John M. Legler

“ It seems remarkable that none of the species of true tortoises occurring in the grasslands of the world has developed insectivorous habits or utilized the unique food niche (in regard to dung-foraging) filled by ornate box turtles in the Great Plains; tortoises are, as far as is known, strictly herbivorous. The ranges of Gopherus and Terrapene are now almost mutually exclusive and the two kinds do not compete with each other for food in the few places where they occur together. ”

“ The only methods that I know of to determine successfully the longevity of long-lived reptiles would be to keep individuals under observation for long periods of time or to study populations of marked individuals. Both methods have the obvious disadvantage of requiring somewhat more than a human lifetime to carry them to completion. Restudy, after one or more decades, of the populations of turtles marked by Fitch and myself may provide valuable data on the average and maximum age reached by T. ornata. ”

“ Early hatching and precocious development presumably confer an advantage on the individual, since turtles that grow rapidly are able better to compete with smaller individuals of the same age. Theoretically, turtles growing more rapidly than usual in the first two or three years of life, even if they grew subsequently at an average rate, would attain adult size and sexual maturity one or more years before other turtles of the same age. ”

“ Nests are dug in the earth by the female using her hind legs. Preferred nesting sites are open, well-drained places with a soft substrate. The nesting site is selected after a period of wandering, in which the female tests the substrate at a number of places; some females search for a nest site for more than a week. Nest digging begins in the evening and is usually completed after dark. Captive females dug a preliminary cavity in which the body rested during the digging of the main nest cavity. The entire clutch of eggs is laid in one nest. ”

“ Practically nothing is known about longevity in T. ornata or in other species of Terrapene although the several plausible records of ages of 80 to more than 100 years for T. carolina (Oliver, 1955:295-6) would indicate that box turtles, as a group, are long-lived. There is no known way to determine accurately the age of an adult turtle after it has stopped growing. It was possible occasionally to determine ages of 20 to 30 years with fair accuracy by counting all growth-rings (including those crowded into the interabdominal seam) of specimens having unworn shells. ”

“ Even in years when summer temperatures are much cooler than normal, eggs probably hatch by the end of October. Hatchlings or eggs would have a poor chance of surviving a winter in nests on exposed cut-banks or in other unprotected situations. Overwintering in the nest, hatchlings might survive more often than eggs, since hatchlings could burrow into the walls and floor of the nest cavity. Unsuitable environmental conditions that delay the nesting season and retard the rate of embryonic development may, in some years, be important limiting factors on populations of ornate box turtles. ”

“ Several females at the Damm Farm traveled long distances from their home ranges to nest but other females nested within their known or estimated home ranges. Seemingly a complex of environmental factors, including soil texture, weather, availability of water, and possibly the urge for random wandering in the breeding season, governs the distances traveled by gravid females and the ultimate selection of a satisfactory nesting site. Females, because of their more extensive travels in the nesting season, seem more likely than [Pg 635] males to have multiple or changing home ranges. ”

“ Activities of man have probably affected the density of populations of the ornate box turtle in many parts of its range but appear not to have acted as limiting factors except in certain areas along the northern edge of the range (Blanchard, 1923:19-20, 24) where disruption of grassland through intensive cultivation probably has excluded the species. Unlike certain other reptiles of the Great Plains (Fitch, 1955:64) , T. ornata seems not to have been affected—either by direct decimation of populations or by disruption of habitat—by intensive zoological collecting in restricted areas. ”

“ The ovarian cycle begins in midsummer, soon after ovulation, and continues up to the time of the next ovulation. Follicular growth is rapid in the period from spring emergence to ovulation. Large follicles remaining after ovulation represent, in many instances, eggs that will be laid later in the same season. Follicular atresia is never great enough to account for the destruction of all large follicles remaining after ovulation. All mature females lay at least one clutch of eggs per year. It is estimated that one-third of the females produces two clutches of eggs in a single season. ”

“ Piles of dung, disturbed by turtles in the course of their foragings, were characteristic "sign" of T. ornata in the areas studied.
Insects form the bulk of the diet for most of the year, although certain other foods, when especially abundant for short periods (mulberries for example) , are eaten in large quantity or eaten to the exclusion of all other foods. Ornate box turtles occasionally eat the eggs and young of ground-nesting birds and slightly damage vegetables, but in no instance do these feeding habits significantly affect the economy of man. ”

“ Observations by William R. Brecheisen and myself on winter activity of aquatic turtles indicate that, in Anderson County, Kansas, the commoner species (Chelydra serpentina, Chrysemys picta, and Pseudemys scripta) are more or less active throughout the year; although they usually do not eat in winter, they are able to swim about slowly and in some instances (P. scripta) even to carry on sexual activity at body temperatures only one or two degrees above freezing. But, ornate box turtles hibernating in the ground a few yards away are incapable of purposeful movement at such low body temperatures. ”

“ Records for males and females are combined.
Attainment of sexual maturity, in the population studied, was more closely correlated with size than with age. For example, nearly all males were mature when the plastron was 100 to 110 millimeters long, regardless of the age at which this size was attained. The smallest mature male had a plastral length of 99 millimeters; according to the data presented in Figures 9 and 10, therefore, a few males reach sexual maturity in the fourth year, and increasingly larger portions of the population become mature in the fifth, sixth, and seventh years. ”

“ Frequent imbibing of water is probably necessary if the search for a nesting site is continued for more than a day or two. Standing water was usually available in ponds, ravines, ditches, and other low areas at the Damm Farm in June. Nesting in June, therefore, is advantageous not only because of the greater length of time provided for incubation and hatching but also because of the amount of water available for drinking. Females can probably be more selective in the choice of a nesting site if their explorations are not limited by lack of water. ”

“ No clearly defined external characteristics of senility were discovered in the populations studied. A male that I examined had one atrophied testis. In another male both testes were shrunken and discolored and appeared to be encased by fibrous tissue. Both males were large, well past the age of regular growth, and had smoothly worn shells. Several old females had seemingly inactive ovaries. Reproductive processes probably continue throughout life in most members of the population, although possibly at a somewhat reduced rate in later life. ”

“ I suspect that, in these old individuals, the germinal layer of the epidermis does not become active each year but retains the capacity to elaborate new epidermis if the shell becomes worn thin enough to expose and endanger the bone beneath it. The germinal layer of old turtles loses the capacity to produce color. ”

“ Ornate box turtles are able to detect the presence of intruders, by sight, from a distance of several hundred feet in open country; apparently, intruders are not detected until seen. Defensive behavior is passive; the shell is closed tightly in response to painful stimuli and, in some instances, at the sight of an intruder. Juveniles usually void odoriferous fluid from the musk glands when handled but adults do so only in response to pain or injury. ”

“ Body temperatures one to three degrees higher were common in the coldest part of the winter. Turtles in shallow hibernacula, like those observed in wooded areas at the Reservation, are probably subjected to freezing temperatures at least for short periods but I have no records of body temperatures this low, except where they were induced experimentally. Turtles exposed to temperatures of zero degrees or slightly lower would retain enough heat to survive without freezing for a period of several hours or even a day if well insulated. A temperature gradient exists within the body ”

“ Of the hind feet, the medial surfaces are the principal parts that touch the ground but some traction is derived from the hind claws at the beginning of each cycle of the hind leg. Under normal conditions, box turtles move slowly and pause to rest and examine their surroundings every few feet. When resting, the plastron is in contact with the ground, the legs relaxed, and the head and neck are extended upward. Some turtles seeking shelter from the heat of sunshine walk rapidly for a hundred feet or more without pausing. ”

“ Sternotherus, Terrapene, and Thamnophis are alike in completing spermatogenesis late in the season and storing spermatozoa, in the seminiferous tubules or in the epididymides, during hibernation.
It is noteworthy that, in the turtles and snakes mentioned above, sperm produced in autumn are used to fertilize eggs laid in the following year, and mating [with the exception of Thamnophis elegans, (Fox, 1956) ] occurs in both spring and autumn. It is not definitely known in any of these instances, whether sperm resulting from autumn or spring inseminations (or both) fertilize the eggs. ”

“ Ornate box turtles are solitary except during periods of mating. Meetings with other individuals in the course of foraging, basking, or seeking shelter, are fortuitous and have no social significance. A broad overlapping of home ranges of both sexes at the Damm Farm suggests that box turtles do not intimidate other individuals in the home range or exclude them from it. ”