[Table 9] shows that there are two basic groups of birds according to peak of egg-laying and incidence of precipitation; raptors, birds of Eurasian origin, resident birds, and birds of grassland habitats tend to have their peaks of egg-laying prior to the peak of spring-summer rains, and the other six categories tend to have their peaks of egg-laying occur in the time of spring-summer rains. Regarding temperature, there are four categories of birds; these are evident in the table.
Some of the correspondences deserve comment. Residents and grassland species both breed before the rains come and before mean temperatures reach 70°F., and this correspondence probably results from most of the grassland species being residents. Contrariwise, most birds of Eurasian stocks are residents, but not all residents are of such stocks; the two groups are discrete when mean temperature at breeding is considered. Woodland birds, aerial foragers, and birds of South American evolutionary stocks breed after temperatures surpass 70°F. on the average. Almost all such species are migrants, but many migrants have different temporal characteristics, and the categories thus are shown to be discrete on the basis of temperature at time of breeding. The change through spring and summer of temperature and precipitation delineates the inception and waxing of the growing season of vegetation and of the subsequent arthropod populations, on which most of the birds feed in the breeding season. The temporal characteristics of growing seasons in North America have been treated by Hopkins (1938) and have been related to timing of breeding seasons in Song Sparrows (Passerella melodia) of the Pacific coast of North America (Johnston, 1954).
Significance of Phylogeny to Breeding Schedules
Evidence from a variety of sources demonstrates that timing of breeding seasons is either broadly or specifically genetically-determined. For some species in some situations major environmental variables are paramount in regulating timing of breeding, but in others the innate, regulatory "clock" is less closely tied to conspicuous exogenous stimuli. The work by Miller (1955a, 1955b, 1960) with several species of Zonotrichia strongly indicates that endogenous timing is most important for these birds, and there is ecological evidence for Song Sparrows that supports the same point (Johnston, 1954, 1956). It is, in any event, possible to treat breeding schedules as species-specific characters, for any one geographic area.
In an attempt to relate a breeding schedule to previous ancestral modes, that is by extension to phylogeny, it is necessary to know how often ancestral adaptations can persist in the face of necessity to adapt to present environmental conditions. It is necessary to know how conservative or how immediately plastic breeding schedules can be. The disadvantage of using available information about configurations of breeding seasons (as shown in Figs. 3 to 9) is that it is extremely difficult to compare visually at one time more than six or eight histograms as to the trenchant similarities and differences regarding times of inception and cessation of breeding, and time of peak egg-laying. It is possible, however, to reduce these three variables to one variable (as described below), which allows the necessary comparisons to be made more easily; this variable may be called the breeding index.
Calculation of Breeding Index
The chronological year is broken roughly into ten-day intervals numbered 1 to 36. The histogram describing the temporal occurrence of the breeding season of a species in our area usually will lie within intervals 7 to 25. The modal date for completion of clutches is given a value corresponding to the number of ten-day intervals beyond interval 7 (March 1-10); this describes the modal variable. The date of completion of 83 per cent of all clutches is given a value corresponding to the number of ten-day intervals it lies from interval 11 (April 11-20); this describes the 83 per cent variable (and is a measure of the length of the season in terms of its inception). The breeding index can then be calculated as follows:
I = Xm + Xsd,
| where: | I is the breeding index, |
| Xm is the modal variable, and | |
| Xsd is the 83 per cent variable. |
This is obviously an arbitrary scheme to gain a simple measure of beginning, peak, and end of a breeding season. Other schemes could be devised whereby different absolute values would be involved, but the relative nature of the results would be preserved. The values under the present system for 73 species of Kansan birds run from -5 to +22; early modal dates and cessation to breeding give low values, late dates high values.