From the first day the young are able to squeak. Poking a young bird was sufficient to elicit this sound, phonetically a nasal _peek_. The only other vocalization noted throughout the nestling period was an abbreviated _chee_.

For the first three days tapping the nest or even movement of it caused by wind would elicit begging. By the fifth day at nest 2-a (1959) only vigorous agitation of the branch to which the nest was attached evoked any response. At this nest on June 16, 1959, one young begged while the other cowered. Cowering is correlated with opening of the eyes, as the young bird that begged had its eyes only partly open.

Both young cowered on June 19, 1959. Table 9 summarizes the maturation of the nestling Bell Vireos.

TABLE 9. MATURATION OF NESTLING BELL VIREOS. THE FIRST DAY THAT AN ACTIVITY WAS OBSERVED IS SHOWN.

================================================================== | Day of nestling life +---+---+---+---+---+---+---+---+---+----+---- | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 | 11 --------------------+---+---+---+---+---+---+---+---+---+----+---- | | | | | | | | | | | Eyes open | | | | x | | | | | | | Feathers erupt | | | | | x | | | | | | Sound: Squeak | x | | | | | | | | | | _Chee_ | | | | x | | | | | | | Begging | x | | | | | | | | | | Cowering | | | | | | | | x | | | Head scratching and | | | | | | | | | | | Preening | | | | | | | | | x | | Hopping to rim of | | | | | | | | | | | nest | | | | | | | | | x | | Fledging | | | | | | | | | | |x[G]

[G] This is the commonest fledging day.

_Parental Behavior_

No eggsh.e.l.ls were found in nests on the days of hatching. Presumably they had been removed by the parents. Nolan (1960:234) indicates immediate disposition of the eggsh.e.l.l after hatching. Lawrence (1953:62) suggests that conspicuous removal of eggsh.e.l.ls by the female Red-eyed Vireo informs the male that the young have hatched.

Both s.e.xes brood and the exchange of partners resembles that described for the incubation period. Decrease in brooding in the daytime begins about the sixth day of nestling life. Nolan (1960:235) reports a sharp decrease in brooding when the oldest nestlings are seven days old.

Brooding decreases notably on the sixth day of nestling life in the Red-eyed Vireo (Lawrence, 1953:62). Nice (1929:17), Hensley (1950:244), and Nolan (1960:235) report that the female Bell Vireo a.s.sumes a slightly greater role in brooding than the male.

Apparent sun-shading was noted at nest 3-b (1959) at 2:00 p.m. on June 17, 1959, on the fifth day of the nestling period. The nest contained three young. An adult flew to the nest; while standing on its rim the bird dipped its head into the nest six times, afterward appeared to be eating a fecal sac, than shifted position to the unattached portion of the rim, gaped three times, thereupon spread its wings, and sat motionless 35 minutes. In this att.i.tude it formed an effective shield sheltering the young from direct sunlight penetrating the thin foliage of the honey locust in which the nest was situated.

The temperature at this time was 95 F., but the sky was partly cloudy. By 2:30 p.m. the sky had become overcast and the sun pa.s.sed behind a cloud. Although sunlight no longer fell directly upon the nest, the bird remained in the shielding posture for another five minutes before flying from its perch. Sun-shading was not observed at either of the other nests containing young; dense overhead vegetation protected those nests. Sun-shading has been noted in other species where the nest was poorly protected from the sun. Lawrence (1953:62) observed this behavior at two Red-eyed Vireo nests in conifers. The "sun-shield" posture of the Bell Vireo does not correspond to any of the sunning postures described by Hauser (1957).

_Feeding of the Nestlings_

Both s.e.xes fed the young, and presumably began shortly after the first nestling hatched. My data indicate that the female does more feeding than the male (Table 10); in about eight hours of observation a total of 67 morsels were brought, 43 by the female and 24 by the male, for an average of once every 7.6 minutes. Nice (1929:17), however, observed a male to bring food 53 times as compared to 21 visits by the female. In five and one-half hours of watching, meals were brought once every 4.9 minutes. Du Bois (_in_ Bent, 1950:257) recorded seven trips in an hour and forty minutes, or one every fourteen minutes.

At three nests containing young the adults were sometimes silent and sometimes vocal on their approach. The female often emitted a subdued _chee_ which, coupled with the vibration of the nest caused by her arrival, elicited begging behavior from the young. None of the males was heard to utter such a call, but I have the impression that they often did call although I failed to hear the sounds. The males did, on occasion, sing several songs as they approached, even with food held in their beaks. Such singing elicited begging from the nestlings. Once the eyes of the young were open they often began begging when a silent adult was within two or three feet of the nest; begging behavior probably is elicited by tactile, auditory or visual stimuli in that order, or, as the nestling period proceeds, by any combination of these stimuli.

TABLE 10. FEEDING OF THE NESTLINGS.

==================================================== Day of | Length of | Adult involved nestling period | observation +---------+--------- | | Male | Female -----------------+--------------+---------+--------- 1 | 30 min. | 3 | 5 2 | 60 min. | 1 | 4 3 | 60 min. | 2 | 5 4 | 30 min. | 1 | 4 7 | 60 min. | 4 | 7 2 | 60 min. | 3 | 3 6 | 60 min. | 3 | 6 7 | 30 min. | 3 | 3 9 | 60 min. | 4 | 6 +--------------+---------+--------- Totals | 510 min. | 24 | 43 -----------------+--------------+---------+---------

Not all trips made by parents resulted in successful feeding of young; some visits seemed to be purely for inspecting the young. On other occasions the adults experienced difficulty in transferring food to the young, and, thus thwarted, would themselves eat the food. Nice (1929:17) estimated that from five to twelve of a total of seventy-five meals were eaten by adults.

_Nest Sanitation_

Both parents regularly removed fecal sacs from the nest, eating them for the first five days and thereafter carrying them off and presumably dropping them. It is doubtful that fecal sacs were actively removed in the last two days of nestling life as the bottoms of nests from which young flew away were invariably covered with excrement.

On several occasions a parent brought food to the nest and then remained perched on the rim alternately peering into the nest and then preening. Once bill swiping was observed and another time an adult male sang once. The adult remained at the nest from twenty seconds to a full minute.

_Fledging_

Eight young were fledged from the four nests in 1959. The nestling period lasted from nine to twelve days. Human interference may have been largely responsible for the fledging of the young at nine days.

Pitelka and Koestner (1942:100) found nestling life to last eleven days. Nolan (1960:235) reports nestling periods varying from 10.5 to 12 days. The young Red-eyed Vireo is ready to leave the nest at ten days but often remains an additional day before departing (Lawrence, 1953:68).

The oldest nestling at nest 2-a (1959) hopped out on June 17, 1959, when I disturbed the parents. On this date the juvenal plumage was only partly developed and the young bird was incapable of flight. By the tenth day of nestling life the young in all the nests were observed to hop to the rim, flutter their wings, hop back into the nest and also to preen and scratch their heads. The young at fledging are usually completely feathered, but have notably short tails and relatively short, stubby wings. According to Ridgeway (1904:205) the juvenal plumage is much like that of the adult.

_Nest Parasites_

Pitelka and Koestner (1942:103) found that incubating adults and later the young suffered infestation of the northern fowl mite, _Ornithonyseus sylviarum_. Nolan (1960:241) reports a heavy infestation of this mite at four nests. Unidentified mites were noted at four nests in my study area in 1959. Incubating adults were observed to peck at their b.r.e.a.s.t.s and scapulars from the eleventh through the fourteenth day of incubation. Serious infestations were not noted at the nests until the ninth day of nestling life. At this time the young were observed to scratch their heads and peck at their b.r.e.a.s.t.s, scapulars, and the base of their tails. On the day of fledging the nests were a seething ma.s.s of crawling mites; the mites also extended well up the branches to which the nests were attached.

Nest 1-a (1959), which was discovered on June 18, 1959, presumably on the day after fledging, was densely covered with mites. Some mites were still crawling on this nest on June 20, 1959.

FLEDGLING LIFE

On June 20, 1959 I located one young 80 feet northeast of nest 2-a (1959), about five hours after it had left the nest. One parent was observed to feed it once. No young were seen thereafter from this or any other nest. Extreme agitation on the part of one or both parents on several occasions shortly thereafter, however, suggested the proximity of the young. Search in the immediate vicinity on each of these occasions proved fruitless. Three days after fledging their young, pair 2 (1959) was primarily occupied with courtship activities.

Pair 1 (1959) was involved in courtship and nestbuilding one and one-half days after the apparent fledging of their young. Nolan (1960:238) indicates that the young remain within the territory and perhaps are fed by the parents up until an age of about 40 days.

Sutton (1949:25) and Lawrence (1953:68) present contradictory reports on fledgling-parent relationships in the Red-eyed Vireo. Sutton concluded that the young quickly took leave of their parents whereas Lawrence reported a young bird being fed 35 days after fledging.

_Second Broods_

The curve based on 66 nesting records of the Bell Vireo representing the breeding activity in northeastern Kansas demonstrates a tendency toward double-broodedness (fig. 6). The peak of the breeding season is from May 20 to June 20. The large number (20) of replacement nests built in late May of 1960 tends to distort the curve of the breeding data; a second peak about 35 days after the first is evident.

I am of the opinion that the vast majority of vireos are single-brooded solely by virtue of the limited success of early nesting efforts, and that in "good" years most pairs would be double-brooded. Each of the four pairs that successfully raised one brood in my study area in 1959 renested within a day or two after the fledging of the young. I do not know the fate of these nests. Nolan (1960:237) reports at least one instance of a second brood in the course of his study. Nolan (_op. cit._) notes that the literature, in general, indicates that vireos are double-brooded, but that his evidence, mentioned previously, is the only evidence based on banded birds.

[Ill.u.s.tration: FIG. 6. Breeding season in northeastern Kansas based on the number of completed clutches in each 10-day period from May through July.]

REPRODUCTIVE SUCCESS

Only four nests were successful; all of these were observed in 1959.

The princ.i.p.al external factors responsible for nesting failure were severe weather, predation, parasitism by Brown-headed Cowbirds (_Molothrus ater_) and human interference (Table 11).

In late winter and early spring of 1960 heavy snow, continuously at a depth of at least 10 inches, covered most of the Mid-west from February 20 through March 20. Consequently, the growing season was some two weeks behind that of 1959. Of all the species in the study area, the Bell Vireo is the most dependent on dense foliage for cover and concealment for its nests. Consequently the tardiness of the season seemingly negatively influenced reproductive success of this more than any other species of bird in the study area.

_Behavior_

Several aspects of the behavior of the Bell Vireo tend to contribute to nesting failure. They include:

1. Nest-site. Nests are occasionally suspended from exposed branches.

Occurrences of this sort suggest that the dimensions of the fork are more important in the choice of a site than availability of cover.

2. Song. The loud, continuous song of the male during nestbuilding alerts cowbirds and predators to the presence of a nest. The incongruous habits of the male of singing in the nest tree and while sitting on the nest may facilitate location by some enemies, particularly cowbirds.

TABLE 11. EGG MORTALITY IN BELL VIREOS.