Part 6

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 = X{m} + X{sd},

where: I is the breeding index, X{m} is the modal variable, and X{sd} 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.

Within this framework there are other, presumably subordinate, factors that influence the values of breeding indices, as follows:

1. Migratory habit. Any migrant tends to arrive on breeding grounds relatively late, hence migrants ordinarily have higher index values than do residents.

2. Colonial breeding. The strong synchrony of colonially-breeding species tends to move the modal egg-date toward the time of inception of breeding; as a result colonially-breeding species probably have lower index values than they would have if not colonial.

3. Single-broodedness. Species having only one brood per season tend to have shorter seasons than double-brooded species, and their index values tend to be lower than those of double-brooded species.

Migratory habit unquestionably has considerable influence on index values in some species. It is not, however, as important as other matters, such as the condition of the food substratum or sensitivity of the pituitary-gonadal mechanism, in determining timing and mode of breeding activity. The schedule of the Purple Martin is the extreme example showing that time of spring arrival on breeding grounds is not necessarily related to time of inception of breeding. It should be emphasized that the factors leading to northward migratory movement may be involved in timing of the annual gonadal and reproductive cycle.

Figure 2 presents a graphic summary of values of breeding indices for many groups of Kansan birds. The values for species of a given family have been linked by a horizontal line. The length of this line is proportional to the degree to which the index values for the species concerned resemble one another. Note that the plottings for the Picidae, Corvidae, Turdidae, Tyrannidae, and Icteridae each contain one point that is well-removed from a cluster of points. This can be interpreted as a measure of the frequency of adaptive plasticity versus adaptive conservatism; five of the 24 plottings show a plastic character, 19 a conservative. There are 26 plottings that show temporal consistency, all of which may be taken as evidence of adaptive (or relictual) conservatism of the species in question.

Conclusion

Such patterns of breeding chronology support the idea that seasonal response to the necessities of breeding is conservative more often than plastic. Most students of breeding schedules believe that since these are highly adaptive, they must also be capable of flexibility to meet variable environments within the range of the species. Such thinking receives support when different geographic localities are considered for one species (Johnston, 1954), or when specific features of a special environment are considered (see Miller, 1960; Johnston, 1956).

Yet, if one, relatively restricted locality is considered, as in the present study, evidence of a conservative characteristic in breeding schedules can be detected. This conservatism may result from the historic genetic "burden" of the species; that is to say, previous adaptive peaks may in part be evident in the matrix of contemporary adaptation. Adaptive relicts of morphological nature have been many times documented, but characteristics associated with seasonality and timing schedules have not.

In any event, genetic relationships are evident in the configuration of breeding seasons of many species here treated. Thus, any consideration of variation in breeding schedules must be sensitive to the limits, whether broad or restricting, that the heritage of a species sets on its present chronological adaptation.

Regulation of Breeding Schedules

Regulation of breeding schedules in birds always involves some exogenous, environmental timing or triggering mechanism. Broad limits to functional reproductive activity seem to be set by the photoperiod--neuroendocrine system. This basic, predominately extra-equatorial, regulator can be ignored by temperate-zone species only if they possess chronological adaptation to special, aperiodic environmental conditions, as does the Red Crossbill (Loxia curvirostra; see McCabe and McCabe, 1933; H. B. Tordoff, ms.), for which the chief consideration seems to be availability of conifer seeds. Environmental phonomena otherwise known to trigger breeding activity include rainfall (Davis, 1953; Williamson, 1956), presence of suitable nesting material (Marshall and Disney, 1957; Lehrman, 1958), temperature (Nice, 1937), and presence of a mate (Lehrman, Brody, and Wortis, 1961). Such regulators, or environmental oscillators, are the "phasing factors" of the physiologic clock that dictate the temporal occurrence of primary reproductive activity.

TABLE 9.--RELATIONSHIP BETWEEN ENVIRONMENTAL FACTORS AND TIMING OF BREEDING IN BIRDS OF KANSAS

==================+=================================================== | Occurrence of Peak of Egg-laying +-------------------+------------------------------- | When | When Mean | Precipitation is: | Temperature (F.) is: +---------+---------+-------+-------+-------+------- | Light | Heavy | < 55 deg. | < 70 deg. | +- 70 deg. | > 70 deg. ------------------+---------+---------+-------+-------+-------+------- Raptors | x | | x | | | O. W. Element | x | | x | | | Residents | x | | | x | | Grassland species | x | | | x | | Marshland species | | x | | | x | N. Amer. Element | | x | | | x | Migrants | | x | | | x | Woodland species | | x | | | | x Aerial foragers | | x | | | | x S. Amer. Element | | x | | | | x ------------------+---------+---------+-------+-------+-------+-------

None of the regulators mentioned above has been specifically investigated for any Kansan bird, but it is reasonable to suppose that, in these temperate-zone species, the photoperiod is the most important general phasing factor in seasonal breeding. Although gonadal response and seasonal restriction of breeding are set by the photoperiod, specific temporal relationships are dictated by more immediate environmental variables.

Table 9, as already noted, shows the gross relationships between certain groups of birds, certain arbitrary indicators of seasonal temperature-humidity conditions bearing significantly on the growing season, and occurrence in time of peak of egg-laying by the birds involved. Some species and groups of Kansan birds breed chiefly under cool-dry environmental conditions, and some under warm-wet environmental conditions. Within each of these categories some variation occurs. Thus, raptors and boreally-adapted species (the Eurasian zoogeographic element) breed under cool conditions prior to rains, and residents and grassland species breed under slightly warmer conditions prior to rains; limnic species, species derived from North American evolutionary stocks, and migrants tend to breed in the cooler segment of the warm-wet period, and woodland birds, aerial foragers, and species derived from South American evolutionary stocks tend to breed in the warmer segment of the warm-wet period.

So much, then, for relationships between birds and their environments at a descriptive level. It would be useful at this point to examine how environmental variables relate to timing of breeding. Certain independent lines of investigation indicate that birds have a well-developed internal timing device; most convincing is the work of Schmidt-Koenig (1960) and the others who have shown that the endogenous clock of birds can be shifted in its periodicity forward or backward in time. This and much other evidence (see Brown, 1960) indicate that many fundamental periodic regulators are extrinsic to the animal; it is thus permissible for present purposes to consider any expression of variation in timing as dependent on environmental oscillators. It is not hereby meant to ignore the fact that differential responses to dominant environmental variables occur within a species, indicating endogenous control over timing of breeding. The work by Miller (1960:518) with three populations of the White-crowned Sparrow, revealing innately different responses to vernal photoperiodic increase, is especially important in this regard. For the moment, however, we may consider exogenous controls only.

Any exogenous control, or environmental variable, can be looked on simply as a timing oscillator. Such variables show regular or irregular periodic activity, and the independent actions as a whole result in the more-or-less variable annual schedule of breeding for any species at any one place. It would seem that some oscillators are linked to one another, but there is a real question concerning the over-all degree to which linkage is present. It is significant that frequency distributions of breeding activity of various species and groups of birds take on the shape of a skewed normal curve. The more information is added to such distributions, the more nearly they approach being wholly normal, with irregularities tending to disappear. This kind of response itself is evidence that most of the variables influencing the distribution are not mutually linked.

This conclusion is warranted if we examine what would happen to frequency distributions if the variables or oscillators regulating timing were linked. The frequency distribution of breeding activity in birds is described by a nonlinear curve (a normal distribution is nonlinear). Let us assume that each of the environmental variables is a nonlinear oscillator, as is probable. A set of nonlinear oscillators mutually entrained or coupled and operating with reference to a given phenomenon would result in that phenomenon being described by a frequency distribution much more stable than if it were regulated by any one oscillator alone. However, the frequency distribution of a set of coupled nonlinear oscillators is non-normal (Wiener, 1958).

We do not obtain such distributions in describing breeding activity, so we may say that the oscillators regulating such activity are not coupled. Present distribution, habitat preference, residency status, foraging adaptation, previous zoogeographic history, and relicts of ancestral adaptation, all bear on the character of the breeding schedule of any bird species. The emphasis above on multiple regulation of breeding schedules conceivably reflects the true picture, but any such emphasis is made at the expense of taking one factor as basic, or reducing the many to one, in order to manufacture simplicity.

ACCOUNTS OF SPECIES

In each account below information is given concerning status, habitat, geographic distribution, seasonal occurrence, schedule of egg-laying, number of eggs laid, and sites of nests, as these pertain to Kansas, unless otherwise stated. The ways in which some of these points were elucidated are as follows.

1.--Breeding schedule. Frequency distributions of egg-laying in time are calculated on the basis of dates of completed clutches, as described earlier (p. 588). Any event in the series of actions of nesting--nestbuilding, egg-laying, incubation, brooding, feeding young out of nests--can be manipulated by adding or subtracting days to or from the date of record to yield the probable date of completion of the clutch. The resulting data are grouped into class intervals of ten days. Extreme dates here given for egg-laying may be as much as nine days off in accuracy, but the error does not often exceed five days. Extreme dates indicated here may be taken as actual or predicted extremes. The raw data used are on file at the Museum of Natural History and are available for use by any qualified individual.

2.--Dates of occurrence. First and last annual occurrences in the State for migrant species are indicated by both a range of dates and a median date. Twenty to 30 dates of first observation in spring are available for most of the common species, and 10 to 20 dates of last observation in autumn are at hand for such species. The median dates, earlier than and subsequent to which an equal number of observations are available, are reliable indicators of the dates on which a species is likely to be seen first in the State in an average year.