[F] These data have been supplemented from the literature pertinent to Kansas.
Caution is necessary in determining mean clutch-size in the Bell Vireo. Eggs occasionally disappear from the nest prior to or during incubation, without subsequent addition of cowbird eggs. Unfamiliarity with the history of such a nest on the part of the observer would lead to an inaccurate determination of clutch-size.
Complete clutches are not replaced with the same regularity as are nests. I have recorded intervals of six to thirty days between successive clutches. Successful replacement of clutches is determined by a number of factors: nest-site, completion of a nest, weather, predation, and parasitism by the cowbird. The difference between the number of renesting attempts and the successful replacement of clutches seems to indicate that different physiological processes are responsible for these two phenomena and that there is lack of synchrony between them. The development of the ovarian follicle requires a specific number of days that is not always coincident with the building of replacement nests. If, in the Bell Vireo, replacing a nest were solely a responsibility of the female, instead of involving the male to a considerable extent, it would seem likely that replacement of nests and the replacement of clutches would be more closely coordinated.
Incubation
Nice (1954:173) considers the incubation period to be the elapsed time between the laying of the last egg in a clutch and the hatching of that egg, when all eggs hatch. My data indicate that, normally, intensive incubation begins when the second egg is laid and lasts fourteen days in the Bell Vireo. Nice (1929:99) also considered the incubation period in this species to be fourteen days but believed it to commence when the third egg was laid. Pitelka and Koestner (1942:99) noted that the first and second eggs hatched fourteen days after laying of the second egg. However, they thought incubation began with the first egg. This would mean a fifteen-day period for this egg. All the eggs that Nolan (1960:234) marked hatched in approximately fourteen days. Eight eggs artificially incubated by Graber (1955:103) required an average of 15.01 days to hatch. As Van Tyne and Berger (1959:293) indicate, periods of sitting on the nest, even all night, do not necessarily mean that incubation has begun, for it has been demonstrated in several species that birds may sit on an egg without actually applying heat. My own observations demonstrate that the first egg may be left unattended for several hours at a time on the day that it is laid.
The Roles of the Sexes in Incubation
Both the male and female sit on the eggs in the daytime. My study of histological sections of ventral epidermis indicates that the male does not possess a brood patch; the increased vascularization typical of the brood patch in females is not evident in males. But, the male loses most of the down feathers of the ventral apterium. Also, according to Bailey (1952:128), the male Warbling Vireo that sits on the eggs lacks a brood patch.
Bailey (1952:128) suggests that male passerines lacking brood patches that habitually sit on eggs do not heat the eggs. Thus it cannot be considered true incubation since no increase of temperature in the eggs is effected by such means. He further notes that it is at night when eggs are likely to experience a drop in temperature that embryonic development will be impaired. I have no data directly pertaining to which sex sits at night, but it is presumably the female, because she is always seen on the nest early in the morning and late in the evening.
Fig. 4. Comparison of periods of incubation by both sexes in cold (54° F.) rainy weather (A) and in warm (82° F.) sunny weather (B).