Richard F. Johnston

“ DISTRIBUTION OF BIRDS IN KANSAS
Birds breeding in Kansas are taxonomically, ecologically, and distributionally diverse. Such diversity is to be expected, in view of the mid-continental position of the State. Characteristics of insularity, owing to barriers to dispersal and movement, tend to be lacking in the makeup of the avifauna here. The State is not, of course, uniformly inhabited by all 176 species (Table 1) of breeding birds; most species vary in numbers from one place to another, and some are restricted to a fraction of the State. ”

“ Two adult salamanders not included in Table 1 were found, in the course of examination for parasites, to have empty stomachs. One was a male, and the other was a female taken from a chamber that held an egg cluster. It would not be surprising regularly to find stomachs empty in "incubating" females, but the fact is that the one other such female collected by us had a small amount of food in the gut; probably these individuals take anything that enters the egg chamber, but do not leave for active pursuit of food. ”

“ 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. [Pg 595] 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. ”

“ Subterranean winter inactivity is thought to be an integral part of the salamanders' lives, and is suggested in part [Pg 585] by the life cycles of the worms. Summer activity appears to occur at the ground surface in logs and talus, and underground; the latter site is suggested by certain ratios obtained in the samples, showing adults to outnumber young and males to outnumber females. The season for egg deposition seems to be in July and August. Clutch-size is lower than for any other plethodontid on record. "Incubation" of eggs apparently parallels that characteristic of other plethodontids. ”

“ 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. ”

“ Table 4.—Sex Ratios in Aneides hardii, Adults
[Pg 582]
Age-ratio
The data in Table 5 show adult salamanders to outnumber young at each collecting locality. This is probably not an accurate reflection of actual age composition in this species. Yet, we obtained the same general result in all three years of the study. We assume, therefore, that young were located where we could not catch many of them; probably they were underground. Sites of hatching and of the activities of early life would thus occur where we think the bulk of eggs are laid. ”

“ 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. ”

“ If it is true that parasites are reacquired each spring—we assume that no temperature factors or immune reactions are delaying development of the worms, and no unusually long external ovic or free-living phase is a necessary part of their life-history—then the host-parasite data can be used as a basis for hypothesizing about the winter life of the salamander. ”

“ The regular recurrence of warm gulf air is responsible for the characteristically high relative humidity in summer over eastern Kansas and it has an ameliorating effect on winter climate in this region. Almost immediately to the north in Nebraska and to the west in the high plains, summers are dryer and winters are notably more severe. The breeding distributions of some species of birds fairly closely approximate the distribution of these warm air masses ”

“ During "surface" life the incidence of parasitism is high (90 per cent and 83 per cent: see Table 2) , indicating that salamanders are readily invaded in times of activity. Salamanders examined in September were all parasitized and probably carried nematodes with them into their winter retreats. This part of their habitat should thus be contaminated with infective stages of both parasites. Yet the salamanders seem to become re-infected when the period of summer activity starts (note the high incidence of immature parasites in salamanders taken in July) ”

“ The present work treats mainly certain temporal relationships of breeding birds in Kansas, but also geographic distribution, habitat preferences, and zoogeographic relationships to the extent necessary for a useful discussion of temporal breeding phenomena.
Information on breeding of some of the 176 species of birds known to breed in Kansas is relatively good, on a few is almost non-existent, and on most is variously incomplete. It is nevertheless possible to make meaningful statements about many aspects of the breeding biology and distribution of most species of Kansan birds ”

“ In the summer rainy season A. hardii lives in and under downed timber and under talus accumulations. Occurrence, however, seems to be partly subterranean and always local; seemingly good habitat frequently appears to lack the animals. Our observations and collections were made in July, August, and September in 1956, 1957, and 1958. Two hundred seventy-seven individuals were taken; these were measured, sexed and examined for breeding status. The food and parasite content of the guts of a few individuals was determined. Thirteen salamanders were kept for varying lengths of time in captivity. ”

“ 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. ”

“ One hundred one species of Kansan birds are woodland species (tables 1 and 2) . The analysis of Udvardy (1958) showed woodland birds to be the largest single avifaunal element in North America, with 38 per cent of North American birds relegated to it. It is likewise the largest element in the Kansan avifauna, representing 58 per cent of Kansan [Pg 582] birds. Although woodland makes up a relatively small fraction of the vegetational complexes in Kansas, a large number of habitats exist in what woodland is present. ”

“ Nevertheless, less than half the stomachs contained ants; this may mean that salamanders do not make an effort to take ants over any other prey. Such foraging behavior would result in random capture of ants, and it is noteworthy that the frequency distribution of ants in stomachs suggests a Poisson distribution, a mathematical description of one kind of random distribution.
Table 1.—Numbers of Food Items Found in Stomachs of 16 Specimens of Aneides hardii
Adult and larval beetles comprised about 28 per cent of the total items, but were found in only seven of the stomachs. ”

“ Some individuals, usually in small groups, can be seen throughout the winter in eastern Kansas, and their presence makes it difficult to document dates of arrival and departure of the strictly summer resident birds; these can be said to arrive in March and to leave in October, but these indications are the barest approximations.
Breeding schedule.—The 334 records of breeding span the period April 1 to July 20 (Fig. 7) ; the modal date of laying of first clutches is April 25, but subsequent peaks are indistinct. Nearly half of all eggs are laid in the period April 11 to 30. ”

“ Breeding activities are ordinarily associated in time with greatest food abundance. [Pg 583]
Clutch-size
By clutch-size we refer to the number of eggs in laid clusters. We collected clutches of six, four, four and one; adding one more of three (Lowe, op. cit.) gives an average of 3.6 eggs per cluster; the average is 4.2 eggs if our clutch of one is discarded on the grounds it was incomplete.
For comparison we have listed (Table 6) clutch-sizes for some other plethodontids. It should be noted that these numbers refer only to eggs deposited in clusters, and not to large ovarian eggs. ”

“ Twenty-three species of our total can be called grassland species (Table 1) . The subtotal is less than one-fifth of the Kansan avifauna, but it represents 72 per cent of the grassland birds of North America; grassland habitats abound in Kansas. Only 5.3 per cent of all North American birds are grassland species (Udvardy, 1958) .
About 56 per cent of these birds are of North American stocks, nine per cent of Eurasian stocks, and three per cent of South American stocks. The percentage of North American species is the greatest for any habitat group here considered. ”

“ Icterus galbula (Linnaeus) .—This common summer resident is most numerous in the east, in woodland and riparian timber. The species hybridizes freely with the Bullock Oriole in western Kansas, and individuals morphologically typical of Baltimore Orioles are rare west of the 100th meridian. Evidence of such hybridization can be found in specimens taken in eastern Kansas, but the linear nature of distribution along water-courses to the west restricts gene-flow, and evident hybrids are not yet conspicuous. ”

“ 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.—Relationship Between Environmental Factors and Timing of Breeding in Birds of Kansas
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. ”

“ Such corrected dates can be used in making histograms that show the time of primary breeding activity, or the "egg-season." All such schedules are generalizations; data are used for a species from any year of observation, whether 50 years ago or less than one year ago. One advantage of such procedure is that averages and modes are thus more nearly representative of the basic temporal adaptations of the species involved, as is explained below. ”

“ Conspicuous breeding activity occurs from the first third of May to the last third of June.
The Influence of Dominant Foraging Adaptation.—Five categories here considered reflect broad foraging adaptation: woodland species, taking invertebrate foods in the breeding season from woody vegetation or the soil within wooded habitats; grassland species, taking invertebrate foods in the breeding season from within grassland situations; limnic species, foraging within marshy or aquatic habitats; aerial species, foraging on aerial arthropods; raptors, feeding on vertebrates or large insects. ”

“ 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. ”

“ 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. ”

“ Twenty-four species, furnishing 875 records of breeding, are here considered to be resident birds in northeastern Kansas. These species are Cooper Hawk, Red-tailed Hawk, Prairie Chicken, Bobwhite, Rock Dove, Great Horned Owl, Red-bellied Woodpecker, Hairy Woodpecker, Downy Woodpecker, Horned Lark, Blue Jay, Common Crow, Black-billed Magpie, Black-capped Chickadee, Tufted Titmouse, Carolina Wren, Bewick Wren, Mockingbird, Eastern Bluebird, Loggerhead Shrike, Starling, House Sparrow, Eastern Meadowlark, and Cardinal. ”

“ 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. ”