Part 3

Territories 1 through 7 (1960) contained from three to ten trees surpassing 40 feet in height. They grew singly or in small groves. The Bell Vireos foraged fully 20 per cent of the time in these trees. Food was sought throughout the leaf canopy.

Behavior in foraging in larger trees followed a routine pattern. Typically a pair alighted in a tree at a height of 15 feet. Then the female hopped to a perch a foot above the one upon which she landed. The male succeeded her to the perch she had previously occupied. The pair in effect spiraled around some large, essentially upright, branch, in foraging. The birds usually reached higher perches in this manner rather than by flying upward 10 to 15 feet to them. This manner of progression within a tree is reminiscent of a similar habit of the Cyanocitta jays. Presumably, the habit of the Bell Vireo of foraging in higher strata is facilitated by the absence of other species of arboreal foraging vireos.

Chapin (1925:25) found the Bell Vireo to be more insectivorous in its food habits than any other North American vireo. He found 99.3 per cent of all food contained in 52 stomachs to be of animal origin. Only three times have I seen a Bell Vireo take food of vegetable origin. On September 9, 10, and 14, 1959, I noted a male eating wild cherries over a period of 65 minutes of observation. Chapin (1925:27) noted that beginning in July vegetable matter represented 1.57 per cent of the bird's subsistence, and thereafter slightly more until fall migration.

Animal food, consisting primarily of insects and spiders, is actively sought along branches and under leaves. Often a foraging bird will leap to the underside of a branch and hover, mothlike, beneath a cluster of leaves while extracting some insect. Some individuals hung upside down on small branches, paridlike, while foraging. Lawrence (1953:710), and Southern (1958:201) have recorded similar behavior of the Red-eyed Vireo. Occasionally, I have seen a Bell Vireo fly from a perch and capture an insect in the manner of a flycatcher. The birds do not appear to be adept at this type of food-getting. Nolan (1960:242) mentions Bell Vireos holding hard-bodied insects by means of their feet while breaking the exoskeleton with the beak to obtain the soft parts. Southern (1958:201) recorded a female Red-eyed Vireo foraging on the ground; I have seen a Bell Vireo on the ground but once, and it was gathering nesting material.

Bathing

On May 14, 1960, in a rill that empties into the northeastern edge of the reservoir a female flew down from a perch six inches above the surface, barely dipped into the water, flew to a perch 12 inches above the water, violently shook her ruffled body feathers, quivered her wings, and rapidly flicked her fanned tail. The entire procedure was repeated three times in five minutes. She was accompanied by a singing male that did not bathe.

Nolan (1960:241) reports a male Bell Vireo bathing by rubbing against leaves wet with dew; he notes that the White-eyed Vireo bathes in a similar manner. Southern (1958:201) twice observed Red-eyed Vireos bathing in water that dropped from wet leaves. In my study area in 1960, only territories 7, 8, 9, and 10 were not immediately adjacent to permanent water. The pairs of Bell Vireos in those territories presumably had to reply on wet vegetation for bathing.

VOCALIZATIONS

The male Bell Vireo begins to sing regularly soon after its arrival in spring. Some daily singing continues following the cessation of breeding activities until departure of the species in late summer or early fall. The highest sustained rate of song occurs on the first and second days of nest building. Because careful records of meteorological data were not kept, I cannot significantly correlate rates of song and specific temperatures and other weather conditions. Frequency of song was reduced when the temperature rose above 90° F., as it did on many days in June, 1960. Nice (1929:17) mentions a similar decrease in singing when the temperature exceeded 85° F.

Passerine birds typically sing at a high rate throughout courtship and nestbuilding, but at a markedly lower rate thereafter. Most vireos are atypical in this respect. In the study area in 1960 Bell Vireos sang more often than Robins, Mockingbirds, Field Sparrows, Brown Thrashers, Catbirds, and Doves breeding in the same habitat, about as often as the Meadow Larks in the adjacent fields, and less often than Painted Buntings.

The Bell Vireo seems to sing less often in the undisturbed state than when aware of the presence of an observer. Observations from my car, at a site approximately equidistant from territories 1 (1960), 2 (1960), 4 (1960), and 6 (1960) indicate that the rate of song during incubation is decidedly less when no disturbing influence is present. Normally, in this period, song aids in maintaining contact between the members of a pair, serving to locate the male as he forages. Mumford (1952:230) noted that the males often came out to meet him as he entered their territories, singing as they approached. The male typically continues to sing for some time after the intruder has departed. Here the song acquires the additional functions of alerting the female to danger and threatening the trespasser. Even after allowance is made for this reaction to disturbance, Bell Vireos sing more often than most of their nesting associates, and, on a seasonal basis, they are vocal for a much longer time.

Singing Postures

In the normal singing posture the body of the Bell Vireo is maintained at an angle of 35° to the horizontal. Occasionally, during nest building, I have observed the body held at angles as severe as 80° from the horizontal.

The head of the White-eyed Vireo is distinctly bobbed up and down, two or three times, during the utterance of a song phrase. A bob involves a deliberate withdrawal of the head towards the body and subsequent sharp, almost vertical, extension of the neck. The head of the Bell Vireo does not bob, although it vibrates as the song is delivered.

Flight Song

The Bell Vireo does not have a distinctive flight song; in fact, it rarely sings or calls while in flight. Nolan (1960:240) has recorded a male singing the normal song while in flight. Sharp scold-notes are uttered in mid-air when a bird is agitated or actually attacking an enemy. These notes and songs recorded by Nolan hardly qualify as flight song, for this term implies use of a distinctive vocalization not uttered in other circumstances.

Daily Frequency of Song

In the morning, Bell Vireos usually began singing a few minutes before sunrise. Their songs were invariably preceded in the study area by those of Western Kingbirds, Robins, Mourning Doves, Mockingbirds, Cardinals and Meadow Larks. Bell Vireos sang relatively little after 6:30 p.m., even on the longest days of the year. The latest daytime singing that was recorded was seven songs at 7:18 p.m. on June 20, 1960. A Cardinal in the vicinity sang for a full hour after this.

Types of Vocalizations

Six vocalizations were readily distinguishable in the field. These are divisible into songs and call notes.

1. Primary song. It has been described by Pitelka and Koestner (1942:103) as an "irregular series of harsh and sharp, but slurred notes preceded by a few distinct notes of the same quality and ending with a decided ascending or descending note of similar harshness." The terminal note may also be somewhat abbreviated and intermediate between an ascending or descending note. The song is sometimes delivered as a couplet that consists of a phrase ending on a descending note. This delivery is typical of incubation and later renesting. During early season activities, the bird utters a phrase ending on the descending note as many as 15 times before a phrase ending on an ascending note is heard.

A sonagram of a single phrase, one of several recorded on May 9, 1960 (the third day of building of nest 1-b 1960), consists of 10 notes, the first of which is distinct. The remaining notes are slurred. This phrase is 1.4 seconds in length.

Songs are delivered most rapidly in the course of territorial disputes and defense. The song is loudest in times of nestbuilding and periods of aggressive behavior. At these times, on clear, calm days, the songs are audible 100 yards away. Singing in the nestling period and post-breeding season is audible at distances of no more than 50 feet; such notes have been termed "whisper songs." Table 1 summarizes singing rates at different periods of the nesting cycle in several situations and under various weather conditions.

Songs are of equal frequency in the immediate vicinity of the nest and elsewhere in the territory. Nice (1929:17) also found this to be true. Perches can be almost at ground level or as high as 60 feet. Forty per cent of my data on song concern singing at heights of more than 20 feet. As indicated in foraging, the lack of competition from aboreal species of vireos presumably contributes to the use of higher perches by Bell Vireos.

No female song was recorded in 1959, but on May 26, 1960, a female was heard to sing once. She appeared at nest 1-f (1960) shortly after the male arrived. Unlike him, she did not participate in building, but seemed to be inspecting the nest. After 30 seconds she sang once--a low garbled phrase--and also scolded once. After this she left. In the meantime the continuously singing male moved two feet away from the nest, then back to it and resumed construction.

The song of the female signaled to the male her departure. Pitelka and Koestner (1942:103) heard a female sing twice after she replaced the male on the nest. Females of three other species of vireos, the Black-capped Vireo, V. atricapillus (Lloyd, 1887:295), the Philadelphia Vireo, V. philadelphicus (Lewis, 1921:33), and the Latimer Vireo, V. latimeri (Spaulding in Pitelka and Koestner, 1942:103) have been heard singing. Lewis and Spaulding also suggest that the song of the female functions as a signal prior to exchange at the nest.

The primary song identifies the singer as a male Bell Vireo. It aids in securing a mate and in warning potential adversaries; also, the song is a signal in certain situations and serves to locate the male.

TABLE 1. REPRESENTATIVE SINGING RATES OF BREEDING BELL VIREOS. ALL RATES WERE AT AIR TEMPERATURES LESS THAN 86° F. EACH INSTANCE REPRESENTS APPROXIMATELY 30 MINUTES OF OBSERVATION.