Natural History of the Bell Vireo, Vireo bellii Audubon Part 4

Swaying has been recorded in a variety of situations of a s.e.xual and semi-s.e.xual nature for the Solitary Vireo (_V. solitarius_; Townsend, 1920:158) and the Red-eyed Vireo (Tyler, 1912:230; Bent, 1950:342). In every instance the body feathers of the swaying birds were sleeked.

Courtship behavior in any species of North American vireo seems closely to resemble that of any other; pairing and nestbuilding of a female _V. solitarius_ and a male _V. flavifrons_ as reported by Hauser (1959:383) support the idea of close resemblance.

A marked similarity will be detected between certain basic elements of aggressive and epigamic displays. These basic elements are wing- and tail-flicking, tail-fanning, and high-intensity delivery of the _chee_. Pouncing and supplanting attacks are essentially similar. Such similarities suggest either a common origin for certain aggressive and epigamic displays or the derivation of one from the other.

High-intensity _cheeing_ is obviously a function of excitement, whether in conjunction with hostility or s.e.xual behavior. According to Andrew (1956:179), flicking of wing and tail in pa.s.serines are intention movements of flight. These actions have been emanc.i.p.ated from incomplete take-offs and incorporated in ritualized courtship and agonistic behavior. In incipient courtship behavior the male is governed by three conflicting tendencies; to flee, to attack, or to behave s.e.xually before his mate (Tinbergen and Hinde, 1958:256). When pairing, Bell Vireos interrupt s.e.xual chase with "greeting ceremonies," the male's tendency to attack and the female's tendency to flee are momentarily reduced, and the forming bond is strengthened.

Thus, the intention movements become an integral part of courtship.

In situations where attacking and fleeing are the two conflicting tendencies, wing-flicking and tail-flicking are incorporated into threat display, but do not lose all of their original function, for they facilitate attack. Tail-fanning, as a display element, increases the awesome aspect of the threatening bird and in courtship presumably makes the s.e.xes more attractive to one another.

Courtship feeding has not been recorded for the Bell Vireo. In general, it is unknown in North American vireos, with the exception of the red-eye (Lawrence, 1953:53). It would serve no "practical" purpose in the Bell Vireo since the male regularly relieves the female during incubation, thus allowing her ample opportunity to forage. In the Red-eyed Vireo, only the female regularly incubates, and courtship feeding is definitely functional. Nolan (1960:228) described a brief pecking or pulling with their bills between pairing birds. This may be incipient "symbolic" courtship feeding, or perhaps mutual preening.

SELECTION OF NEST-SITE AND NESTBUILDING

As far as can be determined, the nest-site is selected by the female.

Typically, the pair makes short, low-level flights from tree to tree with the female invariably in the lead. The birds usually forage within each tree; the female interrupts this activity to inspect small forks of low, pendant branches and the male occasionally pauses to sing. The singing is loud but not particularly regular, as it is later when the male accompanies the female during actual nestbuilding.

Method of selection of site resembles that described by Lawrence (1953:53) for the Red-eyed Vireo.

Nests are suspended from lateral or terminal forks about 27 inches high in bushes and small trees that, in the study area, averaged 11 feet, four inches in height (Table 5). The height above ground of the nests does not vary appreciably as the season progresses as is the case with nests of Red-eyed Vireos, for which Lawrence (1953:54) noted that late nests were placed higher than those built earlier in the season.

Most nests are so situated that they are protected and concealed by the dense foliage of trees. Where nests are placed in low bushes, as coralberry or dogwood, the bush is invariably overhung by the foliage of a much taller shrub or tree.

The nest tree or shrub was in every instance situated at the edge of a thicket or isolated from adjacent trees by several feet. Preference for open situations is characteristic of the species. In contrast, the nest of the White-eyed Vireo (Bent, 1950:229) is placed toward the center of thickets.

In the choice of sites in the study area, the Bell Vireos were almost unopposed by other avian species, owing to the size of the fork utilized and the fact that the nests are located peripherally, rather than centrally, in the bush or tree. This lack of compet.i.tion for a nest-site provides a Bell Vireo with an ample supply of nest-sites within any one territory.

TABLE 5. NEST-SITES UTILIZED IN 1960.

==================================================================== | Number | Average | Average Plant | of | height of | height of | nests | plant | nest -----------------------------+--------+---------------+------------- _Ulmus americana_ | 4 | 7 ft. 6 in. | 2 ft. 3 in.

_Maclura pomifera_ | 20 | 13 ft. 11 in. | 1 ft. 11 in.

_Crataegus mollis_ | 1 | 11 ft. | 3 ft. 1 in.

_Gleditsia triacanthos_ | 2 | 15 ft. 6 in. | 1 ft. 9 in.

_Acer negundo_ | 4 | 8 ft. 9 in. | 2 ft. 5 in.

_Cornus drummondi_ | 2 | 8 ft. | 2 ft. 8 in.

_Symphoricarpos...o...b..culatus_ | 3 | 3 ft. | 1 ft. 10 in.

| | | 7 | 36 | 11 ft. 4 in. | 2 ft. 3 in.

Selection of the first nest-site may take as long as two days, possibly owing to incomplete development of the nesting tendency, but more likely to a general lack of familiarity with the territory.

Red-eyed Vireos require five to six days to choose the first nest-site (Lawrence, 1953:54). Later sites of the Bell Vireo are chosen in as little as three hours. Nest 1-c (1960) was abandoned at about 11:00 a.m. on May 14, 1960, when part of the thicket on the edge of which this nest was located was removed by brush-cutters clearing a power line right-of-way. By 2:00 p.m. this pair had begun construction of 1-d (1960) in an Osage orange 110 feet southwest of 1-c (1960).

This particular site is of further interest because it is the same one utilized for nest 1-a (1960). In all, four instances of utilization of a nest-site a second time were recorded. Two-a (1960) and 2-d (1960) were built in the same fork; 1-c (1960) and 1-h (1960) were in the same tree, but not the same fork. It should be mentioned that 1-a (1960) and 2-a (1960) were abortive attempts that did not progress beyond the suspension apparatus. Nice (1929:16) recorded a similar instance of the re-use of a nest tree, but different forks were used.

Re-use of an exact nest-site would ordinarily be impossible if the initial attempt were not abortive, because the presence of a completed nest would pose problems in construction with which the birds would probably be unable to cope. (A report by Morse in Bent, 1950:256 of a double nest indicates that this may not always be true. At the time of discovery one nest contained two eggs and the other nest contained young.) Since nests are used only once there would be no tendency to adopt the old nest. However, abortive nests, usually little more than a few strands of nesting material secured to the fork, might stimulate the birds to continue building. Re-use of a single nest-site in 15.8 per cent of 38 nests built in 1960 seems to be more than fortuitous circ.u.mstance. This re-use may have physiological benefits in conjunction with apportionment of energy for other nesting activities, because rapid location of a nest-site would mean that energy normally expended in searching and selecting could be rechanneled for actual construction. In each of the instances of rebuilding, the new nest was begun on the same day that the previous nest was abandoned.

The re-nesting of pair 9 (1960) is worthy of note. These birds were established in the elm thicket on Clark land. Elm was by far the most abundant tree, with dogwood, Osage orange and honey locust also relatively common. There were only six boxelders in the territory and yet the four nests built by this pair were placed in them. This is the only instance of seeming preference.

_Building_

Nestbuilding by Bell Vireos can be best discussed in terms of the phases of construction described for the Red-eyed Vireo, Lawrence (1953:57), which are: (1) construction of the suspension apparatus, (2) construction of the bag, (3) lining of the bag and smoothing and polishing of the exterior, and (4) adornment of the exterior. Red-eyes (Lawrence, 1953:59) may continue adornment far into the period of incubation. Both the male and female Bell Vireo have been observed to add spider egg sacs and other silk to the exterior of the nest as late as the sixth day of incubation.

Nice (1929:16) recorded only the female Bell Vireo building, but she did recall, from previous studies, having seen males aiding somewhat.

Pitelka and Koestner (1942:102) wrongly concluded that the female Bell Vireo builds unaided, but Hensley (1950:243) observed that both s.e.xes partic.i.p.ated in nestbuilding, and Mumford (1952:229) reported two instances of building by both adults. His description of the activities viewed in mid-May suggest that they were of the transitional period between the first and second phases. On the second occasion he recorded both adults building during the second phase.

Since no details accompany this second observation I a.s.sume that it pertained to activity not necessarily typical of this phase of construction. Whereas both s.e.xes of the Bell Vireo cooperate in building the nest, only the female Red-eyed Vireo builds according to Lawrence (1953:56). But Common (1934:242) saw both Red-eyed Vireos building a nest.

The suspension apparatus is constructed by only the male on the first day. He punctuates each trip to the nest with song. The single song phrase is given from three to eight times when the male, carrying nesting material in his bill, arrives in the tree. Typically, he alights on several perches within the nest tree before flying to the nest. He often interrupts his work with several songs; when he has finished adding a load of material he sings from several perches within the nest tree before departing. The male periodically stops building to court the female.

In eight hours (494 minutes) of observing the first phase of construction at five different nests, I saw the female come to the nest 28 times; the male made 95 trips. The female came alone only once, and brought nesting material ten times, but did not build; on the other 18 occasions her visits were brief and she usually confined her activities to an inspection of the nest. Twenty of the visits by the female were made late in the first phase, marking a gradual transition to her a.s.sumption of building responsibility. (The delay by the female in beginning to build is puzzling; because all evidence indicates that she helps select the nest-site, I would expect her to help with the initial building. There seems to be no clear advantage in her delay in beginning to build.) The courtship and building activities of the male plus the presence of a partly completed nest seem to stimulate the female to commence building. Her visits become more frequent as construction of the suspension apparatus nears completion. At a time early in the second day the transition has taken place, and the female becomes the sole worker.

On May 7, 1960, male 2 (1960), at the time unmated, was observed as he came upon a nest of the previous year. The nest, after a year's weathering, suggested in appearance perhaps an early second-day nest.

The bird flew to the nest and tugged and wove loose strands of gra.s.s for three minutes. Before leaving the site, the bird sang twice from different perches. This observation suggests that a partly constructed nest can elicit nestbuilding behavior, even in an unmated male.

The techniques of building by the male consist primarily of laying pieces of gra.s.s or bark across the fork, or along one of its branches, and then fastening them in place with pieces of animal silk. Once a "racket" has been formed, spider egg cases and plant down are emplaced among the fibers. The male employs weaving, twisting, and pecking motions of the head to emplace material.

As previously indicated, the female is the princ.i.p.al worker in the second and third phases of construction. The male infrequently visits the nest, but regularly visits the nest tree. The molding of the bag is accomplished by piling leaves, gra.s.ses and plant down onto the suspension apparatus. This material is also bound in with animal silk.

As the amount of material acc.u.mulates, the female begins to trample it and gradually the bag takes shape. When trampling is first attempted, the nest often fails to support the female and she falls through the bottom of the nest. Such an occurrence was observed on May 23, 1960, on three consecutive trips by female 1 (1960), in constructing nest 1-e (1960). As the bag deepens, additional strands of gra.s.s are added to the wall and woven into place.

The male is extremely attentive during this and the following phase.

He follows the female as she gathers nest-material accompanying both this activity and her building with rapid song; he may give an average of seven song phrases per minute. The male brings to the nest a strand of gra.s.s, or some other material, about every twentieth trip. He frequently inspects the nest and the activities of the female from perches near the nest. Construction of the bag is ordinarily completed in the third day.

The third phase, the lining of the interior and the smoothing of the exterior, involves an additional one and one-half to two days.

Smoothing of the exterior refers to tightening of the gra.s.ses woven into the bag and addition of more animal silk. In lining the nest, the female stands on one of the branches of the fork and emplaces one end of a long, thin strand of some relatively stiff piece of gra.s.s or strip of bark. She then jumps into the bag and, while slowly turning around, pecks it into place, thus coiling the strand neatly around the interior of the bag.

As previously mentioned, the fourth phase overlaps the periods of lining, smoothing, egglaying, and incubation. The princ.i.p.al activity is the addition of white spider egg sacs to the exterior. The trips are infrequent; but, occasionally, birds will interrupt an hour of incubation with three or four minutes of active adornment, during which several trips may be made. Both s.e.xes partic.i.p.ate in this phase.

_Gathering of Nesting Material_

Nesting materials were gathered anywhere within the territory.

Occasionally materials were collected from within the nest tree, but usually they were obtained 20 to 200 feet from the nest-site. On several occasions I observed birds inspecting stems or branches where bark was frayed. Loose ends are grasped in the beak and torn free with an upward jerk of the head. Possibly the notch near the distal end of the upper mandible aids in grasping these strands. Plant down is first extracted and then rolled into a ball by means of the beak while held with the feet before being transported to the nest.

_Length and Hours of Nestbuilding_

As indicated by Nolan (1960:230), accurate determination of the length of nestbuilding is difficult because of continued adornment and polishing after the nest is functionally complete. Most of the early nests for which I have records took from four and one-half to five days to construct. A four-to five-day period of building is reported by other observers (Nice, 1929:16; Pitelka and Koestner, 1942:99; Hensley, 1950:242; Nolan, 1960:230).

One instance of protracted building was recorded. Nest 6-d (1960) was begun on May 29, 1960, and not completed until nine days later on June 6, 1960. In contrast nest 1-g (1960) begun on May 31, 1960, was finished three days later on June 2, 1960. Nestbuilding occurs between the hours of 6:00 a.m. and 5:30 p.m. Heavy rain in the early morning may delay building.

_Abortive Nestbuilding Efforts_

Eight of 38 nests started in 1960 were never completed (Table 6). Six of these abortive attempts were abandoned during, or shortly after, the completion of the suspension apparatus. Five of these nests were abandoned because the female did not begin building following the end of work by the male. The early abandonment of the other three nests 1-a (1960), 2-c (1960) and 6-e (1960) was attributable to the interruption of building by the male because of heavy rain and protracted territorial conflicts. The occurrence of these abortive nests at any time within the nesting efforts of a single pair indicates that such attempts are not examples of "false nestbuilding."

_Renesting_