SAN CLEMENTE TOWHEE
PIPILO ERYTHROPHTHALMUS CLEMENTAE Grinnell
HABITS

Contributed by RICHARD FOURNESS JOHNSTON

This subspecies was described by Grinnell (1897a). Ridgway (1901) states it is “Similar to P. megalonyx but bill and feet larger (at least relatively) and coloration grayer; adult male with the black of a duller or grayer cast, and the adult female with coloration much lighter * * *ï” Miller (1951b), however, states that he is not able to demonstrate the difference in bill length between clemerdae and megalo’nyx.

This bird is a permanent resident on Santa Rosa, Santa Catalina, and San Clemente Islands off the coast of southern California. It inhabits, according to Grinnell and Miller (1944), “Fairly tall chaparral, especially along watercourses. Wild cherry thickets are favored both because of associated ground conditions and the protection and fruit supply they afford. Towhees also have been noted in cactus patches and in scrub oak and toyon.” It is remarkable that this race is found, among the northern channel islands, only on Santa Rosa Island, and not on Santa Cruz (Miller, 195 ib); these islands are closer to one another than are either to Santa Catalina and San Clemente.

Eggs: The four eggs in the Harvard Museum of Comparative Zoology measure 25.7 by 18.5, 25.3 by 17.9, 25.0 by 17.8, and 25.0 by 18.2 millimeters.

DISTRIBUTION
Range: The San Clemente towhee is resident on Santa Rosa, Santa Catalina, and San Clemente islands off southwestern California.

Egg date: California: 1 record, May 9.

CAPE COLNETT TOWHEE
PIPILO ERYTHROPHTHALMUS UMBRATICOLA Grinnell and Swarth
HABITS

Contributed by RICHARD FOURNESS JOHNSTON

This race of towhee is restricted to northern Baja California and is characterized in the original description (Grinnell and Swarth, 1926) as follows: “Differs from P. m. megalonyx, to which it is nearest both geographically and in appearance, in smaller bill and darker coloration. Color dilferences are most apparent in females, these being decidedly slaty dorsally in umbraticola as compared with the browner tinge seen in female megalonyx. * * * The bill and feet * * * are on the average decidedly blacker than in any other of the western subspecies of Pipilo maeuk~tue.” Morphologic intergradation between the two races is extensive near the northern boundary of the range outlined below for umbraticola.

Grinnell (1928b) says the distribution of this resident race extends “locally, north of latitude 300, from San Rani6n, at the mouth of the Santo Domingo River, north, centrally, to very near the United States boundary, and east in the San Pedro M~rtir section from the seacoast at Colnett to mouth of El Caj6n Cation at east base of the San Pedro M4rtirs; reaches an altitude of 7500 feet toward the tops of those mountains.” The preferred habitat is found within chaparral associations. Deep, shaded ravines are especially favored. In ecology and general behavior umbraticola probably differs little from megalonyx.

DISTRIBUTION
Range: The Cape Colnett towhee is resident in northwestern Baja California between latitudes 320 and 300 N. (Sierra Juarez and Sierra San Pedro MArtir west to the coast).

GUADALUPE TOWHEE
PIPILO ERYTHROPHTHALMUS CONSOBRINUS Ridgway
HABITS

Contributed by RICHARD FOURNESS JOHNSTON

Ridgway (1901) remarks skins of this towhee to be “Similar to P. m. ore gonu.s in restriction of the white markings on the wings, tail, scapulars, etc., but wing and tail much shorter, and hind claw much larger; adult male with the black much duller, dark sooty rather than black.”

The bird, now extinct, formerly was resident on Guadalupe Island, off the coast of Baja California. It apparently inhabited the understory of the cypress (Cupressus) grove, large individuals of which are still to be found on the high part of the island. Howell and Cade (1954) note with reference to this bird, “We found no trace, of course, of any of the endemics considered extinct. Indeed, the complete absence today of shrubs or understory of any kind in the forests of the island make it difficult to believe that the towhee, Pipilo erythrophthalmus consobrinus * * * once existed there, and this utter lack of suitable habitat should convince even the most hopeful skeptic that [this form is] totally extinct.

“The goats continue to be the greatest threat to the biota of the island through their destruction of vegetation.” The towhee was last seen alive in June, 1897 (Thoburn, 1899).

DISTRIBUTION
Range: Extinct. It was formerly resident on Guadalupe Island off northwestern Baja California.

LARGE-BILLED TOWHEE
PIPILO ERYTHROPHTHALMUS MAGNIROSTRIS Brewster
HABITS

Contributed by RICHARD FOURNESS JOHNSTON

Ridgway (1901) says this large-billed, pale spotted towhee is “similar to P. m. megalonyx but wing and tail decidedly shorter, bill larger, hind claw averaging larger, white on outermost tail-feathers decidedly more extensive, and color of sides and flanks much paler (buff-tawny instead of cinnamon-rufous).”

P. e. magnirostris is one of the most highly geographically isolated forms of the species; this is to a certain extent evident in its gross morphological characteristics. The range is restricted in southern Baja California. Grinnell (1928b) states that the bird is a “Common resident of mountainous portions of the Cape district. Appertains to brushy tracts chiefly within the Upper Sonoran life-zone. A few come down to sea level in winter (C. C. Lamb, MS.). * * * Northernmost known station of occurrence, Triunfo * * * southernmost, Miraflores * * *”

DISTRIBUTION
Range:The large-billed towhee is resident in the mountains of southern Baja California (Triunfo, Sierra de la Laguna).

Casual record: Casual at lower levels at Miraflores.

NEVADA TOWHEE
PIPILO ERYTHROPHTHALMUS CURTATUS Grinnell
HABITS

Contributed by WENDELL TABER

Joseph Grinnell (191 ic) described this race as being most nearly like P. e. montanus, from which it differs mainly in being shorter winged and darker colored. (For a lengthy discussion of these races, see Swarth, 1913.)

Grinnell and A. H. Miller (1944) describe its habitat as similar to that of the race montanus, in brushy cover including willow thickets, artemisia, and rabbit brush. In winter in the Colorado River Valley, it occurs in thickets of arrowweed and in atriplex bushes. J. M. Linsdale (1951) states that in eastern and southern Nevada this race is resident in the mountains and higher valleys, but that an appreciable movement takes place in winter to lower valleys and more southern parts of the state.

It is doubtful that the habits and behavior of this race vary greatly if at all from those of other races described in greater detail.

The measurements of 20 eggs average 24.0 by 17.9 milhinieters; the eggs showing the four extremes measure 26’.O by 17.5, 23.9 by 18.9, £~.9 by 17.5, and 24.9 by 17.3 millimeters.

DISTRIBUTION
Range: Southern British Columbia and Idaho to northwestern Mexico.

Breeding range: The Nevada towhee breeds from central southern British Columbia (Lillooet, Okanagan Landing, Robson) and northern Idaho (5 miles west of Cocolalla) south, east of the Cascades, to northeastern California (south to Mono Lake), western and central Nevada (Tybo), and southeastern Idaho (Craters of the Moon).

Winter range: -Winters in part in the breeding range and also south to southeastern California (Potholes), northwestern Sonora (Sonoyta), and southeastern Arizona (Huachuca and Chiricalina mountains).

Egg dates: British Columbia: 30 records, Apr. 21 to July 8; 15 records, May 5 to June 2.

TEXAS TOWHEE
PIPILO ERYTHROPHTHALMUS GAIGEI Van Tyne and Sutter
HABITS

Contributed by KEITH L. DIXON

The rufous-sided towhee of western Texas was separated under the name Pipilo mactdatus gaigei by Van Tyne and Sutton in 1987 during their avifaunal studies in Brewster County, Tex. The type locality of this subspecies, which they named in honor of Frederick M. Gaige, is “southeast of Boot Spring, 6800 feet” in the Chisos Mountains.

In voice and behavior, the rufous-sided towbees occupying the mountain ranges of east-central and southeastern New Mexico, western Texas, and northern Coahuila appear to differ in no important respects from other populations of the species. For the most part, these birds have been reported from forested areas above 6,000 feet, but they are limited either to wooded areas with a brushy understory or to rather dense shrub communities. A feature common to the places I have seen them is the presence of clumps of shrubby canopy from one to several feet above the ground and leaf litter on the surface of the ground. In the Chisos Mountains of Texas, such cover is provided by a variety of shrubs, including Rku~ triobata, Salvia regla, and Cercocarpus eximius, as well as the oaks, Quereus gri~ea and Q. emoryi. On a slope west of the Laguna in the Chisos Mountains in late July 1955, this towhee was found with black-chinned sparrows (Spizella atroguiaris) and Bewick’s wrens (Thryomanee be’wickii) in a shrub growth which included squaw bush (Bhus trilobata), silk tassel bush (Garrya lindheimeri), scrub oak (Querous intricata), and Viguiera stenoloba.

The distribution range of this form, according to Van Tyne and Sutton (1937), extends northward to the vicinity of Cabra Springs in east-central New Mexico. In that area, some 30 miles north of Santa Rosa, Vernon and Florence M. Bailey found this towhee in the oak brush understory of yellow pine timber ~at 7,400 feet elevation in June 1903 (F. M. Bailey, 1910). The form is known from the Capitan and Guadalupe mountains of New Mexico and also from the latter range in Texas, from the Davis (A. P. Smith, 1917) and Chisos mountains of Texas, and the Sierra del Carmen of Coahiula (A. H. Miller, 1955a). Van Tyne and Sutton’s report (1937) of specimens of gaigei taken in late May on Mount Ord and the Glass Mountains, which lie in northern Brewster County between the Chisos and Davis mountains, suggests its breeding there also. However the rufous-sided towhee is absent from the low mountain ranges between the Davis and Guadalupe mountains, as the woodland of juniper and pin~ion there appears too open for it (William B. Davis, pers. comm.).

Spring: A. H. Miller (1955a) reported only sporadic singing of males in the higher parts of the Sierra del Carmen in the period Apr. 4: Apr. 18, 1953, and he noted a preponderance of males present at this time. These males, although not in full breeding condition, appeared to be spaced on territories.

Nesting: Although W. W. Cooke (F. M. Bailey, 1928) stated that this species “nests commonly in June” in New Mexico, the breeding season appears to be a long one. Van Tyne and Sutton (1937) reported that “specimens taken on April 27, 1935, in the Chisos Mountains were obviously breeding,” and my observation of a male gathering food at 5,700 feet elevation on the north side of that range on May 28, 1957, indicates early nesting. Further, I found that singing was not conspicuous in Boot Canyon in the Chisos Mountains (6,500 to 7,000 feet) from July 11: 26, 1955, suggesting that the peak of breeding had passed. However, in the Guadalupe Mountains of New Mexico, Vernon Bailey found a nest with eggs on August 12 (F. M. Bailey, 1928) and A. P. Smith (1917) took “young, barely able to fly” at 6,500 feet elevation in the Davis Mountains on Sept. 10, 1916.

Although Van Tyne and Sutton (1937) spoke of this towhee as being of “common” occurrence in the Chisos Mountains, I recorded it as “infrequent” in the upper Chisos as a whole in the latter half of July, due in part to its restricted habitat distribution. Miller (1955) likewise found rufous-sided towhees “sparsely distributed” in the neighboring Sierra del Carmen.

Winter: The 1957 A.O.U. Check-List uses the expression “resident, in part at least” in referring to the montane distribution of P. e. gaigei. Some evidence for altitudinal migration of this form in the Guadalupe Mountains was provided by T. D. Burleigh and G. H. Lowery, Jr. (1940). In December and January they found P. e. monianus the only wintering towhee (two specimens), and neither of these was taken or recorded above 6,000 feet elevation, whereas gaigei ranged upward to 8,750 feet in June. W. B. Davis took a specimen in pinyon-juniper woodland at 4,400 feet elevation in the Delaware Mountains (south of the Guadalupes) on Mar. 8,1942, and Van Tyne and Sutton (1937) reported a few individuals wintering in live oak groves at 5,000 feet elevation in the vicinity of Alpine. In the area surrounding the Chisos Mountains, however, there appears to be little habitat suitable for wintering rufous-sided towhee, and I know of no records of their occurrence in winter below 5,200 feet. Although dense brush may be found, in the desert lowlands, there usually is no accumulation of leaf litter.

DISTRIBUTION
Range: The Texas towhee breeds, and is resident at least in part, in mountains of central eastern and southeastern New Mexico (Cabra Springs, Guadalupe Mountains), western Texas (Guadalupe, Davis, and Chisos Mountains), and northern Coahuila (Sierra dcl Carmen).