The California Towhee is a common resident in a variety of West Coast habitats, including areas near human occupation. California Towhees make a wide variety of sounds, though none are particularly musical. Singing commences well before sunrise, and often takes place from the top of a tree.
California Towhees are rarely parasitized by Brown-headed Cowbirds. Western Scrub-Jays, which can eat eggs and young, are a greater danger to nesting towhees.
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Description of the California Towhee
The California Towhee is a chunky, grayish-brown towhee with orange undertail coverts and a faintly streaked throat. Length: 9 in. Wingspan: 11 in.
Seasonal change in appearance
Similar to adults.
Coastal sage scrub, brushy areas, and city thickets and gardens.
Primarily seeds and insects, and will visit the ground underneath bird feeders.
California towhees forage on the ground, often scratching with both feet simultaneously in typical towhee fashion.
California Towhees are found in western California and in Baja California. The population appears to be stable.
Bent Life History
Visit the Bent Life History for extensive additional information on the California Towhee.
The California Towhee used to be lumped with Canyon Towhee as a single species called the Brown Towhee.
Pairs typically remain together on their territory year-round.
The song is a series of high notes that accelerates in cadence. The call is a hard “teek”.
Abert’s Towhee has a black face. Ranges generally do not overlap, area of nearest crossover is southern California.
The Canyon Towhee has a reddish crown and more heavily marked breast.
The nest is an open cup of grasses, twigs, and weed stems, usually placed in a shrub or low tree.
Number: Usually lay 3-4 eggs.
Color: White or bluish-white and spotted with brown or black.
Incubation and fledging:
The young hatch at about 11 days and leave the nest in another 8-9 days, but continue to associate with the adults for some time.
Bent Life History of the California 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 California Towhee – the common name and sub-species reflect the nomenclature in use at the time the description was written.
OREGON BROWN TOWHEE
PIPILO FUSCUS BULLATUS Grinnell and SwarthContributed by HENRY E. CHILDS, JR.
This, the northernmost subspecies of the brown towhee, is limited to the chaparral areas of southwestern Oregon and northern Siskiyou County, Calif. In California, John Davis (1951) states it is resident “north of the yellow pine: Douglas fir belt rimming the northern end of the Sacramento Valley, and between the Cascade Range to the east and the Trinity Mountains and their northern continuation to the west.” In Oregon, Ira N. Gabrielson and Stanley G. Jewett (1940) consider it “a characteristic bird of the bushy hillsides in the interior valleys of Douglas, Jackson, and Josephine Counties. ‘where in the lowlands between the Coast and Cascade Ranges in the Umpqua and Rogue watersheds its characteristic metallic alarm note can be heard throughout the year.” In the early 1900’s it was found considerably north of its present range near Corvallis, Oreg., where it no longer occurs.
The same authors state: “Patterson (MS.) has found numerous nests about Ashland and Pinehurst with extreme dates of May 2 and 20.” They describe the nest as “In bushes and trees, usually within a few feet of the ground, made of inner bark, twigs, and weed stems and lined with plant stems, wool, and hair. Eggs: 4 or 5, pale blue, spotted with purplish brown.”
Range: The Oregon brown towhee is resident in the Umpqua River and Rogue River valleys of southwestern Oregon (Roseburg, Takilme, Ashland) and the Klamath River and Shasta valleys of north-central California (Berwick, Hornbrook, Edgewood).
SACRAMENTO BROWN TOWHEE
PIPILO FUSCUS CAROLAE McGregor
Contributed by HENRY E. CHILDS, JR.
In the dry foothills of the western slopes of the Sierra Nevada, the valley quail is the only bird more conspicuous in the brushland avifauna than the brown towhee. The principal use of this land area is for grazing, and the white-face Herefords contrast markedly with the somber brown bird which in great measure depends on the cattle troughs for its water supply. The brown towhee is essentially a resident of the upper Sonoran zone, but it may be found occasionally in suitable habitat both above and below it.
As this race, like petulans, is largely dependent on “edge” habitat, its abundance depends to some degree on grazing. Tall grass and thick brush do not suit it; consequently the thinning of such vegetation by the grazing and browsing of cattle benefits the towbees. This race also has responded to the artificial habitats produced in residential areas. J. G. Tyler (1913) notes that it occurs among the shrubs and trees along the irrigation canal and creeks from the foothills, and was most common in Kearney Park in the city of Fresno. He found nests 3 to 8 feet above the ground and lined with horsehair. He states:
“The usual complement of eggs is four, but I have found several sets of but three, and in at least three different instances the birds began the duties of incubation with just two eggs to their credit. The sets of two were in each case the first ones laid, so far as I could determine. May and June are the nesting months, my earliest record being May 1 (1906) for considerably incubated eggs. A set well along in incubation was found June 30 of the same year, while all other dates have fallen between these two extremes.”
Range: The Sacramento brown towhee is resident in California ea.st of the humid coastal region, from Humboldt County (Hoopa Valley) to Napa County, east to the foothills of the Cascade Mountains and the Sierra Nevadas, and south along the eastern side of San Joaquin Valley to Kern County (Piute Mountains, Fort Tejon).
Egg date: California: 1 record, May 9.
ARGUS BROWN TOWHEE
PIPILO EUSCUS EREMOPHILUS van Rossem
Contributed by HENRY E. CHILDS, JR.
This very restricted race of the brown towhee is considered a relict population of a group formerly more widely distributed but now confined by the deserts surrounding the Argus Mountains. It was described in 1935 from specimens taken in Mountain Spring Canyon at 5,500 feet in the Argus Mountains of Inyo County, Calif. As John Davis (1951) points out: “Nearly all [known] specimens have been collected at this locality and only at this station has breeding been established. It seems likely that breeding colonies of this form will be found in other canyons in the Argus Mountains in which willow thickets are present, such as Wilson Canyon.”
Nothing is known about possible variations in its patterns of life history and behavior from those of other races. Davis (1951) surmises that a local migration takes place in winter, as he found no brown towhees at Mountain Spring Canyon in December when the ground above 4,300 feet was covered with snow. An apparent vagrant taken at Lone Pine, Inyo County, is the only record for the race away from the Argus Mountains.
Range: The Argus towhee is resident in the Argus Mountains of Inyo and San Bernardino counties in California.
SAN FRANCISCO BROWN TOWHEE
PIPILO FUSCUS PETULANS Grinnell and Swarth
Contributed by HENRY E. CHILDS, JR.
The brown towhee is a widespread species over much of the brushcovered areas of California and the southwest. It is, perhaps, as typically Californian as the chaparral it so commonly inhabits. Because of its drab appearance and uninteresting song, it has until recently received little more than casual study by bird enthusiasts. Yet the brown towhee is one of the most common dooryard species in suburban areas near the large cities of California, and an important element of the avifauna of much of the west.
The many subspecies of the brown towhee range from southern Oregon to southern Mexico and east into Texas. In California it occurs from the lower Sonoran to the Transition Zone (A. H. Miller, 1951c). Apparently the species originated in Mexico and spread northward during the Pliocene through a generalized sclerophyll woodland whose elements are called the Madro-Tertiary flora by D. I. Axelrod (1939). This floral association occurred north into California and contained many of the trees and shrubs found in that area today.
John Davis (1951) has shown that the brown towhees of the United States are divisible into two main groups of subspecies, each geographically isolated from the other. The crissalis group is characterized by a low degree of contrast between pileum and dorsum, a dark dorsal coloration, and a relatively long tail. These are the birds of California. The perpallidus group is distinguished by a high degree of contrast between pileum and dorsum, pale dorsal coloration, and a black pectoral spot. These are the birds of Arizona, New Mexico, Texas, and parts of Mexico.
Throughout its range the species is strictly resident and highly territorial. Movements within each population can only be interpreted as dispersal of young individuals.
The climate varies considerably within the range of this species, with maximum rainfall and lowest temperatures in southern Oregon and minimum rainfall and highest temperatures in the deserts of the southwest. In the coast ranges and in the foothills of the Sierra Nevada, which may be considered typical of the habitat of this species in California, the temperatures are moderate and rainfall is from 10 to 20 inches a year. These factors appear to limit the distribution of this species only when they result in little or no brush vegetation.
Within its range the brown towhee meets with the closely related Abert’s towhee (Pipio aberti) to which it is very similar in appearance. Ahert’s towhee is a bird of desert riparian and lacustrine vegetation, a much more restricted habitat than that of the brown towhee. In southern Mexico another close relative, Pipilo rutilu.s, is found. The brown towhee also comes in contact with the rufoussided towhee (Pipilo erythrophthalmu.s), a ground-nesting species inhabiting more dense vegetation than that of its congener. The rufous-sided and brown towhees differ considerably in habits, song, and coloration, indicating a much more distant relationship than that between the brown and Abert’s towhees. Davis suggests that jwscus may be related only distantly to erytkrophthalmus and may be more closely related to the genus Melozone.
The brown towhee is primarily associated with the chaparral and secondarily with riparian vegetation. Davis (1951) describes its vegetation requirements as follows:
The growth form most desirable for protection and for location of the nest may be provided by many plants or combination of plants. However, the association utilized to the greatest extent is Californian chaparral. Of primary importance in this association are A denostoma fa.sciculatum and Ceanothus cuneaius. Both species are widely distributed in the Coast ranges, the Cascade and Sierra Nevada foothills, and the mountains of Southern California. Both extend into northwestern Baja California, and Ceanothus cunea4us extends northward into Oregon. Other important chaparral shrubs are Prunus iticifolia, Pickeringia montana, Dendromecon rigida, Photinia arbutifolia, and various species of Arctostaphytos, Rhus, Quercus, Rhamnus, Gerr ye, and Eriodict yen. The most important trees associated with the chaparral are Arbutus menziesii, various species of Qucrcus, Pinus sabiniana, and P. coulteri (MeMinn, 1939:6).
The Californian races are found secondarily in associations other than chaparral. They have been recorded frequently in riparian vegetation, especially in Alnus and Saliz and in tangles of Rubus vitifolius, R. parviflora, and Ribes. Plants such as Baccharis, Diplacus, Vitis, and &zmbucus may also be utilized. A few specimens have been taken in Artemisia. Disturbed areas may be used when the proper growth form has succeeded the original vegetation or when the original vegetation has been modified to provide suitable habitat for the brown towhees. Grazing, farming, logging operations, road building, and landscaping have opened areas to invasion by towbees. Local spread or increase in numbers has resulted.
Two Californian races occur mainly or entirely in associations other than chaparral. P. f. petulans of the humid north-central coast occupies an area in which chaparral is much restricted. This race is found in chaparral where this plant formation occurs within its range, but the birds have been restricted for the most part to the secondary association just listed. P. f. eremophilis of the Argus Mountains of Inyo and San Bernardino counties is found in rocky canyons that support extensive thickets of Salix.
Within these plant associations the brown towhee is an inhabitant of the brush edges. It is found along roads, trails, clearings, on lawns, wherever it can forage in the open with a clear view of predators and close to safe cover. Edge also appears to be of importance in determining territorial size.
During the winter towhees are found in association with other species of ground-feeding birds, particularly the white and goldencrowned sparrows, song sparrow, and house finch. It is the despot wherever it is found, putting all smaller birds to ffight by strongly attacking those in its path. At Berkeley only the larger Steller and scrub jays dominate any chance encounter.
That the brown towhee is a conspicuous bird in its habitat is indicated by T. L. Rodgers and C. G. Sibley (1940), who recorded it in 99.6 percent of trips taken on the Berkeley campus throughout the year.
Territory: Although lacking in brilliance of plumage and beauty of song, the brown towhee is well able to establish and maintain a typical passerine territory providing all the necessities for its comparatively sedentary life. In it are found mating and nest sites, food gathering areas, and escape areas for flight and roosting. A strictly resident bird, the male towhee defends his territory vigorously from all encroachment by others of his species.
Territorial defense is first noted in late fall after a period following the major dispersal of juveniles. Chasing and fighting between individuals becomes more frequent and pronounced in spring with the onset of breeding. Normally a relatively quiet bird, the towhee then becomes a loud, dominating tyrant to the hapless neighbor or vagrant towhee caught trespassing on his territory.
Song plays only a small part in the announcement of territory. Birds of neighboring pairs can see one another over most of their territories and trespass is usually observed and reacted to almost immediately. During the morning twilight hours in early spring, the male moves around the limits of his territory giving a rapid loud tsip note. This behavior, constant enough to be useful in census taking, ceases with the onset of nesting activities.
Population density reaches its maximum in disturbed areas such as the campus of the University of California at Berkeley. Here the vegetation is composed of ornamental shrubs, usually in a linear arrangement along buildings, streams, or paths, with much of the area in lawns. This pattern of vegetation provides nesting cover in dense shrubs from 2 to 5 feet tall, foraging areas on the lawns for these primarily ground feeding, seed-eating birds, and escape cover close to the foraging and nesting areas, mandatory because the towhee’s short, rounded wings do not allow long or fast ffight. The habitat thus provided appears optimum for the brown towhee as indicated by its abundance, 48 pairs per 100 acres. The territories, 1: 2 acres in size, are remarkably uniform. In a study of avian responses to artificial habitat near Dillon Beach, Calif., F. A. Pitelka (1942) observed one once in this coastal locality, but found no towhees breeding.
In natural areas such as the Berkeley hills, where the shrub vegetation of the west-facing slopes grows along drainages, territories are larger and longer, following the intersection of shrubs and grass. Here individual territories may be over 5 acres in size.
Courtship: Courtship behavior has not been described in great detail. On several occasions during the breeding season the male has been observed approaching the female with wings slightly drooped and quivering. Both wings may quiver in unison or alternatingly. Copulation may occur on the ground, but on one occasion it was observed on the top of a three-story building. The female assumes a position with the head tipped slightly forward and down and raises her tail. The male then mounts. In all cases observed both sexes were silent during coition.
Nesting: Most brown towhees at Berkeley start nest building by mid-April, and some pairs complete three nestings successfully by the end of summer. Incubation by the female alone takes 11 days and the first clutch of three or four usually hatches in late April. After 8 days in the nest, the young fledge enough to leave it. The period of dependence on the adults varies. If there is no renesting, the young may remain with the adults from 4 to 6 weeks, but when the adults renest they drive the first young from the territory when the next clutch hatches.
The nests are rather bulky affairs of grasses and twigs of many kinds, and not too well constructed. Built from ground level to more than 35 feet up in a variety of trees and shrubs, the optimum height appears to be between 4 and 12 feet. The nest is placed in the densest part of the foliage and is usually supported by several branches. J. Grinnell and M. W. Wythe (1927) report nests being found from midApril to the first week in September, situated well above ground in thick bushes or low trees.
A series of unusual nest sites have been reported. J. Mailliard (1936) reports a nest built a few inches underneath an apple. When the apple finally fell, it did not break any of the four-egg clutch, but the nest was deserted. J. McB. Robertson (1931) reports that towhees will nest in eucalyptus in accumulations of bark not too far from the ground. It is one of the few species to respond to this exotic tree, so plentiful in California. Brown towhees C. D. Scott (1920) kept in captivity nested on the ground two successive years. W. M. Pierce (1915) found a pair of brown towhees nesting in a berry bpsket 10 feet inside a barn door, but they deserted it midway in incubation.
Eggs: The measurements of 40 eggs average 24.9 by 18.3 millimeters; the eggs showing the four extremes measure 26.4 by 18.3, 25.9 by 19.7, 20.9 by 18.0, and 25.7 by 17.5 millimeters.
Plumages: John Davis (1951) bas the following to say about the molts and plumages of the brown towhee:
The only molt of real significance in the brown towhee is the fall molt. At this time adults undergo complete feather replacement. Birds undergoing the postjuvenal (first fall) molt retain the primaries and secondaries, and usually the rectrices. This molt pattern offers plumage characters that may he used to separate first-year birds from second-year or older birds. In fresh fall plumage the primaries of first-year (=~ immature) birds are dull brown and the margins of these feathers are finely erose because of wear. The primaries of second-year and older birds (~= adults) at this time are glossy and nearly black and the margins of these feathers are entire. The rectrices of immature birds have pointed tips * * * and show signs of wear. The rectrices of adults are obtuse at the tip * * * and are glossy and unworn. Since rectrices are often replaced in the post-juvenal molt, the amount of wear on the primaries offers the most reliable criterion for judging age.
The period of time over which these criteria may be used depends entirely on the amount of wear to which the plumage of the individual is subjected. As the adult primaries become worn they take on the dull brown color and erose margins of the immature primaries. As the tips of the adult rectrices become worn they become so ragged that it is impossible to distinguish them from the rectrices of unmatures. A few individuals may lose the characters of their age group as early as December; others may retain them as late as June. Age determination of most individuals becomes impossible by the middle of February. Occasional immatures are encountered which retain a few of the juvenal upper tail coverts. This has enabled the determination of age of some specimens that were otherwise so worn that the usual criteria were of no use.
The plumage of brown towhees is especially subject to wear. The abrasion of feathers causes a noticeable paling over the entire plumage. This paling is most pronounced on the coloration of the throat, since the color of this region is restricted to the terminal two-thirds of the throat feathers. As the colored portions of these feathers are worn away, the white feather bases arc exposed. In examples of extreme wear the throat may appear white, because only the feather bases remain. The coloration of the crissum is also strikingly affected by wear, for the intensity of the coloration of this area is deterni med largely by the thickness of mass of the under tail coverts. As these become worn the color of the crissum becomes paler, owing to the thinning of the feather mass.
Food: Brown towhees forage on the ground for seeds and insects. Occasionally they pursue large flying insects, but in the main they depend on the small ground surface forms. F. E. L. Beal (1910) made a study of stomach contents of these birds and found weed seeds: 51 percent: to be their most important food, followed by grain: 28 percent. Insects were the principal animal food: 14 percent: and fruit: 4.4 percent: probably windfalls, taken occasionally.
To obtain its food the brown towhee does not scratch as frequently or as violently as does the rufous-sided towhee. Birds are seen occasionally feeding on new grass shoots and drinking the dew collected on them in the morning. They respond well to feeding stations baited with various commercial bird seeds and bread crumbs, oat groats being particularly favored. The young, as in most other fringillids, are fed entirely on insect material, largely moth and butterfly larvae.
Behavior: A behavior pattern often mentioned in the literature is the brown towhee’s so-called “shadow-boxing,” a fighting response aroused when its reflection in a shiny surface suggests the presence of another towhee in its territory. D. P. Dickey (1916) first described this action for senicula as follows: “Perching on the sill, the bird would eye his reflection, and then set systematically to work to kill the supposed rival, with all the ire and intolerance of a rutting moose.” Reflections from window panes near feeding stations frequently stimulate these attacks, and hub caps often receive the same attention in the Berkeley area from both towhees and robins. W. E. Ritter and S. Benson (1934) describe and discuss the meaning of this phenomenon in terms of breeding activity and territorial behavior:
The Towhee, standing on the ledge, would face the window and assume a threatening attitude by lowering its bead, fluffing out its feathers, and drooping its wings. It would then leap up at the window, striking it with its feet, or with the feet and the beak at the height of about ten inches. It would then fall back and immediately leap up to strike again. Sometimes it varied the procedure by continuing up the pane, clawing at its image as it rose.
After May 1 the bird fought the window every day until July 4. Its last visit for the summer was July 14, but it appeared again on September 23 and fought occasionally for some weeks. The amount of fighting was never constant during this period. Beginning April 28 the bird increased its fighting activity until May 15. During this period it conllned its efforts to the windows of our rooms. By May 20 the attacks had fallen off a great deal and the bird had expanded the zone of its operations to include the west-facing window, of the main room of the Museum. By May 25 the attacks had nearly ceased, but after this date they increased again at our windows until on June 24 they were about as vigorous as ever. Subsequently they dwindled again to cease finally after July 14. On September 23, and for a few days subsequent to it more attacks occurred.
An explanation of the variation in the amount of activity was easy to discover. During the first period of increasing activity the fighting bird always accompanied its mate to the window. Their appearance at the window ledge was always heralded by a medley of mewing and squeaking notes from the oak tree outside the window. While the female fed busily the male would fight the window. The birds would usually fly into the oak tree where the female would sometimes preen awhile, but in a short time the female would fly to the west and disappear among the trees bordering Strawberry Creek. The male always closely accompanied the female. In a short time the male would return to the window to fight. The actions of the pair, and the presence of brood patches on the female, led to the belief that the female was incubating, but it appeared that the male was taking little if any part in the incubation other than guarding the female. * * *
It is clearly apparent from the above description that shadow boxing is intimately connected with breeding and defense of territory. The authors further point out that the attacks dropped ofT during the feeding of the young and increased again with the second brood.
E. I. Dyer (1931) observed four or five brown towhees respond to the distress calls of rufous-sided towhees when he approached too closely to their nest.
Voice: Just as uninspiring as its plumage is the voice of the brown towhee. With care, however, it is possible to distinguish many variations of the basic chip note and to interpret the meanings of these variations. Indeed, to understand the mechanics of the species’ population biology, one must become acquainted with its metallic clatter song. C. W. Quaintance (1938, 1941) has reported at length on the voice of the brown towhee and in the main this discussion comes from his work.
The basic note of the brown towhee’s vocabulary is a metallic chip or t.sip, often repeated with monotonous regularity, sometimes as fast as 30 per minute for 25 minutes at a time. Quaintance (1941) thus describes the use of this note:
The tsip note is not necessarily correlated with activity, although it does announce the beginning of activity in the early mornings and again, the cessation of activity in the evenings before the towhees go to roost. On the other hand a towhee may tsip for fifteen minutes without any activity. In flight it may give several tsips or none at all. An emphatic tsip may announce the take-off, and a tsip or two may be given upon alighting from a flight. A towhee may tsip on the ground while it is foraging, or it may forage for minutes without giving a sound. Frequently, in the nesting season, the tsiping is done from a high station such as the top of a laurel tree, or thirty or forty feet up in a eucalyptus, or from the peak of the highest available house or telephone pole.
The general tsip note seems to have different functions. It may serve as a contact note between birds, especially birds of a pair, as a protest note, or as an alarm or warning note. Further study of this note may yet resolve it into at least three different notes corresponding to the behavior induced. In other words, although the tsip may sound the same to us, the variations in manner of delivery or loudness may have meaning.
The tsips of a female disturbed at the nest are immediately answered by her mate and if her notes become hurried and excitable, he may come racing in to her side, no matter what part of the territory he may be in. Both birds may then utter loud isips of protest at an intruder. The protest note invariably brings neighboring towhees close to the nest in disregard of territorial boundaries. Birds of other species are also attracted to the region of the nest in apparent curiosity.
The rapidly repeated trip early in the morning is apparently the means by which the male towhee announces possession of his territory and its boundaries. He starts this tripping while moving about over his territory shortly after dawn starts to break, calling it from the tops of bushes, boulders, and other song perches within the territory at the rate of about 60 notes per minute. He continues this activity until sunrise and then stops. I have observed this behavior only during the early part of the breeding cycle from mid-March to June at Berkeley, and mid-April to June at O’Ncals, Calif.
During the nesting season the trip note may be accentuated to a trinlc note which appears to function in the control and warning of the young. It also serves to warn the female of danger while she is on the nest. A faint sparrowlike tsrp is used to keep in contact with a mate when separated by shrubbery. Conditions of great stress, such as attack by predators or handling by humans, produce a loud squawking. When uttered by a young bird, this squakwing acts as a distress signal that brings the parents rushing to the scene.
Qne of the most interesting combination of notes is the brown towhee’s mate-call, which C. W. Quaintance (1941) describes as follows:
“To me, the basic trip note is distinguishable in this particular series of notes. The utterance sounds something like this: Tsr’ tsr’ tsr’ tsurr trurr trurr, starting with fast staccato notes and getting faster toward the end. Because these unique notes are given almost exclusively between birds of a mated pair, they are referred to in this study as mate-notes or as the mate-call.”
A casual observer seeing two birds fly into shrubbery uttering this call might associate the notes with fighting. The observation of banded birds allows only the interpretation of the mate-call as having a “pair reinforcement” function. Quaintance (1941) continues:
In most of these instances the meaning of the mate-call might have been misconstrued if the whole action was not carefully followed. In over a hundred recorded observations these mate-notes were given between members of a known pair when they met and in no instance did the two fight. Moreover, in approximately one hundred clashes or boundary disputes between birds of different pairs, no mate-call was given * *
Outside of the nesting season the mate-calls are given less often. Before meeting in the morning, the members of a pair may forage separately for awhile; when they meet, the mate-call is given * * *ï Generally speaking, the mated birds forage together in the winter months and since the meetings are relatively fewer, the mate-call is given less.
It is difficult to tell which sex gives the mate-call. Sometimes it appears that only one of the birds gives the call, but more often it seems to be a duet.
Of particular interest is the brown towhee’s only really musical attempt at song, which appears to function chiefly in attracting females to unattached males. As all birds singing it that have been collected have been males, it is apparently a male trait. Its prevalence from late January into June suggests that the sex ratio is often unbalanced in favor of the male.
This song is an elaboration of the basic chip or tsip note repeated three or four times in succession and followed by a rapid, sometimes descending series of notes almost trill-like in quality. R. Hunt (1922) confines the rhythm of this song to that 6f a golf ball dropped on a hard surface and allowed to bounce until it stops. The trill-like ending may vary considerably from a linnetlike trill to the bubbling of a winter wren, and at times may be omitted entirely. This finchlike warble appears to be a retention of an ancestral song which is gradually being lost. As the brown towhee lives where members of a pair can almost always see each other and their neighbors, such elaborate advertising as in the song sparrow or mockingbird is not necessary.
The note of the young in the nest calling for food is a pu/dee, reminescent of the call of the rufous-sided towhee. The fledgling’s hunger note is similar to that of most sparrows, a loud tst, ta, tst. As the young bird becomes more active it gives a faint tssp.
In territorial clashes between neighbors or wandering intruders and when driving juveniles from their natal territory, a throaty techuek note is repeated rapidly during flight. There can be no mistaking the peculiar quality of this note or the intent of the bird making it. It means: “Get out and stay out.”
In rare instances the males utter a whispered finchlike warble not unlike the song of the house finch, but so faint it cannot be heard more than a few feet away. A. H. Miller (MS.) heard it from a male fighting his reflection in a window, and another time from a paired male bidden in the foliage of a tree. Several times I have heard a male give this song near the nest when the incubating female had left it to feed. From the variety of situations in which it is used, and the fact that it is inaudible at a distance of over 10 feet, the function of this song is not clear. Presumably it is another ancestral remnant of song no longer highly important to the species. A. H. Miller (MS.) reports a similar pleasant fineblike song in brown towhees in northern Mexico.
Its varied repertoire is indeed remarkable for so unmelodious a bird, which seems to act on the premise “one tsip is worth a thousand words.”
Field marks: This dull brown bird appears as a very plain, large sparrow, slightly smaller than a robin. The moderately long tail and the cinnamon or rusty under tall coverts are the most noticeable field marks. The throat and upper breast are buff, sometimes with faint dusky streaks. Its hunched over posture and ground feeding habits readily identity it from other sparrows.
Enemies: On the Berkeley campus I found the Norway rat the principal predator of the brown towhee, destroying many nests, eggs, and young along the wooded creeks. The domestic house cat was often seen stalking these birds. Man, through destruction of habitat and disturbance of nests, was the greatest single’ factor in nest desertion.
In November 1948 at Berkeley I watched a pair of towhees chivvy a cat crouched beside an ash can. First one bird, then the other, dropped to the ground about 10 feet from the cat. Both tsipped loudly and continuously, one at the rate of 60 per minute. Whitecrown sparrows and a scrub jay were attracted by the noise. The cat meowed, apparently in frustrat.ion, as the birds moved around it just out of reach. After 5 minutes the birds lost interest and departed.
The same year I reported (1948) the reactions of a pair of brown towhees to a pair of scrubjays intent on robbing their nest. Another pair of towhees joined them, and all four flew at the jays, fluttering widespread wings and tail and uttering the squawk note. Several times they dropped to the ground within a few feet of me, showing no fear. In their excitement the resident pair attacked the other towhees trying to help them, territorial defense against their own kind becoming momentarily paramount. One jay was driven off, but the second, after being driven from the nest bush three times, was no longer molested. The following day the nest was empty, the towhees had left the area, and did not return.
In a series of food habit studies of important predators on range lands at the San Joaquin Experimental Range at O’Neals, Calif., H. S. Fitch (1948a, 1948b, 1949) found the towhee to be a minor prey subject. Ground squirrels, although serious predators of valley quail eggs, were seen being chased by adult towhees away from a nest containing four young. Coyotes were abundant, but among 2,250 identified items in their diet he found only two towhees. Of the snakes in the area the Pacific rattlesnake was the only species known to take towhees, and only two were found in 285 identified prey items. Fitch et al. (1946a), in their study of the red-tailed hawk report only 3 towhees in 625 articles of food brought to the nest, and 7 towhees in 2,094 pellets. Fitch et al. (1946b) found only 1 towhee in 41 items at two nests of the Cooper hawk. Apparently the predation rate on towhees in this area is low and has little effect on the population.
J. M. Linsdale (1931), investigating the destructive effects on wildlife of a rodenticide then in common use, found only one towhee killed by this material. Compound 1080, now widely used for rodent control, particularly ground squirrels, is mixed with oats and colored a bright canary yellow that apparently make the seeds unattractive to birds.
Towhees evince a mobbing reaction to owls during the mating season, but not a very intense one, according to S. A. Altmann (1956), and Fitch found no towhees among the owl foods he identified.
Young brown towbees in the nest are occasionally parasitized by fly (Protocalliphora) larvae. 0. E. Plath (1919) examined eight nests and found two of them heavily infested with 53 larvae per nest. None were observed in the Berkeley population. C. M. Herman et al. (1942) found five out of eight towbees infected with Isopora. Hippoboscid or flat flies are found on towbees during the warm months.
Because of their liking for roadside vegetation and their poor powers of flight, brown towhees are frequent victims of automobiles. J. McB. Robertson (1930) listed four such casualties in a study in southern California.
Range: The San Francisco brown towhee is resident in California in the humid north central coastal region from Humboldt County (Korbel) to Santa Cruz County (Corralitos), and inland to the western edge of the northern San Joaquin Valley.
Egg dates: California: 8 records, May 9 to June 28.
CALIFORNIA BROWN TOWHEE
PIPILO FUSCUS CRISSALIS (Vigors)
Contributed by HENRY E. CHILDS, JR.
Most of our information about this south central coastal race comes from the Hastings Natural History Reservation in the northern Santa Lucia Mountains in Monterey County near Robles del Rio, Calif., in publications by Jean M. Linsdale. Since the late 1030’s many students have recorded information on the animal life of this 1600-acre study area.
An intense banding program was carried out during the winter on a section of the canyon near the living quarters (Lindsale, 1949). Brown towhees are one of five most plentiful species here, the others being the scrub jay, the rufous-sided towhee, the plain titmouse, and the Oregon junco. Brown towhees were banded in some numbers, 452 from November 1937 to June 1948 or about 41 per year. Later recaptures of 361 of these provide some information on survival and longevity in the species: 184 (51 percent) survived to at least 1 year of age, 83 (23 percent) lived 2 years, 51 (14 percent) 3 years, 29 (8 percent) 4 years, 9 (2 percent) 5 years, 3 (0.8 percent) 6 years, and 2 (.06 percent) 7 years, the oldest recorded in the study. Thus seven years may be assumed as the potential ecological life span for this species. From these data the average individual life span is 1.9 years.
Eggs: The measurements of 50 eggs average 24.9 by 18.4 millimeters; the eggs showing the four extremes measure 26.2 by 19.8, 25.7 by 20.3, and 22.0 by 17.0 millimeters.
Range: The California brown towhee is resident in California from the central coast area (Seaside) to the western edge of the San Joaquin Valley (Orestimba Peak) and south to western Kern (Temblor Range), Santa Barbara, and Ventura Counties.
Egg dates: California: 13 records, April 27 to June 20.
ANTHONY’S BROWN TOWHEE
PIPILO FUSCUS SENICULA Anthony
Contributed by HENRY E. CHILDS, JR.
This is by far the darkest colored of the California races of the brown towhee. John Davis (1951) suggests this is due to its habitat. He states that senicula is resident in extreme southwestern California in “The area between the seacoast and the western edge of the deserts in Los Angeles County, San Bernadino County * * a”, Riverside, Orange, and San Diego counties, thence southeast in Baja California between the coast and the lower edge of the coniferous forest on the Sierra Juarez and the Sierra San Pedro Martir south to latitude 29o.~~ The chaparral the species inhabits in this region contains less Ceanothus than the chaparral farther north and more Adenostoma, which gives the vegetation, commonly (ailed “black chaparral,” its (larker aspect. This dark background, augmented by the frequent reduction of illumination by fog (and smog), tends to select the darker variants in the population.
Though essentially an inhabitant of the wild sage and chaparral country, this subspecies has made itself at home in the settled suburbs of Los Angeles, Pasadena, San Diego, and other cities that have sprung up within its range during the past century. Wherever open lawns are adjacent to dense base-plantings and other ornamental shrubbery, the brown towhee becomes as common a dooryard resident as the mockingbird.
At their home in Pasadena, Harold and Josephine Michener (MS.) banded more than 1,300 brown towbees over a 25-year period. From most of these nothing more was ever heard, but a number of individuals came back to their traps year after year, the oldest one 8 years after they had banded it as a juvenal. In common with others who have tried to study this species through banding, the Micheners found that most individuals soon became “trap-wise.” Though birds holding territory in the vicinity may visit the feeding trays regularly and glean bait spilled outside the traps, after once being caught they are reluctant to enter the trap again immediately. By marking some of the resident individuals with colored bands, the Micheners were able to keep some track of their movements by observation alone.
One of their most interesting case histories involves a male they banded as a juvenal in August 1933, who established his territory on their property and maintained it there until he was last recorded in March 1938, when he was almost 5 years old. “He lived in the yard as we did, and was so easily found that to record each time he was seen would have seemed absurd. We soon learned that his territory, defined as the space within which a great many sight records were made and beyond which few or none are noted, extended approximately 150 by 175 feet. This area contained a great many shrubs and trees for its size, several water drips, and food was always available at a shelf on the corner of the house, on a window ledge, and on the drive.”
During the four breeding seasons this bird lived with the Micheners, though they were unable to keep track of all his nesting activities, they were able to determine he had at least three different mates. Each nest these built within his territory was in a different place. The first nest found, with four eggs in early April 1935, was on top of an old mockingbird nest in a melaleuca bush. This nest was unsuccessful, probably owing to the effects of an unprecedented heavy rain, and its female was last recorded near it on May 3. By June 1 a new female that had been banded as a juvenal 6 years earlier in June 1929, appeared on the territory and, though no evidence of nesting was noted, remained in occupancy with the male throughout the winter. Her first nest was found Apr. 14, 1936, 7 feet up in a buddleja on the opposite side of the yard. This nest fledged its young and on May 2 the female was found sitting on a new nest 4 feet up m a small crataegus, from which she also fledged her brood. Late that autumn her dried remains were found among the broken eggshells in a thrid nest she had built 20 feet up in a eucalyptus. She was thus 7 years old when she died. No records of the male’s nesting activities or mate were obtained in 1937, though he remained on the territory throughout the year. The last time he was taken, in midMarch 1938, he had a new mate, a female banded as a juvenal in June 1935. She built her nest in a small acacia, from which she fledged a single young on Apr. 11, 1938.
The reactions of this male and his various mates to other brown towhees intruding on their territory varied at different times of the year. In the fall and winter they showed no resentment when pairs from adjoining territories joined them at the feeders, but ate with them peacefully. On Mar. 2, 1935, “he and his mate were eating on the ground at the corner of the house in company with the pair holding the adjacent territory to the west. The four birds were in perfect harmony. The following November, December, and January, the color-banded pair holding the territory to the east were frequent visitors as well. There was no fighting. From these and later observations we concluded that this peaceful behavior was due to the fact that the pairs were settled for the season, that they knew each other and, certainly at this time of year, our resident pair did not regard neighboring mated territory holders as a threat.”
Most territorial clashes were noted during the nesting season, and they usually involved comparatively strange birds. Whenever one of these managed to enter a trap on his territory, the male always attacked it viciously. “On May 1,1936, a towhee entered the trap near his nest. At once there was the sound of fighting and commotion. The trap was of 1-inch poultry wire, and the trapped bird kept putting his head through the mesh, where our male could and did inflict severe injuries. By the time I reached the trap the intruder was bleeding badly and in such a serious state that after reading his band number I released him at once. He did not fly, but ran across the yard, our male after him, looking curiously like two rats rather than birds. When he disappeared under an old barn our male returned and seemed to regard the incident as closed. We had banded the intruder as a juvenile July 24, 1935, and this was his only repeat record. The continued and violent attacks on trapped birds within a territory suggest that, as with mockingbirds, an intruder is always at a disadvantage and usually leaves at any sign of hostility. But a trapped bird cannot leave, and this fact is ignored by the territory owner.~~ In an anlysis of nest-site data in California, John Davis (1951) notes: “In the vicinity of towns and farms the birds seem to nest frequently in trees, especially fruit trees, and to a lesser extent, in ornamentals such as pines, palms, and poplars. Tree nests may be located up to twenty-five feet from the ground.” Of 20 nests taken under natural conditions in Reche Canyon, near Colton, Riverside County, “eleven were found in Eriogonum Ja8ciculatum. They were placed from two and one-half to three and one-half feet above the ground. Two nests were found in Ceanothus cuneatu.s, two and onehalf and three feet above the ground. The rest were found in various situations, including twenty feet up in a sycamore, and one each in Convolvuh~.s, Rhu.s dii’ersiloba, sunflower, ‘wild holly,’ nettle s, Salvia apiana, scrub oak, and Lupinus. One was in a cleft five feet above the ground on the face of a rock wall. Only one was in tuna cactus and only one on the ground.”
Eggs: The measurements of 13 eggs average 24.8 by 17.8 millimeters; the eggs showing the four extremes measure 27.7 by 18.2, 26.4 by 18.6, 21.1 by 17.5, and 23.8 by 17.5 millimeters.
Records of egg weights are not plentiful. Hanna (1 924a) recorded the weights of a clutch of six eggs found at Colton, Calif., in a nest in black sage, 2~4 feet above the ground. These eggs weighed 4.18, 4.14, 4.05, 4.05, 3.98, and 3.90 grams. As six is an unusual number for a brown towhee clutch, it would be interesting to compare these with the weights of eggs in the normal three- or four-egg clutches. DIsmIBuTION
Range: Anthony’s brown towhee is resident in southern California, west of the Mohave and Colorado deserts, from Los Angeles County southward through northwestern Baja California, west of the montane coniferous forests, to lat. 290201 N. (Yubay). Recorded once from Todos Santos Island, lat. 310481 N.
Egg dates: California: 3 records, March 25 to June 6.
SAN PABLO BROWN TOWHEE
PIPILO FUSCUS ARIPOLIUS Oberho1ser
Contributed by JOHN DAVIS
Described by H. C. Oberholser (1919), this race differs from senicula to the north in the paler and grayer coloration of its pileum and dorsum; whiter, less huffy mid-ventral area; pallor of the posterior part of the throat patch; and longer bill. In most characters, it is intermediate between 8enwula and albigula, which occurs to the south in the cape region of Baja California. The intermediacy of aripolius between senicula, a typical representative of the subspecies of the brown towhee found on the Pacific Coast, and albigida, which resembles the geographically isolated subspecies of the southwestern United States and northern Mexico, led Oberholser to the conclusion that all the brown towbees were conspecific and should be regarded as belonging to the single species Juscu.s, rather than divided into the species crissalis, for the birds of the Pacific Coast, and the speciesjuscus for the towhees to the east and south. The validity of his conclusion has been upheld by other workers.
The northern limits of the San Pablo brown towhee’s distribution occur near the sharp vegetational break between the mixed foothill and desert flora of northwestern Baja California and the extremely rich and varied desert flora that has its northern limits between latitudes 290 and 300 N. The life history of this form is virtually unknown, although it is a common bird over much of its range.
Range: The San Pablo brown towhee is resident in the middle section of the peninsula of Baja California, from Playa Maria Bay, lat. 28o55~ N. (Grinnell, 1928b) south to Guajademi, lat. 26035k N.
Egg date: Baja California: 1 record, April 26.
SAN LUCAS BROWN TOWHEE
PIPILO FUSCUS ALBIGULA Baird
Contributed by JOHN DAVIS
Described originally by Spencer F. Baird (1859) as Pipilo albigula, this well-marked subspecies differs from the race aripolius to the north in its paler, more rufescent pileum; paler dorsum, sides, flanks, throat, and under tail coverts; and in its greater area of ventral white. Superficially it resembles the canyon brown towhee (Pipio Jwscws mesoleucus), from which it is separated by a broad geographic hiatus, but it is easily differentiated from that form by its generally smaller size and lack of a black breast spot. Further, its voice is similar to that of the Pacific coast races rather than to that of mesoleucus (Marshall, 1964).
The San Lucas brown towhee is restricted to the cape regioxi, or terminal third of the peninsula of Baja California, from latitude 26~ 35′ N. south to Cape San Lucas. It is found in the rich desert vegetation of the lowlands, and it also occurs in the mountains of the cape region, in a vegetation containing such highland elements as Pinus cembroides, Glaueothea brandegeei, Populus monijeola, Nolina beldingi, Arbutus penirisularis, and Qu~ercus desia. Most museum specimens of this form have been collected in the lowlands, and this suggests that brown towhees are more common in the deserts of the cape region than in the mountains. Judging by the large number of specimens in collections, it is a common bird over most of its range.
Nests: W. Brewster (1902) stated that three nests collected by M. Abbot Frazar in late July were composed of dry grass, weed stalks, and twigs. Two were lined with horsehair, and one with horsehair and fine grass. Two nests were located in bushes, and one in a tree. All were 6 to 8 feet above ground. Each nest contained three eggs. Baird, Brewer, and Ridgway (1874b) describe two nests found by John Xantus; one, containing four eggs, was located in a wild Humuhts thicket; the other was found in a thicket of wild roses in a garden fence.
Eggs: Brewster (1902) described the three sets of eggs collected by Frazar as having a ground color “greenish white with a tinge of blue,” each egg marked “chiefly about the larger ends, with irregular spots, dashes, and pen-lines of lavender and purplish black.” Of the eggs collected by Xantus, Baird, Brewer and Ridgway (1874b) state: “They bear a strong resemblance to those of the P. Juscus, but the markings are darker and more distinctly defined, standing out with a clear and striking effect, in marked contrast with the light background. The ground-color of the eggs is a light tint of robin-blue. The markings of dots, dashes, and lines are all about the larger end, and are of a deep dark shade of purplish-brown, so dark as, except in a strong light, to be undistinguishable from black.”
II. M. Hill and I. L. Wiggins (1948) found evidence of fall breeding in a male collected five miles northwest of Canipol, Nov. 18, 1946. The left and right testes of this individual measured 6.0 X 3.5 mm. and 5.0 X 4.0 mm.
The measurements of 32 eggs average 23.5 by 17.2 millimeters; the eggs showing the four extremes measure 85.2 by 18.1, 20.8 by 16.3, and 23.9 by 15.8 millimeters.
Range: The San Lucas brown towhee is resident in southern Baja California from lat. 26o35~ N. south to Cape San Lucas.