Part 2

Individual trees in which males sang most were marked over a three-week period. Then the distances between the most remote perches were paced. These distances aided in determining the size of the territories. The general configuration of the vegetation within each territory determined the location of one or more boundaries of the territory. Each territory was given a number, 1, 2, 3, etc., as it was discovered; consequently there is no numerical relationship between the designations of the territories established in 1959 and 1960. Nests within a territory were designated as 1-a, 1-b, 1-c, etc.

Although experimentation was not a primary source of data, it proved useful in certain instances. A stuffed Blue Jay elicited mobbing behavior from nesting pairs. A dummy Bell Vireo elicited both agonistic and epigamic behavior from nesting pairs, depending on the phase of the nesting cycle.

The temperature at the beginning of each day's work was taken by means of a Weston dial thermometer. A hand counter and a pocket watch having a second hand were used in determining such data as frequency of song and periods of attentiveness by the sexes. Histological cross-sections, prepared by A. Wayne Wiens, of the ventral epidermis of both sexes were used to study brood patches.

STUDY AREA

The intensive field work was on a 39-acre tract (fig. 1) extending approximately 7/10 of a mile west from U. S. highway 59, which in 1959-1960 constituted the western city limit of Lawrence, Douglas County, Kansas. The eastern boundary of the study area is approximately 1-1/2 miles southwest of the County Courthouse in Lawrence. The eastern ten acres is associated with the Laboratory of Aquatic Biology of the University of Kansas. The 15 acres adjacent to this on the southwest is owned by the University of Kansas Endowment Association, but is used by Mr. E. H. Chamney for the grazing of cattle. This portion is bounded on the west by a stone fence, beyond which lies a 14-acre field of natural prairie owned by Dr. C. D. Clark that is bordered on the extreme west by a narrow thicket of elm saplings.

The principal topographic feature of the area is an arm of Mount Oread, that rises some 80 feet above the surrounding countryside. About 200 feet from the crest of the southwestern slope of the hill a 40-foot-wide diversion terrace directs run-off toward the two-acre reservoir that is the source of water for eight experimental fish ponds of the laboratory.

The predominant shrub-vegetation consists of Osage orange (Maclura pomifera), honey locust (Gleditsia triacanthos), and American elm (Ulmus americana). These saplings, ranging in height from 3 to 25 feet, grow in dense thickets as well as singly and in clumps of twos and threes. Larger trees of these same species grow along the crest of the hill, along the eastern and southeastern boundaries of the area, and along the stone fence separating University land from that owned by Dr. Clark. A dense growth of coralberry (Symphoricarpos orbiculatus) forms the understory just below the crest of the hill. Isolated clumps of dogwood (Cornus drummondi) and hawthorn (Crataegus mollis) are scattered throughout the area. These species of shrubs grow densely along the stone fence. The isolated thicket on the Clark land is composed primarily of elm and boxelder (Acer negundo), but includes scattered clumps of dogwood, Osage orange, and honey locust. Poplars (Populus deltoides) are the only large trees in this area.

The open areas between the thickets are grown up in red top (Triodia flava), bluestem (Andropogon scoparius), Switchgrass (Panicum virgatum), Kentucky bluegrass (Poa pratensis), bush clover (Lespedeza capitata) and mullen (Verbascum thapsus). Shrubby vegetation occupies about 65 per cent of the total area, but in the Clark portion constitutes only about 35 per cent of the ground cover.

Considerations of Habitat

Nolan (1960:226), summarizing the available information on habitat preferences of the Bell Vireo, indicates that this species tolerates "a rather wide range of differences in cover." He pointed out that a significant factor in habitat selection by this species may be avoidance of the White-eyed Vireo (V. griseus) where the two species are sympatric.

In Douglas County where the Bell Vireo is the common species, the White-eyed Vireo reaches the western extent of its known breeding range in Kansas. At the Natural History Reservation of the University of Kansas, where both species breed, the Bell Vireo occurs in "brush thickets in open places" (Fitch, 1958:270) and the White-eyed Vireo occupies "brush thickets, scrubby woodland and woodland edge" (Fitch, op. cit., 268). Along the Missouri River in extreme northeastern Kansas, Linsdale (1928:588-589) found the White-eyed Vireo "at the edge of the timber on the bluff, and in small clearings in the timber," while "the Bell Vireo was characteristic of the growths of willow thickets on newly formed sand bars." Elsewhere in northeastern Kansas I have found the Bell Vireo in shrubbery of varying density and often in habitat indistinguishable from that occupied by White-eyed Vireos at the Natural History Reservation. In the periphery of the region of sympatry the rarer species is confronted with a much higher population density of the common species and consequently might well be limited primarily to habitat less suitable for the common species. This would seem to be the case in eastern Kansas, presuming that interspecific competition exists.

The Bell Vireo has followed the prairie peninsula into Indiana, aided by the development of land for agriculture. In nearby Kentucky where thousands of miles of forest edge are found, and where little brushy habitat of the type preferred by the Bell Vireo occurs, the White-eyed Vireo is abundant whereas the Bell Vireo is unknown as a breeding bird (R. M. Mengel, personal communication).

In more central portions of the area of sympatry, nevertheless, the two species do occur within the same habitat (Ridgway, 1889:191; Bent, 1950:254) and occasionally within the same thicket (Ridgway, in Pitelka and Koestner, 1942:105); their morphological and behavioral differences, although slight, probably minimize interspecific conflict. The Bell Vireo and the Black-capped Vireo (V. atricapillus) have been found nesting in the same tree in Oklahoma by Bunker (1910:72); the nest of the black-cap was situated centrally and that of the Bell Vireo peripherally in the tree. Bell Vireos invariably place their nests in the outer portions of trees and peripherally in thickets. This placement would further obviate interspecific conflict with the white-eye since its nests are placed centrally in the denser portions of a thicket.

A critical feature of the habitat preferred by the Bell Vireo is the presence of water. In far western Kansas this species is restricted to riparian growth along the more permanent waterways. This in itself is not adequate proof of the significance of water supply because thicket growth in that part of the state is found only along waterways. The 20 areas over the state that I have visited where Bell Vireos were present were closely associated with at least a semi-permanent source of water. Fifteen other areas indistinguishable from the 20 just mentioned, but lacking a permanent supply of water, also lacked Bell Vireos. Nevertheless areas in which Bell Vireos typically nest are decidedly less mesic than those frequented by White-eyed Vireos.

Once the Bell Vireo was probably more local in its distribution being restricted to thickets associated with permanent water. Clearing of woodland for agricultural and other use, and subsequent encroachment of second growth concomitant with the creation of man-made lakes and ponds, has greatly increased the available habitat for this bird. The preferred species of shrubs for nesting are reported (Bent, 1950:254) to be various wild plums (Prunus sp.). The widespread distribution and abundance of the exotic Osage orange has greatly augmented the supply of trees suitable for nesting.

SEASONAL MOVEMENT

Arrival in Spring

The subspecies of the Bell Vireo breeding in Kansas, V. b. bellii, winters regularly from Guerrero and the Isthmus of Tehuantepec south to Guatemala, El Salvador, and northern Nicaragua (A. O. U. Check-list, Fifth Edition, 1957:469-470). In the United States migrating birds are first recorded in early March (Cooke, 1909:119). The Bell Vireo is a relatively slow migrator, moving primarily at night and covering little more than 20 miles at a time (Cooke, op. cit. 119). The average date of arrival, based on 27 records, for northeastern Kansas is May 8; the earliest record is April 22, 1925, from Manhattan, Riley County, Kansas (fig. 2-A).

In 1959 the first bird arrived at the study tract about May 5. No additional birds were heard singing until the third week of the month, in which eight new males were noted. As mentioned, in 1960 field work was begun in mid-April and the study area was traversed daily. No birds were detected until late afternoon of May 3, when one, presumably a male, was seen foraging.

Lawrence (1953:50) has reported that males of the Red-eyed Vireo precede females in the breeding area by as much as two weeks; the male Red-eyed Vireo forages but sings little in the pre-nesting period. The male Bell Vireo arrives first at the breeding area but precedes the female by only a few days. On the morning of May 4 the first male was singing from a number of perches while ranging over an area of seven acres. This area encompassed territories later occupied by three pairs, 2 (1960), 4 (1960), and 5 (1960). Late on the afternoon of May 4 the first courtship songs were heard and the first male was seen with a mate at 6:20 p.m. Eight additional males arrived from May 6 through May 18. A tenth male was discovered in the vicinity of territory 9 (1960) on June 18, 1960.

Fall Departure

The average date of departure for 21 years in northeastern Kansas is September 3 (fig. 2-B). The earliest date is August 14 from Concordia, Cloud County, Kansas (Porter, unpublished field notes). The latest date is September 27 (Bent, 1950:262) from Onaga, Pottawatomie County, Kansas. In 1959 the last vireo was seen at the study tract on September 14. The birds do not all depart at the same time. On September 1 there were still five singing males in the study area; by September 10 there were three and on September 13, only one.

GENERAL BEHAVIOR

Flight

In "straight-away" flight the Bell Vireo undulates slightly. In a typical flight several rapid, but shallow, wing beats precede a fixed-wing glide of from 1 to 15 feet in length. Because the wings are extended horizontally during the glide, the bird does not move distinctly above or below the plane of flight. The White-eyed Vireo generally appears to be slower and more lethargic in flight than the Bell Vireo. In the breeding season most flights of the Bell Vireo are no longer than a few feet between adjacent shrubs and trees, but occasional sustained flights are as long as 300 feet. The birds fly as low as 2 feet above ground, but have often been observed as high as 70 feet above the ground.

In courtship and protracted territorial disputes, where chase between sexual partners or a pair of antagonists occurs, looping flights are observed. The wings are beaten as the birds climb and many aerial maneuvers are performed in the course of the glide.

Foraging and Food Habits

The Bell Vireo has been characterized as a thicket forager (Hamilton, 1958:311; Pitelka and Koestner, 1942:104), but in my experience it is not restricted to low level strata; birds forage from ground level upward, both in thickets and isolated trees ranging in height from 3 feet to 65 feet. The tendency to forage at higher levels is in part dictated by the presence of tall trees within the various territories.