Fig. 28.The movements of an adult (non-gravid) female ofT. o. ornatain the house pond area at the Damm Farm during a period of 24 days in July, 1955 (solid line), and a period of three days (broken line) in July, 1956. Solid dots represent the points where the turtle was found as her thread trail was mapped; hollow symbols represent points of recapture when no trailing thread was attached to the turtle.The actual home range of almost every individual studied, even of those individuals for which the most data were available, probably differed at least slightly from the observed or estimated home range. One adult female, for example, was captured six times intwo years within a radius of approximately 50 feet. Another female was found 2780 feet from her last point of capture. These last two records were regarded as unusual; when they were grouped with records of the 44 individuals mentioned above, the average radius of home range for the entire group was much larger (327 feet).Fig. 29.The movements of a gravid female ofT. o. ornatain the southern ravine area at the Damm Farm in a period of ten days in June, 1956. Her movements were, for the most part, in and around several ravines (shown on map by broken lines) where she was searching for a nesting site. For explanation of symbols see legend forFig. 28.Homing BehaviorGould (1957) reported that 22 of 43T. carolinamoved in a homeward direction when they were released in open fields up to 5.8 miles from their original points of capture. Turtles oriented themselves by the sun; homeward headings were inaccurate or lacking on overcast days and, light reflected from a mirror caused turtles to alter their courses. Seven of ten turtles released more than 150 miles from home headed in directions that corresponded most nearly to the headings last taken (at release-points near home base) and did not necessarily correspond to the direction of home. Gould's studies point out that box turtles perhaps practice a kind of "solar navigation." His work raises the question of whether the movements of box turtles are guided by the sighting of local landmarks or whether such landmarks alter the course of movement only when acting as barriers.In the present study two experiments were made to determine the homing ability ofT. ornata. An adult male, taken from his normal home range in the house pond area and released 1200 feet away in the southern ravine area, traveled a generally northward course (not northeastward in the direction of home) for five days, moving a distance of approximately 1900 feet. His detached trailer was recovered several days later 740 feet southeast of the last known point in his travels (a distance that could have been covered in two days) and 150 feet from the point of original capture; he had returned to his home range by a circuitous route in a period of approximately seven days. Another adult male, captured in the southern ravine area, and released in the house pond area 1900 feet away, traveled on a course that bore approximately 25 degrees north of true homeward direction; after five days he was approximately 600 feet north of the original capture point. He then began a northeastward course that took him back to the house pond area where he remained for several days; no further data are available for this individual. It is significant that the homing males discussed above traveled greater average distances per day (based on records for nine days of trailing) than any of the other turtles studied (Fig. 27). Fitch (1958:101) released an individual one half mile from where he captured it and, one year later, recovered the turtle near the point of release.Social RelationshipsOrnate 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. No instances of fighting were observed.Allard (1935:336), Perm and Pottharst (1940:26), and Latham (1917) recorded instances of fights between individuals ofT. carolina; in the latter two instances fights were between males. Stickel (1950:362) observed an incident between two males that may have been a fight; however, she was of the opinion that fights rarely occur in nature and that box turtles do not defend territories. Evans (1954:23-25) considered the behavior ofT. carolinareported by Perm and Pottharst (loc. cit.) to represent "territoriality." He found "… a true hierarchy…." existing betweenfour captive males ofT. carolinaand another between three captive females of the same species; young individuals in the group raised their social level in the hierarchy after receiving experimental doses of male hormone. Evans (op. cit.:25) pointed out that true tortoises (family Testudinidae) have a more complex pattern of social behavior than do emyid turtles.Observations made with binoculars from the vantage point of a blind provide the only information that I have concerning the reactions of box turtles to one another under natural conditions. Turtles foraging in a bare area were not startled by the approach of other turtles, and turtles moving across the area seemed to take no notice of turtles already there, regardless of whether these turtles were moving or not. Adults and subadults behaved in approximately the same manner.Individuals traveling or foraging in rough terrain or in grassy areas probably are unable to see each other even when they are close to one another. Conversely, box turtles can see each other and are surely aware of each other's presence in bare, flat areas. These facts suggest that no social hierarchy exists inT. ornata. On one occasion an adult male and a juvenile (hatched the previous autumn) were found foraging next to one another on the same pile of cow dung.When an individual became motionless in an attitude of wariness after having detected me in my blind, its behavior evoked no response on the part of other turtles, a few feet away.INJURIESFire, freezing, molestation by predators, and trampling by cattle or native ungulates are only a few natural sources of injury to which box turtles have always been exposed. Man's civilization in the Great Plains, chiefly his automobile and other machines, have compounded the total of environmental hazards. Automobiles now constitute a major cause of death and serious injury to box turtles. Each year thousands are struck on Kansas highways alone, not to mention the many casualties resulting from mowing machines, combines, and other farm machinery.Although grass fires usually occur in early spring or late fall when box turtles are underground, some turtles are surely killed by fires and many are injured. In early April of 1955 the pasture at the Damm Farm was burned. Similar burnings, I discovered, had occurred both intentionally and accidently in past years atirregular intervals. No deaths or injuries, attributable to fire were discovered in the course of intensive field work in the spring and summer of 1955, when the new grass was short and conditions for finding and marking box turtles were ideal. Badly burned individuals, if any, may have secreted themselves until their wounds had healed. In June, 1957, an adult female, that had been burned severely, was taken from a small puddle in a ravine on the Damm Farm. The soft parts of her body, excepting her head and neck, were a nearly solid mass of smooth scar tissue, the scales and rugosities of the skin being practically obliterated. The tail was reduced to a mere knob surrounding the anus and dead, exposed bone was visible on most of the dorsal part of the carapace. Possibly this female was burned in the fire of 1955. Lack of injury to the head and neck can probably be accounted for by the additional protection afforded these parts by the folded forelegs when the turtle was withdrawn in the shell.Turtles that are smashed flat on the highway, of course, have no chance of survival. Highway fatalities are usually the result either of "direct hits," where the tire of a vehicle passes directly over the turtle, or of repeated pummeling by subsequently passing vehicles. The writer, while driving behind other cars that struck turtles or by sitting beside roads, has observed numerous turtle casualties. Most are struck a glancing blow by a tire and are propelled some distance through the air or on the surface of the pavement, often to the side of the road. Such a blow is usually sufficient to crack or chip the shell, or at least to scuff away parts of the epidermal covering. Turtles, so injured, usually survive.Parts of the shell do not break away easily, even when several deep cracks are present, and only a little bleeding occurs. A common injury inflicted on the highway is the wrenching and subsequent dislocation of the carapaco-plastral articulation. In such instances the ligamentous tissue joining the two parts is torn extensively. Under these circumstances the movable shell parts seem to act as a safety device, giving way underpressurethat would crack the shell of a turtle with rigid, fixed buttresses. Dislocations of the carapaco-plastral articulation that have healed are characterized by abnormally heavy development of ligamentous tissue, which may elaborate a horny, scutelike substance on its outer surface.The extent to which serious injury incapacitates a turtle is not known. Surely open wounds are susceptible to infection and tovarious kinds of secondary injury; normal activity is probably interrupted by a period of quiescence, at least in the period of initial healing.An injured female had a hole, slightly more than one inch in diameter, in the right side of the carapace at the level of the second lateral lamina. A tight, thin membrane stretched between the broken edges of the opening; this membrane contained no bone and was covered externally by scar tissue. It was obvious that this turtle had recovered, at least in part, from a serious injury (inflicted probably by a piece of heavy farm machinery).Minor chips, scratches, and abrasions on the shell result from a variety of sources, some of them mentioned above. Small rounded pits in the bony shell (shell pitting) due to causes other than mechanical injury, are found in nearly all kinds of turtles according to Carpenter (1956), Hunt (1957), and my personal observation. InT. ornata, however, the condition is less common than in the specimens ofT. carolinadescribed by Carpenter and in the remaining species ofTerrapenethat I have examined.Carpenter (1956:86) came to no conclusion as to the cause of shell pitting inTerrapene carolinabut suggested that a variety of factors including parasitic fungi, parasitic invertebrates, and simple shell erosion, might be responsible.According to my own observations on turtles in the University of Kansas collections, shell pits range in size and shape from shallow, barely discernible depressions to deep borings; I suspect that shell pitting for turtles in general has many causes, some of which may be of more frequent occurrence in one species than in another.Hunt (1957:20) presumably was referring to shell pitting by a more suitable name when he wrote of, "… necrosis … of mycotic origin." Hunt (loc. cit.) stated that "Of those cases which have been recently examined, the author found all were due to the invasion of Mucorales beneath the plates of the epidermal laminae. This disease is of extremely common occurrence and has been found in all members of the order but is seldom found in marine species. Mycosis more frequently occurs on the plastron than on the carapace." Hunt presented no evidence to support his statement regarding invasion of the shell by Mucorales.Evidence that injury to the soft parts of the body is also fairly common is seen in the manyT. ornatawith missing feet and legs. Stumps resulting from amputations are covered with tough, calloused skin and sometimes by horny tissue similar to that of theantebrachial scales. Amputees are incapacitated only slightly in normal locomotion if a functional stump remains; probably a cripple is somewhat handicapped in other functions, such as burrowing, nest digging (females), and copulation (males). Causes of amputation are discussed in the section on predators.Fractures of the limb bones are common. A female from Stafford County, Kansas (Pl. 29, Fig. 4), showed a typical case of fracture and subsequent repair; the right fibula had been broken and the ends dislocated; a great mass of bone joined the repaired break to the middle of the tibia, giving the entire skeleton of the leg the appearance of the letter "H." The fibula, shortened by the dislocation, no longer articulated by its proximal end with the femur; the tibia probably bore the entire load in the period of repair and the transverse connection that formed between the bones later took over the function of the fibula.There is little doubt that ornate box turtles are stepped on or trampled by cattle, at least occasionally, but I never observed such an incident; the predilection of ornate turtles for dung insects and for moving along cattle pathways brings them to close quarters with cattle and probably did likewise with native ungulates. A steer, stepping on a box turtle, could inflict superficial damage to the shell or cause broken limbs but would probably not crush the turtle unless on a hard substrate.REPAIR OF INJURIES TO THE SHELLMost adults and a few juveniles examined in the field and laboratory had one or more small injuries on the carapace that had healed or were undergoing repair. Such injuries almost never occurred on the plastron. In an injury that was undergoing repair, a small piece of smooth, whitened bone was exposed where a piece of epidermis was missing from the shell. One or more edges of the exposed bone characteristically projected over the surrounding epidermis, making the bone appear as though it had been driven forcefully, like a splinter, into the shell (Pl. 29, Figs. 1 and 2). Because of their curious appearance, small areas of repair were referred to in my notes as "splinter scars." The position and number of splinter scars were often recorded as supplementary means of individualizing turtles in the field.Splinter scars result from minor abrasions that damage a few square millimeters of the shell. Larger areas of exposed bone were noted in only a few specimens. Two turtles at the Damm Farm hadbone exposed on more than one-half the surface area of the carapace; both of these turtles were probably burned in the grass fire of 1955. Ordinarily, a break in the shell does not induce extensive regeneration of tissues; when shells are damaged by crushing or cracking, regeneration of epidermis and bone occurs only along the lines of fracture, unless the broken parts have been dislocated. Ligamentous tissue develops in some breaks on the plastron, the broken area remaining slightly movable after healing is completed (Pl. 24).Dissection of injured shells revealed the mode of shell regeneration to be the same whether a large or small portion of the shell had been damaged. An abrasion may gouge out a small portion of the shell; burning, freezing, or concussion may kill a portion of the epidermis and a corresponding portion of bone beneath it without actually disfiguring the shell. Dead bone and epidermis become loosened at the margin of the wound. The epidermis sloughs off soon afterward but the bone adheres to the wound. New epidermis and new bone, growing from undamaged tissues at the edges of the wound, encroach on the wound beneath the layer of dead bone. The piece of dead bone is thereby gradually isolated from the rest of the shell and is sloughed off when healing is complete. The dead bone may come off in one piece or slough off gradually at its edges as healing proceeds toward the center of the wound. The layer of dead bone protects the wound during the process of regeneration (Pl. 30). Areas of exposed bone become white and shiny, nearly enamellike in appearance, as a result of wear on the shell.The above conclusions, in regard toT. ornata, agree basically with the findings of Woodbury and Hardy (1948:161-162) and Miller (1955:116) on regeneration of the shell in desert tortoises (Gopherus agassizi). Danini (1946:592-4, English summary) made histological studies on regeneration of the shell in specimens ofEmys orbicularis; he found that new bone trabeculae formed on the surfaces of undamaged trabeculae at the edge of the wound and formed also in connective tissue at the center of the wound. Regeneration of bone was incomplete in some instances where total extirpation of a portion of the shell had occurred. Regenerated epidermis was usually thicker than the original scute.Exposed bone on the shells of turtles that have been injured in fires, although dead, is unmarked and shows no evidence of being burned. Exposure to fire kills the growing portions of both the epidermis and the bone but seemingly does not actually char or disfigurethe bone (although the epidermis may be so affected) (Pl. 29, Fig. 3). Injuries from fire result probably from brief encounter with the fire itself or from more prolonged contact with some surface heated by the fire. A turtle that remained in a fire long enough to have its shell charred would presumably have little chance of survival. Grossly disfigured shells therefore do not result directly from burns but are due to the gnarled texture of the regenerated bone and epidermis remaining after the dead portions of the shell have been sloughed off. Information on injuries from fire was supplemented by examination of several badly burned specimens ofT. carolina. Their shells were nearly covered with exposed bone and regenerated epidermis. One specimen was so badly damaged that the entire anterior rim of its carapace was loose and could be pulled away easily to disclose a gnarled mass of regenerating bone beneath it (Pl. 29, Fig. 3). There were areas near the posterior margin of the carapace of each specimen where regenerated epidermis was evident but where the bone was seemingly uninjured; the regenerated epidermis was nearly transparent.Areas of regenerated epidermis on specimens ofT. ornatawere rough in texture and slightly paler than the surrounding scutes. Color-pattern is not reproduced in the process of regeneration but irregularly shaped light blotches sometimes occur in the places where radiations or other distinct markings formerly were present. A slight depression remains on the shell after regeneration is completed. I suspect that small injuries may be repaired in the course of a single growing season but that injuries involving a large part of the shell may take several years to heal completely. Cagle (1945:45) reported that a bullet wound in the shell of a painted turtle (Chrysemys picta) healed completely in approximately 23 months. Danini (loc. cit.) found that regeneration of the shell inEmys orbiculariswas complete in as short a time as 225 days. Woodbury and Hardy (loc. cit.) stated that small injuries to the shell ofGopherus agassizimay take as long as seven years to heal.ECTOPARASITESTwo kinds of ectoparasites were found on ornate box turtles in the course of the present study; larvae of chigger mites (Trombicula alfreddugesi) were abundant on specimens collected in summer and, larvae of the bot fly (Sarcophaga cistudinis) were found on specimens throughout the season of activity, and, in a few instances, on hibernating turtles. In general, these ectoparasitesdo little or no harm to ornate box turtles, although heavy infestations may cause temporary interruption of normal activity or may even cause occasional death.Concerning the larvae ofT. alfreddugesi, Loomis (1956:1260) wrote, "In northeastern Kansas, larvae become numerous in early June (shortly after they first appear), increase in numbers to greatest abundance throughout late June and July, decrease slightly in August, become markedly reduced in September, and only a few larvae (mostly on hosts) remain in October and early November." He consideredT. alfreddugesito be the most abundant chigger mite in Kansas and stated (op. cit.:1265) that it is most common "… in open fields supporting good stands of grasses, weeds and shrubs, and where moderate to large populations of vertebrates are present." Loomis listed ornate box turtles (op. cit.:1261-2) as important hosts ofTrombicula alfreddugesibut noted that box turtles are not so heavily infested as are certain other reptiles. The two other species of chigger mites that Loomis (op. cit.:1368) found onT. ornatain Kansas (T. lipovskyanaandT. montanensis) were not found in the present study.Box turtles were considered to have chigger infestations when the reddish larvae could be detected with the unaided eye. No chiggers were seen on turtles in the period from spring emergence until June 13, 1955. On the latter date a few scattered chiggers were noted on several individuals and it was on this same date that the writer received his first "chigger bites" of the year. Numbers of chiggers increased in the latter half of June and heavily infested turtles were noted throughout July. No chiggers were seen on box turtles after mid-September in 1955.Chiggers were ordinarily found only on the soft parts of the turtles' bodies. Early in the season infestations were chiefly on the head and neck. Favorite sites of attachment were the point where the skin of the neck joins the carapace and on the skin around the eyes. Later in the season some chiggers could be found on nearly every part of the body where soft skin was present; concealed areas of skin, such as the axillary and inguinal pockets, the anal region, and the inner rim of the carapace (where it joins the skin of the body), harbored concentrations of chiggers. Juveniles were relatively more heavily infested than adults and, even early in the season, had chiggers attached along many of the interlaminal seams of the shell. Broad areas of soft, newly-formed epidermis on the shells of juveniles probably afforded a betterplace of attachment to chiggers than did the interlaminal seams of adults. The interlaminal seams and transverse hinges of adults were not infested until the height of the season of chigger activity. Heavily infested adults, observed in early July, were literally covered with chiggers; red larvae outlined nearly all the scutes of the shell, the anus, the mouth, and the eyes. When turtles were picked up for examination, chiggers could be seen moving rapidly from one interlaminal seam to another.Box turtles kept in outdoor pens and in the laboratory did not long maintain visible infestations of chiggers, even during the time in summer when turtles found in the field were heavily infested.A four-year-old juvenile was found nearly immersed in the shallow water of a pond on July 4, 1955; its right eye had been damaged by an especially heavy concentration of chiggers. When I released the turtle, some 50 feet from the pond, it returned to the water and spent the next four days there. The turtle was probably in a period of quiescence induced by the eye injury and the heavy infestation of chiggers; immersion in water could be expected to help free the turtle of chiggers and to relieve trauma resulting from the injured eye. Richard B. Loomis told me that larval chiggers are able to survive under water for several days but that warm water will hasten their demise.Infestations of larval bot flies (Sarcophaga cistudinis) were noted in several turtles at the Damm Farm and, upon closer scrutiny, were found to be common in preserved specimens from other areas. Larvae were always found in flask-shaped pockets (Pl. 27, Fig. 2) beneath the skin; the pockets opened to the outside by a small hole, the edges of which were dried and discolored. One larva sometimes protruded from the opening. The inside of the pocket is lined with smooth, skinlike tissue. Heavily infested box turtles may have four or five such pockets, each containing one to many larvae. The most frequent sites of the pockets are the skin of the axillary and inguinal regions, and the skin of the limbs and neck, especially near the bases of these members. Subadults were more heavily infested than older adults; no infestations of hatchlings or small juveniles were noted.An adult female, infested with bot fly larvae when she was removed from her hibernaculum in late October, 1955, bore no trace of larvae or of the pocket that had contained them when she was recaptured the following June. According to Rokosky (1948), the larvae eventually fall to earth and pupate. The individuals ofT.carolinastudied by him were not re-infested by adult bot flies; one turtle ate some of the larvae that dropped from its body.The manner in which box turtles are infested by bot fly larvae is uncertain. Possibly the eggs are picked up accidentally or laid on the skin while box turtles are foraging in dung. Belding (1952:841) classifies the genusScarophagaas semi-host-specific, depositing eggs in open wounds.McMullen (1940), Rodeck (1949), and Rainey (1953), described individuals ofT. ornataparasitized byS. cistudinis. Rokosky (1948) and Peters (1948:473) reported infestations inT. carolina. Infestations were the cause of death in the instances noted by Rainey and Rokosky.PREDATORSFew first-hand observations on predators ofT. ornataare available and I have found little direct evidence of predation in the course of this study. In general, adults of the species seem to have few natural enemies other than man. Several of my colleagues at the University of Kansas have observed dogs carrying box turtles in their mouths or chewing on them. Frank B. Cross told me his dog caught and ate youngT. ornatain Payne County, Oklahoma, and A. B. Leonard once saw a badger carrying one in Dewy County, Oklahoma. At the Reservation, a freshly killed juvenile was found beneath the nest of a crow (Corvus brachyrhynchos) and remains of a hatchling were found in a scat of a copperhead (Agkistrodon contortrix).Dr. Fred H. Dale, Director of the Patuxent Research Refuge, Laurel, Maryland, kindly furnished photostatic copies of cards, from the Division of Food Habits Research of the U. S. Fish and Wildlife Service, recording the instances in whichTerrapene ornatawas listed as a food-item. In one instance the stomach of each of two nestlings, in the same nest, of the White-necked Raven (Corvus cryptoleucus) in Terry County, Texas, contained remains of recently hatched ornate box turtles; the remains of one turtle made up 64 per cent of the contents of one stomach, and parts of three turtles made up 80 per cent of the contents of the other stomach. Each of two stomachs of the coyote (Canis latrans) from Quay County, New Mexico, contained a "trace" of ornate box turtle.Wild carnivores known to occur on the Damm Farm were raccoons (Procyon lotor), striped skunks (Mephitis mephitis), badgers (Taxidea taxus), and coyotes (Canis latrans); all were suspect as predators of ornate box turtles.On December 10, 1953, ten dead box turtles (eight adults and two juveniles) were discovered at the top of a cut bank on the Damm Farm, within a few feet of a burrow that was used at least part of the time by a striped skunk. The condition of the turtles suggested that they had lain in the open for several weeks. The heads and legs were missing from most of the turtles and tooth marks were discernible on several of the shells. A logical explanation of this occurrence is that the turtles, using the burrow as a hibernaculum, were ousted by a predator that also inhabited the burrow. Turtles moving about sporadically in late autumn may be quickly chilled by a sudden drop in temperature and therefore be more susceptible to predation than at other times of the year. Two of my colleagues at the Museum of Natural History informed me that they had observed similar concentrations of deadT. ornatain winter.In July, 1952, H. B. Tordoff collected eight shells of juvenileT. ornatain a dry creek bed near Sharon, Barber County, Kansas. Some of the shells had small tooth-punctures. The stream bed habitat and the appearance of the tooth punctures tended to incriminate raccoons as predators. Raccoons, more than any other carnivore mentioned above, possess the manual dexterity necessary to pry open the shell of a box turtle and bite away the soft parts. Badgers and possibly coyotes are probably the only local carnivores (excluding large dogs) that could crack open the shell of an adult turtle by sheer force.Adults ofT. ornata, since they occasionally molest small juveniles, must be considered in the category of predators. When captive adults and juveniles were fed from the same container in the laboratory, the turtles occasionally bit one another accidently. Serious injury to the young was prevented by watching the adults closely and moving them away when they caught a smaller turtle by the leg or head. Similar accidents presumably occur in nature; juveniles and adults were sometimes found feeding side by side. William R. Brecheisen told me that adults kept in a stock tank at his farm in the summer of 1955 regularly and purposefully chased and bit small juveniles in the same tank. Brecheisen gave me a juvenile that had been so bitten; the right side of its head was badly damaged (the eye gone and a portion of the bony orbit broken) but was partly healed. Ralph J. Donahue told me that he saw an adultT. ornataattack a juvenalT. carolina, and provided a photograph of the incident. The juvenile was not injured.Although small box turtles may occasionally be caught and killed by adults in nature, this seems not to constitute a major source of predation on the young.Other animals that may prey upon young box turtles occasionally (and that were known to occur at the Damm Farm) are bullsnakes (Pituophis catenifer), red-tailed hawks (Buteo jamaicensis), marsh hawks (Circus cyaneus), crows (Corvus brachyrhynchos), and opossums (Didelphis marsupialis), and domestic cats.Nest predators probably have greater effect on populations ofT. ornatathan do predators of hatchlings, juveniles, and adults. Four robbed nests were found at the Damm Farm; in each instance, striped skunks were thought to be the predators. E. H. Taylor told me that he once saw a bullsnake swallow an entire clutch of newly laid eggs before the female turtle could cover the nest.DEFENSEBox turtles rely for protection on the closable shell and on inconspicuousness; defense reactions, except in the rare instances that biting is provoked, are purely passive.Box turtles handled in the course of field work varied widely in their reactions. Many struggled violently when being measured or marked whereas others were completely passive, closing the shell tightly and making it difficult for me to examine the soft parts of the body. These differences in behavior did not seem to be correlated either with sex or with age; generally lessened activity was associated with suboptimum body temperatures. All box turtles found in the field were extremely wary. As soon as one sighted me (sometimes at a distance of 200 feet or more), it became motionless with shell raised from the ground and neck extended (Pl. 28, Fig. 5). Some turtles remained in this motionless stance for half an hour or more, finally moving slowly away if I remained motionless. Turtles made no attempt to escape until I approached them closely or until they were in danger of being trampled by my horse; they would then move away with remarkable rapidity. Box turtles seemed unaware of an intruder until he could be seen or until he touched the turtle. When a turtle was approached from the rear, whistling, finger snapping, and normal footfalls did not attract its attention. Latham (1917:16) observed corresponding behavior inT. carolina. Wever and Vernon (1956) found the ear ofT. carolinato be keenly sensitive to sounds in the range of 100-600 cycles per second but progressively less sensitive to sounds of higher and lower frequencies. Surely a predator as stealthy as a coyote could approach a box turtle unseen and could quickly bite off at least one of the turtle's legs. Many of the mutilated box turtles that I observed may have survived such encounters with carnivores. The tendency of some individuals, when handled, to over-extend the limbs and neck (rather than closing the shell) in an attempt to escape, would make them easy victims for any predator.Ornate box turtles were kept in my home, along with several cats. Initial behavior was characterized by mutual wariness; subsequently the cats would follow a turtle about the house for a time, occasionally pawing at an exposed limb. The turtles withdrew only when touched or when approached from the front. After a day or two the cats and turtles ignored each other, often eating and drinking from the same dishes without incident. Under these circumstances the cats, I believe, could easily have killed or injured the turtles. A turtle would occasionally gain the respect of a cat by biting it.The strong odor sometimes given off by box turtles is produced by the secretions of four musk glands, two situated anteriorly on each side and opening by small, nearly invisible apertures beneath the fourth marginal scute. According to Hoffman (1890:9), two other musk glands, opening beneath the eighth marginal scute on each side, are also present inTerrapene; these posterior glands were not found in the several specimens ofT. ornatathat I dissected.Strong odors were produced by nearly all small juveniles until they became accustomed to being handled. Older juveniles and adults produced strong odors only in response to pain or injury, as, for example, when they were killed in the laboratory prior to preservation or when they were being marked in the field. Young box turtles were capable of producing strong odors as soon as they hatched.Norris and Zweifel (1950:3) considered the odor produced byT. o. luteolato issue from the "… concentrated, highly pungent urine…." voided by individuals when they were disturbed, and thought the production of odor to be a defense mechanism. Neill (1948b:130) reported that hatchlings ofT. carolinawith unhealed umbilical scars emitted a musky odor comparable to that of the stinkpot,Sternotherus odoratus; he thought the capacity to produce this odor was lost at about the time that the plastral hinge became functional.The function of musk glands inTerrapeneand, in all other turtles, is unknown. Since biting and nuzzling of the edges of the shell isan integral part of the courtship of many turtles, odor produced by the musk glands may well be a means of social recognition or of sexual stimulation. Repellant odor may have a protective value in young box turtles but it is unlikely that larger predators would be frightened away or even discouraged by odor alone. In this respect Neill (loc. cit.) and I concur.DISCUSSION OF ADAPTATIONSMost of the morphological characteristics distinguishing box turtles from other North American emyid turtles, the most notable of which is the movable plastron, are modifications that have evolved as a result of selectional pressures favoring adaptation to more or less terrestrial existence. Similar adaptations have arisen independently in several branches of the emyid stock (see introduction). The genusTerrapeneseems to have departed farther from a generalized emyid form than have other kinds of box-turtle-like chelonians. In a morphological sense,Terrapene ornatais clearly the most specialized member of its genus now occurring in the United States (my own studies have revealed that populations in western Mexico now referred to asT. klauberiandT. nelsoniare as specialized asT. ornatain some respects but more generalized in others). The present ecological study has demonstrated thatT. ornatais specialized in habits as well as in structure. It is concluded that these specializations (of more generalized and perhaps more primitive conditions as, for example inT. carolina) constitute adaptation for terrestrial existence in open, semiarid habitats. These adaptations inT. ornatahave resulted, in a few instances, in unique habits and structures; however, in most instances the adaptations have produced slight but recognizable changes that are definable only by degree of difference from other species of box turtles.The closable shell of box turtles is of obvious survival value in providing protection for the soft parts of the body. In most of the species ofTerrapene, the lobes of the plastron completely close the openings of the shell; closure is so tightly effected in some individuals that it is difficult to insert the blade of a knife between the adpressed margins of carapace and plastron. InT. ornatanevertheless, both lobes of the plastron are deficient on their lateral margins; four narrow openings remain when the lobes are drawn shut. Emargination of the plastron has occurred at the places where the limbs rub against it during locomotion. Thisreduction of the plastron permits the body to be held off the ground during forward locomotion and seemingly permits a generally freer range of movement for the limbs. The possible disadvantages of an imperfectly closable shell seem to be compensated for by increased mobility. Reduction of the plastron is correlated with a general lightening of the shell, probably associated with the increased vagility of this species. Lightening of the shell is evident also in the relatively thin, loosely articulated bony elements. Shells of adultT. ornatathat are old and weathered, or macerated (unless they are partly co-ossified because of injury), can nearly always be disarticulated with ease, whereas the bony elements in the shells of adultT. carolina(all races) are nearly always co-ossified or separable only after prolonged maceration.The relatively low, flattened shell ofT. ornatais an adaptation associated with the tendency to seek shelter in the limited space of earthen forms, burrows, or small natural cavities in the course of the warm season and to burrow more deeply into the ground in winter.Terrapene ornatais, in fact, the only species of the genus that may be considered an habitual burrower. Individuals ofT. carolinatend to seek shelter in the warm season by making forms in dense vegetation or by digging into yielding substrata such as mud or humus, although they may burrow deeply into the earth in winter. Extreme weakness or absence of the middorsal keel ofT. ornataseems to be a modification associated with burrowing habits and general adaptation to terrestrial life; the keel is similarly reduced in testudinids.Retention of epidermal laminae (as opposed to regular exfoliation of the older parts of scutes) occurs in all box turtles, in several other groups of terrestrial emyids, and in testudinids. The phenomenon is here considered to be a specialization of scute shedding—developed in terrestrial and semiterrestrial chelonians—that provides additional protection to the shell against wear and minor injuries.General shortening of digits—the result of reduction in number of phalanges as well as in their length, and to a lesser degree the shortening of metapodial elements—has occurred in several groups of chelonians with terrestrial tendencies (the opposite—lengthening of phalanges and metapodials, and hyperphalangy—has occurred in certain groups that are highly aquatic). The pes of box turtles has remained relatively unchanged in this respect; a fewphalanges on the lateral digit have been lost (especially in three-toed forms), but little reduction in length has occurred. The chief modification of the pes is a general narrowing brought about by the tendency of the digits to be crowded together, one on top of the other, rather than spread in a horizontal plane. Considerably more modification is seen in the manus ofTerrapene. Phalangeal formulae (expressing the number of phalanges from the first digit outward) range from 2-3-3-3-2 (primitive inTerrapene) to 2-3-3-2-2 in the races ofcarolinaand have the same range in the species of eastern Mexico. Extreme reduction in number (2-2-2-2-2) as well as general shortening of phalanges occurs inT. ornata. The formula is the same in the one specimen ofT. klauberithat has been skeletonized. This modification of the forelimb inT. ornatahas produced a more rigid, stronger manus that is well adapted to the requirements of burrowing and to locomotion over unyielding substrata. Shortening of the manus (and, to a lesser extent, the pes) has been accompanied by reduction and loss of interdigital webbing. It is noteworthy thatT. ornatahas achieved the same reduction in number of phalanges asGopherus, which displays the extreme of specialization in this respect among North American turtles. The manus inT. ornatais not shortened so much as inGopherus.The first toe in males ofT. ornatais uniquely widened, thickened, and inturned. Males of some other species ofTerrapenehave greatly enlarged rear claws, some of which turn slightly inward, but none has the flexed first toe hooklike as it is inornata(a modified first toe, resembling that described forT. ornata, has been observed in a live male ofT. klauberi[now KU 51430] since the preparation of this manuscript). In males ofT. ornatathe penultimate phalanx of the first toe has a normal, vertical articular surface on its proximal end. However, the distal articular surface (when viewed from the distal end of the phalanx) has its axis rotated away from the vertical plane approximately 45 degrees in a counterclockwise direction. As the foot is pronated and extended, and as the digits are flexed, there is a concomitant inward rotation of the first metatarsal at its proximal joint; this rotation, combined with the divergent planes of the articulating surfaces on the penultimate phalanx, cause the ungual phalanx to be flexed at right angles to the inner side of foot, in a plane perpendicular to that of the other toes (Fig. 21).The precise function of the modified first toe of males is unknown,although it is reasonably safe to assume that the modification is closely associated with clasping during coition. In the matings that I observed, the inturned first claw of the male secured a hold on the female's rump or just beneath her legs, whereas the remaining three toes gripped the edge of her plastron. The combined hold, on shell and skin, clearly affords the male a more secure position during coitus (whether the female clasps his legs with hers or not) than would a hold on skin or shell alone. Possibly intromission can be maintained in this position even when the female is attempting to escape. In males the plastron is less concave inT. ornatathan inT. carolina. Furthermore, males ofT. ornataare, on the average, smaller than females, whereas the reverse is true inT. carolina. Possibly the ability of the male to secure an especially firm grip on the female enhances the probability of small males mounting and inseminating larger females, whereas successful matings might otherwise be limited to pairs in which the male was the larger member.It is worthy of note that turtles of the genusTerrapeneare seemingly the only North American emyids that carry out the entire process of mating on land; other, semiterrestrial emyids (for example,Clemmys insculptaandEmydoidea blandingi) return to water for actual coition, although the precoital behavior sometimes occurs on land.Nearly all gradations from a fully developed zygomatic arch to a greatly reduced arch can be observed in skulls of the various species ofTerrapene(Fig. 2) (Taylor, 1895:586, Figs. 2-7). The highest degree of reduction is achieved inT. ornataandT. klauberi, both of which lack the quadratojugal bone and have no zygomatic arch whatever (except for an occasional, poorly defined anterior vestige formed by the postfrontal, the jugal, or both). Reduction of the zygoma clearly represents modification of a more generalized, complete arch. As yet there is no clear evidence that reduction of the zygomatic arch is of adaptive value. It is noteworthy, however, that similar reduction of the arch has occurred independently in a number of emyid and testudinid groups, nearly all of which have terrestrial or semiterrestrial habits. Although discussion of phyletic lines inTerrapeneis beyond the scope of this report, I tentatively suggest that reduced zygomatic arches have arisen independently in more than one group ofTerrapeneand that similar reduction of the arch in two species of the genus does not necessarily indicate an especially close relationship of such species.In a recent survey of cloacal bursae in chelonians, Smith and James (1958:88) reportedT. ornataandT. mexicanato be among the few emyids that lacked these structures; in the opinion of the authors (op. cit.:94) cloacal bursae evolved in chelonians that required an accessory respiratory organ for long periods of quiescence (hibernation or aestivation) under water, and were secondarily lost in terrestrial forms that hibernated on land. The assumption is a reasonable one, at least in regard to emyids and testudinids. Lack of cloacal bursae inT. ornataand in all testudinids, can be correlated with the completely terrestrial habits of those turtles. Cloacal bursae seem to be vestigial in the species ofTerrapenepossessing them and to be of little or no use as respiratory structures (except perhaps inT. coahuila).In most of the species ofTerrapenethe carapace has a pattern of pale markings on a darker background; however, unicolored individuals are the rule in certain populations (for example, at the western edge of the range ofT. carolinaand inT. ornata luteola) and occur as occasional variations in other populations (inT. yucatana,T. mexicana, and, throughout the range ofT. carolina, albeit more commonly in the southeastern part of the range). Personal observation of interspecific and ontogenetic variation of color patterns of box turtles has convinced me that a basic pattern of more or less linear radiations is the one from which all other patterns (including spots, blotches, rosettes, and the unicolored condition) can be derived, and, that the radial pattern is generalized and primitive forTerrapene(possibly for all emyids and testudinids as well). In the light of this conclusion, the radial pattern ofT. ornatamay be considered generalized. I suspect, however, that the pattern of a living species most closely approaching that of the primitive ancestral stock ofTerrapeneis the pattern of fine, wavy, dark radiations (on a paler background) present in young examples ofT. coahuila.Box turtles in general have lower reproductive potentials (as indicated by fewer eggs and longer prepuberal period) than do most aquatic emyids. This low potential seems to be compensated for by a lower rate of postnatal mortality (especially in adults) due to the protection afforded by the closable shell and the ability to recover from serious injury.Terrapene o. ornataandT. c. carolinaare the only box turtles the life histories of which are known well enough to permit significant comparison. The reproductive potentials ofT. o. ornataandT. c. carolinaseem to be much the same.
Fig. 28.The movements of an adult (non-gravid) female ofT. o. ornatain the house pond area at the Damm Farm during a period of 24 days in July, 1955 (solid line), and a period of three days (broken line) in July, 1956. Solid dots represent the points where the turtle was found as her thread trail was mapped; hollow symbols represent points of recapture when no trailing thread was attached to the turtle.
Fig. 28.The movements of an adult (non-gravid) female ofT. o. ornatain the house pond area at the Damm Farm during a period of 24 days in July, 1955 (solid line), and a period of three days (broken line) in July, 1956. Solid dots represent the points where the turtle was found as her thread trail was mapped; hollow symbols represent points of recapture when no trailing thread was attached to the turtle.
The actual home range of almost every individual studied, even of those individuals for which the most data were available, probably differed at least slightly from the observed or estimated home range. One adult female, for example, was captured six times intwo years within a radius of approximately 50 feet. Another female was found 2780 feet from her last point of capture. These last two records were regarded as unusual; when they were grouped with records of the 44 individuals mentioned above, the average radius of home range for the entire group was much larger (327 feet).
Fig. 29.The movements of a gravid female ofT. o. ornatain the southern ravine area at the Damm Farm in a period of ten days in June, 1956. Her movements were, for the most part, in and around several ravines (shown on map by broken lines) where she was searching for a nesting site. For explanation of symbols see legend forFig. 28.
Fig. 29.The movements of a gravid female ofT. o. ornatain the southern ravine area at the Damm Farm in a period of ten days in June, 1956. Her movements were, for the most part, in and around several ravines (shown on map by broken lines) where she was searching for a nesting site. For explanation of symbols see legend forFig. 28.
Homing Behavior
Gould (1957) reported that 22 of 43T. carolinamoved in a homeward direction when they were released in open fields up to 5.8 miles from their original points of capture. Turtles oriented themselves by the sun; homeward headings were inaccurate or lacking on overcast days and, light reflected from a mirror caused turtles to alter their courses. Seven of ten turtles released more than 150 miles from home headed in directions that corresponded most nearly to the headings last taken (at release-points near home base) and did not necessarily correspond to the direction of home. Gould's studies point out that box turtles perhaps practice a kind of "solar navigation." His work raises the question of whether the movements of box turtles are guided by the sighting of local landmarks or whether such landmarks alter the course of movement only when acting as barriers.
In the present study two experiments were made to determine the homing ability ofT. ornata. An adult male, taken from his normal home range in the house pond area and released 1200 feet away in the southern ravine area, traveled a generally northward course (not northeastward in the direction of home) for five days, moving a distance of approximately 1900 feet. His detached trailer was recovered several days later 740 feet southeast of the last known point in his travels (a distance that could have been covered in two days) and 150 feet from the point of original capture; he had returned to his home range by a circuitous route in a period of approximately seven days. Another adult male, captured in the southern ravine area, and released in the house pond area 1900 feet away, traveled on a course that bore approximately 25 degrees north of true homeward direction; after five days he was approximately 600 feet north of the original capture point. He then began a northeastward course that took him back to the house pond area where he remained for several days; no further data are available for this individual. It is significant that the homing males discussed above traveled greater average distances per day (based on records for nine days of trailing) than any of the other turtles studied (Fig. 27). Fitch (1958:101) released an individual one half mile from where he captured it and, one year later, recovered the turtle near the point of release.
Social Relationships
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. No instances of fighting were observed.
Allard (1935:336), Perm and Pottharst (1940:26), and Latham (1917) recorded instances of fights between individuals ofT. carolina; in the latter two instances fights were between males. Stickel (1950:362) observed an incident between two males that may have been a fight; however, she was of the opinion that fights rarely occur in nature and that box turtles do not defend territories. Evans (1954:23-25) considered the behavior ofT. carolinareported by Perm and Pottharst (loc. cit.) to represent "territoriality." He found "… a true hierarchy…." existing betweenfour captive males ofT. carolinaand another between three captive females of the same species; young individuals in the group raised their social level in the hierarchy after receiving experimental doses of male hormone. Evans (op. cit.:25) pointed out that true tortoises (family Testudinidae) have a more complex pattern of social behavior than do emyid turtles.
Observations made with binoculars from the vantage point of a blind provide the only information that I have concerning the reactions of box turtles to one another under natural conditions. Turtles foraging in a bare area were not startled by the approach of other turtles, and turtles moving across the area seemed to take no notice of turtles already there, regardless of whether these turtles were moving or not. Adults and subadults behaved in approximately the same manner.
Individuals traveling or foraging in rough terrain or in grassy areas probably are unable to see each other even when they are close to one another. Conversely, box turtles can see each other and are surely aware of each other's presence in bare, flat areas. These facts suggest that no social hierarchy exists inT. ornata. On one occasion an adult male and a juvenile (hatched the previous autumn) were found foraging next to one another on the same pile of cow dung.
When an individual became motionless in an attitude of wariness after having detected me in my blind, its behavior evoked no response on the part of other turtles, a few feet away.
INJURIES
Fire, freezing, molestation by predators, and trampling by cattle or native ungulates are only a few natural sources of injury to which box turtles have always been exposed. Man's civilization in the Great Plains, chiefly his automobile and other machines, have compounded the total of environmental hazards. Automobiles now constitute a major cause of death and serious injury to box turtles. Each year thousands are struck on Kansas highways alone, not to mention the many casualties resulting from mowing machines, combines, and other farm machinery.
Although grass fires usually occur in early spring or late fall when box turtles are underground, some turtles are surely killed by fires and many are injured. In early April of 1955 the pasture at the Damm Farm was burned. Similar burnings, I discovered, had occurred both intentionally and accidently in past years atirregular intervals. No deaths or injuries, attributable to fire were discovered in the course of intensive field work in the spring and summer of 1955, when the new grass was short and conditions for finding and marking box turtles were ideal. Badly burned individuals, if any, may have secreted themselves until their wounds had healed. In June, 1957, an adult female, that had been burned severely, was taken from a small puddle in a ravine on the Damm Farm. The soft parts of her body, excepting her head and neck, were a nearly solid mass of smooth scar tissue, the scales and rugosities of the skin being practically obliterated. The tail was reduced to a mere knob surrounding the anus and dead, exposed bone was visible on most of the dorsal part of the carapace. Possibly this female was burned in the fire of 1955. Lack of injury to the head and neck can probably be accounted for by the additional protection afforded these parts by the folded forelegs when the turtle was withdrawn in the shell.
Turtles that are smashed flat on the highway, of course, have no chance of survival. Highway fatalities are usually the result either of "direct hits," where the tire of a vehicle passes directly over the turtle, or of repeated pummeling by subsequently passing vehicles. The writer, while driving behind other cars that struck turtles or by sitting beside roads, has observed numerous turtle casualties. Most are struck a glancing blow by a tire and are propelled some distance through the air or on the surface of the pavement, often to the side of the road. Such a blow is usually sufficient to crack or chip the shell, or at least to scuff away parts of the epidermal covering. Turtles, so injured, usually survive.
Parts of the shell do not break away easily, even when several deep cracks are present, and only a little bleeding occurs. A common injury inflicted on the highway is the wrenching and subsequent dislocation of the carapaco-plastral articulation. In such instances the ligamentous tissue joining the two parts is torn extensively. Under these circumstances the movable shell parts seem to act as a safety device, giving way underpressurethat would crack the shell of a turtle with rigid, fixed buttresses. Dislocations of the carapaco-plastral articulation that have healed are characterized by abnormally heavy development of ligamentous tissue, which may elaborate a horny, scutelike substance on its outer surface.
The extent to which serious injury incapacitates a turtle is not known. Surely open wounds are susceptible to infection and tovarious kinds of secondary injury; normal activity is probably interrupted by a period of quiescence, at least in the period of initial healing.
An injured female had a hole, slightly more than one inch in diameter, in the right side of the carapace at the level of the second lateral lamina. A tight, thin membrane stretched between the broken edges of the opening; this membrane contained no bone and was covered externally by scar tissue. It was obvious that this turtle had recovered, at least in part, from a serious injury (inflicted probably by a piece of heavy farm machinery).
Minor chips, scratches, and abrasions on the shell result from a variety of sources, some of them mentioned above. Small rounded pits in the bony shell (shell pitting) due to causes other than mechanical injury, are found in nearly all kinds of turtles according to Carpenter (1956), Hunt (1957), and my personal observation. InT. ornata, however, the condition is less common than in the specimens ofT. carolinadescribed by Carpenter and in the remaining species ofTerrapenethat I have examined.
Carpenter (1956:86) came to no conclusion as to the cause of shell pitting inTerrapene carolinabut suggested that a variety of factors including parasitic fungi, parasitic invertebrates, and simple shell erosion, might be responsible.
According to my own observations on turtles in the University of Kansas collections, shell pits range in size and shape from shallow, barely discernible depressions to deep borings; I suspect that shell pitting for turtles in general has many causes, some of which may be of more frequent occurrence in one species than in another.
Hunt (1957:20) presumably was referring to shell pitting by a more suitable name when he wrote of, "… necrosis … of mycotic origin." Hunt (loc. cit.) stated that "Of those cases which have been recently examined, the author found all were due to the invasion of Mucorales beneath the plates of the epidermal laminae. This disease is of extremely common occurrence and has been found in all members of the order but is seldom found in marine species. Mycosis more frequently occurs on the plastron than on the carapace." Hunt presented no evidence to support his statement regarding invasion of the shell by Mucorales.
Evidence that injury to the soft parts of the body is also fairly common is seen in the manyT. ornatawith missing feet and legs. Stumps resulting from amputations are covered with tough, calloused skin and sometimes by horny tissue similar to that of theantebrachial scales. Amputees are incapacitated only slightly in normal locomotion if a functional stump remains; probably a cripple is somewhat handicapped in other functions, such as burrowing, nest digging (females), and copulation (males). Causes of amputation are discussed in the section on predators.
Fractures of the limb bones are common. A female from Stafford County, Kansas (Pl. 29, Fig. 4), showed a typical case of fracture and subsequent repair; the right fibula had been broken and the ends dislocated; a great mass of bone joined the repaired break to the middle of the tibia, giving the entire skeleton of the leg the appearance of the letter "H." The fibula, shortened by the dislocation, no longer articulated by its proximal end with the femur; the tibia probably bore the entire load in the period of repair and the transverse connection that formed between the bones later took over the function of the fibula.
There is little doubt that ornate box turtles are stepped on or trampled by cattle, at least occasionally, but I never observed such an incident; the predilection of ornate turtles for dung insects and for moving along cattle pathways brings them to close quarters with cattle and probably did likewise with native ungulates. A steer, stepping on a box turtle, could inflict superficial damage to the shell or cause broken limbs but would probably not crush the turtle unless on a hard substrate.
REPAIR OF INJURIES TO THE SHELL
Most adults and a few juveniles examined in the field and laboratory had one or more small injuries on the carapace that had healed or were undergoing repair. Such injuries almost never occurred on the plastron. In an injury that was undergoing repair, a small piece of smooth, whitened bone was exposed where a piece of epidermis was missing from the shell. One or more edges of the exposed bone characteristically projected over the surrounding epidermis, making the bone appear as though it had been driven forcefully, like a splinter, into the shell (Pl. 29, Figs. 1 and 2). Because of their curious appearance, small areas of repair were referred to in my notes as "splinter scars." The position and number of splinter scars were often recorded as supplementary means of individualizing turtles in the field.
Splinter scars result from minor abrasions that damage a few square millimeters of the shell. Larger areas of exposed bone were noted in only a few specimens. Two turtles at the Damm Farm hadbone exposed on more than one-half the surface area of the carapace; both of these turtles were probably burned in the grass fire of 1955. Ordinarily, a break in the shell does not induce extensive regeneration of tissues; when shells are damaged by crushing or cracking, regeneration of epidermis and bone occurs only along the lines of fracture, unless the broken parts have been dislocated. Ligamentous tissue develops in some breaks on the plastron, the broken area remaining slightly movable after healing is completed (Pl. 24).
Dissection of injured shells revealed the mode of shell regeneration to be the same whether a large or small portion of the shell had been damaged. An abrasion may gouge out a small portion of the shell; burning, freezing, or concussion may kill a portion of the epidermis and a corresponding portion of bone beneath it without actually disfiguring the shell. Dead bone and epidermis become loosened at the margin of the wound. The epidermis sloughs off soon afterward but the bone adheres to the wound. New epidermis and new bone, growing from undamaged tissues at the edges of the wound, encroach on the wound beneath the layer of dead bone. The piece of dead bone is thereby gradually isolated from the rest of the shell and is sloughed off when healing is complete. The dead bone may come off in one piece or slough off gradually at its edges as healing proceeds toward the center of the wound. The layer of dead bone protects the wound during the process of regeneration (Pl. 30). Areas of exposed bone become white and shiny, nearly enamellike in appearance, as a result of wear on the shell.
The above conclusions, in regard toT. ornata, agree basically with the findings of Woodbury and Hardy (1948:161-162) and Miller (1955:116) on regeneration of the shell in desert tortoises (Gopherus agassizi). Danini (1946:592-4, English summary) made histological studies on regeneration of the shell in specimens ofEmys orbicularis; he found that new bone trabeculae formed on the surfaces of undamaged trabeculae at the edge of the wound and formed also in connective tissue at the center of the wound. Regeneration of bone was incomplete in some instances where total extirpation of a portion of the shell had occurred. Regenerated epidermis was usually thicker than the original scute.
Exposed bone on the shells of turtles that have been injured in fires, although dead, is unmarked and shows no evidence of being burned. Exposure to fire kills the growing portions of both the epidermis and the bone but seemingly does not actually char or disfigurethe bone (although the epidermis may be so affected) (Pl. 29, Fig. 3). Injuries from fire result probably from brief encounter with the fire itself or from more prolonged contact with some surface heated by the fire. A turtle that remained in a fire long enough to have its shell charred would presumably have little chance of survival. Grossly disfigured shells therefore do not result directly from burns but are due to the gnarled texture of the regenerated bone and epidermis remaining after the dead portions of the shell have been sloughed off. Information on injuries from fire was supplemented by examination of several badly burned specimens ofT. carolina. Their shells were nearly covered with exposed bone and regenerated epidermis. One specimen was so badly damaged that the entire anterior rim of its carapace was loose and could be pulled away easily to disclose a gnarled mass of regenerating bone beneath it (Pl. 29, Fig. 3). There were areas near the posterior margin of the carapace of each specimen where regenerated epidermis was evident but where the bone was seemingly uninjured; the regenerated epidermis was nearly transparent.
Areas of regenerated epidermis on specimens ofT. ornatawere rough in texture and slightly paler than the surrounding scutes. Color-pattern is not reproduced in the process of regeneration but irregularly shaped light blotches sometimes occur in the places where radiations or other distinct markings formerly were present. A slight depression remains on the shell after regeneration is completed. I suspect that small injuries may be repaired in the course of a single growing season but that injuries involving a large part of the shell may take several years to heal completely. Cagle (1945:45) reported that a bullet wound in the shell of a painted turtle (Chrysemys picta) healed completely in approximately 23 months. Danini (loc. cit.) found that regeneration of the shell inEmys orbiculariswas complete in as short a time as 225 days. Woodbury and Hardy (loc. cit.) stated that small injuries to the shell ofGopherus agassizimay take as long as seven years to heal.
ECTOPARASITES
Two kinds of ectoparasites were found on ornate box turtles in the course of the present study; larvae of chigger mites (Trombicula alfreddugesi) were abundant on specimens collected in summer and, larvae of the bot fly (Sarcophaga cistudinis) were found on specimens throughout the season of activity, and, in a few instances, on hibernating turtles. In general, these ectoparasitesdo little or no harm to ornate box turtles, although heavy infestations may cause temporary interruption of normal activity or may even cause occasional death.
Concerning the larvae ofT. alfreddugesi, Loomis (1956:1260) wrote, "In northeastern Kansas, larvae become numerous in early June (shortly after they first appear), increase in numbers to greatest abundance throughout late June and July, decrease slightly in August, become markedly reduced in September, and only a few larvae (mostly on hosts) remain in October and early November." He consideredT. alfreddugesito be the most abundant chigger mite in Kansas and stated (op. cit.:1265) that it is most common "… in open fields supporting good stands of grasses, weeds and shrubs, and where moderate to large populations of vertebrates are present." Loomis listed ornate box turtles (op. cit.:1261-2) as important hosts ofTrombicula alfreddugesibut noted that box turtles are not so heavily infested as are certain other reptiles. The two other species of chigger mites that Loomis (op. cit.:1368) found onT. ornatain Kansas (T. lipovskyanaandT. montanensis) were not found in the present study.
Box turtles were considered to have chigger infestations when the reddish larvae could be detected with the unaided eye. No chiggers were seen on turtles in the period from spring emergence until June 13, 1955. On the latter date a few scattered chiggers were noted on several individuals and it was on this same date that the writer received his first "chigger bites" of the year. Numbers of chiggers increased in the latter half of June and heavily infested turtles were noted throughout July. No chiggers were seen on box turtles after mid-September in 1955.
Chiggers were ordinarily found only on the soft parts of the turtles' bodies. Early in the season infestations were chiefly on the head and neck. Favorite sites of attachment were the point where the skin of the neck joins the carapace and on the skin around the eyes. Later in the season some chiggers could be found on nearly every part of the body where soft skin was present; concealed areas of skin, such as the axillary and inguinal pockets, the anal region, and the inner rim of the carapace (where it joins the skin of the body), harbored concentrations of chiggers. Juveniles were relatively more heavily infested than adults and, even early in the season, had chiggers attached along many of the interlaminal seams of the shell. Broad areas of soft, newly-formed epidermis on the shells of juveniles probably afforded a betterplace of attachment to chiggers than did the interlaminal seams of adults. The interlaminal seams and transverse hinges of adults were not infested until the height of the season of chigger activity. Heavily infested adults, observed in early July, were literally covered with chiggers; red larvae outlined nearly all the scutes of the shell, the anus, the mouth, and the eyes. When turtles were picked up for examination, chiggers could be seen moving rapidly from one interlaminal seam to another.
Box turtles kept in outdoor pens and in the laboratory did not long maintain visible infestations of chiggers, even during the time in summer when turtles found in the field were heavily infested.
A four-year-old juvenile was found nearly immersed in the shallow water of a pond on July 4, 1955; its right eye had been damaged by an especially heavy concentration of chiggers. When I released the turtle, some 50 feet from the pond, it returned to the water and spent the next four days there. The turtle was probably in a period of quiescence induced by the eye injury and the heavy infestation of chiggers; immersion in water could be expected to help free the turtle of chiggers and to relieve trauma resulting from the injured eye. Richard B. Loomis told me that larval chiggers are able to survive under water for several days but that warm water will hasten their demise.
Infestations of larval bot flies (Sarcophaga cistudinis) were noted in several turtles at the Damm Farm and, upon closer scrutiny, were found to be common in preserved specimens from other areas. Larvae were always found in flask-shaped pockets (Pl. 27, Fig. 2) beneath the skin; the pockets opened to the outside by a small hole, the edges of which were dried and discolored. One larva sometimes protruded from the opening. The inside of the pocket is lined with smooth, skinlike tissue. Heavily infested box turtles may have four or five such pockets, each containing one to many larvae. The most frequent sites of the pockets are the skin of the axillary and inguinal regions, and the skin of the limbs and neck, especially near the bases of these members. Subadults were more heavily infested than older adults; no infestations of hatchlings or small juveniles were noted.
An adult female, infested with bot fly larvae when she was removed from her hibernaculum in late October, 1955, bore no trace of larvae or of the pocket that had contained them when she was recaptured the following June. According to Rokosky (1948), the larvae eventually fall to earth and pupate. The individuals ofT.carolinastudied by him were not re-infested by adult bot flies; one turtle ate some of the larvae that dropped from its body.
The manner in which box turtles are infested by bot fly larvae is uncertain. Possibly the eggs are picked up accidentally or laid on the skin while box turtles are foraging in dung. Belding (1952:841) classifies the genusScarophagaas semi-host-specific, depositing eggs in open wounds.
McMullen (1940), Rodeck (1949), and Rainey (1953), described individuals ofT. ornataparasitized byS. cistudinis. Rokosky (1948) and Peters (1948:473) reported infestations inT. carolina. Infestations were the cause of death in the instances noted by Rainey and Rokosky.
PREDATORS
Few first-hand observations on predators ofT. ornataare available and I have found little direct evidence of predation in the course of this study. In general, adults of the species seem to have few natural enemies other than man. Several of my colleagues at the University of Kansas have observed dogs carrying box turtles in their mouths or chewing on them. Frank B. Cross told me his dog caught and ate youngT. ornatain Payne County, Oklahoma, and A. B. Leonard once saw a badger carrying one in Dewy County, Oklahoma. At the Reservation, a freshly killed juvenile was found beneath the nest of a crow (Corvus brachyrhynchos) and remains of a hatchling were found in a scat of a copperhead (Agkistrodon contortrix).
Dr. Fred H. Dale, Director of the Patuxent Research Refuge, Laurel, Maryland, kindly furnished photostatic copies of cards, from the Division of Food Habits Research of the U. S. Fish and Wildlife Service, recording the instances in whichTerrapene ornatawas listed as a food-item. In one instance the stomach of each of two nestlings, in the same nest, of the White-necked Raven (Corvus cryptoleucus) in Terry County, Texas, contained remains of recently hatched ornate box turtles; the remains of one turtle made up 64 per cent of the contents of one stomach, and parts of three turtles made up 80 per cent of the contents of the other stomach. Each of two stomachs of the coyote (Canis latrans) from Quay County, New Mexico, contained a "trace" of ornate box turtle.
Wild carnivores known to occur on the Damm Farm were raccoons (Procyon lotor), striped skunks (Mephitis mephitis), badgers (Taxidea taxus), and coyotes (Canis latrans); all were suspect as predators of ornate box turtles.
On December 10, 1953, ten dead box turtles (eight adults and two juveniles) were discovered at the top of a cut bank on the Damm Farm, within a few feet of a burrow that was used at least part of the time by a striped skunk. The condition of the turtles suggested that they had lain in the open for several weeks. The heads and legs were missing from most of the turtles and tooth marks were discernible on several of the shells. A logical explanation of this occurrence is that the turtles, using the burrow as a hibernaculum, were ousted by a predator that also inhabited the burrow. Turtles moving about sporadically in late autumn may be quickly chilled by a sudden drop in temperature and therefore be more susceptible to predation than at other times of the year. Two of my colleagues at the Museum of Natural History informed me that they had observed similar concentrations of deadT. ornatain winter.
In July, 1952, H. B. Tordoff collected eight shells of juvenileT. ornatain a dry creek bed near Sharon, Barber County, Kansas. Some of the shells had small tooth-punctures. The stream bed habitat and the appearance of the tooth punctures tended to incriminate raccoons as predators. Raccoons, more than any other carnivore mentioned above, possess the manual dexterity necessary to pry open the shell of a box turtle and bite away the soft parts. Badgers and possibly coyotes are probably the only local carnivores (excluding large dogs) that could crack open the shell of an adult turtle by sheer force.
Adults ofT. ornata, since they occasionally molest small juveniles, must be considered in the category of predators. When captive adults and juveniles were fed from the same container in the laboratory, the turtles occasionally bit one another accidently. Serious injury to the young was prevented by watching the adults closely and moving them away when they caught a smaller turtle by the leg or head. Similar accidents presumably occur in nature; juveniles and adults were sometimes found feeding side by side. William R. Brecheisen told me that adults kept in a stock tank at his farm in the summer of 1955 regularly and purposefully chased and bit small juveniles in the same tank. Brecheisen gave me a juvenile that had been so bitten; the right side of its head was badly damaged (the eye gone and a portion of the bony orbit broken) but was partly healed. Ralph J. Donahue told me that he saw an adultT. ornataattack a juvenalT. carolina, and provided a photograph of the incident. The juvenile was not injured.
Although small box turtles may occasionally be caught and killed by adults in nature, this seems not to constitute a major source of predation on the young.
Other animals that may prey upon young box turtles occasionally (and that were known to occur at the Damm Farm) are bullsnakes (Pituophis catenifer), red-tailed hawks (Buteo jamaicensis), marsh hawks (Circus cyaneus), crows (Corvus brachyrhynchos), and opossums (Didelphis marsupialis), and domestic cats.
Nest predators probably have greater effect on populations ofT. ornatathan do predators of hatchlings, juveniles, and adults. Four robbed nests were found at the Damm Farm; in each instance, striped skunks were thought to be the predators. E. H. Taylor told me that he once saw a bullsnake swallow an entire clutch of newly laid eggs before the female turtle could cover the nest.
DEFENSE
Box turtles rely for protection on the closable shell and on inconspicuousness; defense reactions, except in the rare instances that biting is provoked, are purely passive.
Box turtles handled in the course of field work varied widely in their reactions. Many struggled violently when being measured or marked whereas others were completely passive, closing the shell tightly and making it difficult for me to examine the soft parts of the body. These differences in behavior did not seem to be correlated either with sex or with age; generally lessened activity was associated with suboptimum body temperatures. All box turtles found in the field were extremely wary. As soon as one sighted me (sometimes at a distance of 200 feet or more), it became motionless with shell raised from the ground and neck extended (Pl. 28, Fig. 5). Some turtles remained in this motionless stance for half an hour or more, finally moving slowly away if I remained motionless. Turtles made no attempt to escape until I approached them closely or until they were in danger of being trampled by my horse; they would then move away with remarkable rapidity. Box turtles seemed unaware of an intruder until he could be seen or until he touched the turtle. When a turtle was approached from the rear, whistling, finger snapping, and normal footfalls did not attract its attention. Latham (1917:16) observed corresponding behavior inT. carolina. Wever and Vernon (1956) found the ear ofT. carolinato be keenly sensitive to sounds in the range of 100-600 cycles per second but progressively less sensitive to sounds of higher and lower frequencies. Surely a predator as stealthy as a coyote could approach a box turtle unseen and could quickly bite off at least one of the turtle's legs. Many of the mutilated box turtles that I observed may have survived such encounters with carnivores. The tendency of some individuals, when handled, to over-extend the limbs and neck (rather than closing the shell) in an attempt to escape, would make them easy victims for any predator.
Ornate box turtles were kept in my home, along with several cats. Initial behavior was characterized by mutual wariness; subsequently the cats would follow a turtle about the house for a time, occasionally pawing at an exposed limb. The turtles withdrew only when touched or when approached from the front. After a day or two the cats and turtles ignored each other, often eating and drinking from the same dishes without incident. Under these circumstances the cats, I believe, could easily have killed or injured the turtles. A turtle would occasionally gain the respect of a cat by biting it.
The strong odor sometimes given off by box turtles is produced by the secretions of four musk glands, two situated anteriorly on each side and opening by small, nearly invisible apertures beneath the fourth marginal scute. According to Hoffman (1890:9), two other musk glands, opening beneath the eighth marginal scute on each side, are also present inTerrapene; these posterior glands were not found in the several specimens ofT. ornatathat I dissected.
Strong odors were produced by nearly all small juveniles until they became accustomed to being handled. Older juveniles and adults produced strong odors only in response to pain or injury, as, for example, when they were killed in the laboratory prior to preservation or when they were being marked in the field. Young box turtles were capable of producing strong odors as soon as they hatched.
Norris and Zweifel (1950:3) considered the odor produced byT. o. luteolato issue from the "… concentrated, highly pungent urine…." voided by individuals when they were disturbed, and thought the production of odor to be a defense mechanism. Neill (1948b:130) reported that hatchlings ofT. carolinawith unhealed umbilical scars emitted a musky odor comparable to that of the stinkpot,Sternotherus odoratus; he thought the capacity to produce this odor was lost at about the time that the plastral hinge became functional.
The function of musk glands inTerrapeneand, in all other turtles, is unknown. Since biting and nuzzling of the edges of the shell isan integral part of the courtship of many turtles, odor produced by the musk glands may well be a means of social recognition or of sexual stimulation. Repellant odor may have a protective value in young box turtles but it is unlikely that larger predators would be frightened away or even discouraged by odor alone. In this respect Neill (loc. cit.) and I concur.
DISCUSSION OF ADAPTATIONS
Most of the morphological characteristics distinguishing box turtles from other North American emyid turtles, the most notable of which is the movable plastron, are modifications that have evolved as a result of selectional pressures favoring adaptation to more or less terrestrial existence. Similar adaptations have arisen independently in several branches of the emyid stock (see introduction). The genusTerrapeneseems to have departed farther from a generalized emyid form than have other kinds of box-turtle-like chelonians. In a morphological sense,Terrapene ornatais clearly the most specialized member of its genus now occurring in the United States (my own studies have revealed that populations in western Mexico now referred to asT. klauberiandT. nelsoniare as specialized asT. ornatain some respects but more generalized in others). The present ecological study has demonstrated thatT. ornatais specialized in habits as well as in structure. It is concluded that these specializations (of more generalized and perhaps more primitive conditions as, for example inT. carolina) constitute adaptation for terrestrial existence in open, semiarid habitats. These adaptations inT. ornatahave resulted, in a few instances, in unique habits and structures; however, in most instances the adaptations have produced slight but recognizable changes that are definable only by degree of difference from other species of box turtles.
The closable shell of box turtles is of obvious survival value in providing protection for the soft parts of the body. In most of the species ofTerrapene, the lobes of the plastron completely close the openings of the shell; closure is so tightly effected in some individuals that it is difficult to insert the blade of a knife between the adpressed margins of carapace and plastron. InT. ornatanevertheless, both lobes of the plastron are deficient on their lateral margins; four narrow openings remain when the lobes are drawn shut. Emargination of the plastron has occurred at the places where the limbs rub against it during locomotion. Thisreduction of the plastron permits the body to be held off the ground during forward locomotion and seemingly permits a generally freer range of movement for the limbs. The possible disadvantages of an imperfectly closable shell seem to be compensated for by increased mobility. Reduction of the plastron is correlated with a general lightening of the shell, probably associated with the increased vagility of this species. Lightening of the shell is evident also in the relatively thin, loosely articulated bony elements. Shells of adultT. ornatathat are old and weathered, or macerated (unless they are partly co-ossified because of injury), can nearly always be disarticulated with ease, whereas the bony elements in the shells of adultT. carolina(all races) are nearly always co-ossified or separable only after prolonged maceration.
The relatively low, flattened shell ofT. ornatais an adaptation associated with the tendency to seek shelter in the limited space of earthen forms, burrows, or small natural cavities in the course of the warm season and to burrow more deeply into the ground in winter.Terrapene ornatais, in fact, the only species of the genus that may be considered an habitual burrower. Individuals ofT. carolinatend to seek shelter in the warm season by making forms in dense vegetation or by digging into yielding substrata such as mud or humus, although they may burrow deeply into the earth in winter. Extreme weakness or absence of the middorsal keel ofT. ornataseems to be a modification associated with burrowing habits and general adaptation to terrestrial life; the keel is similarly reduced in testudinids.
Retention of epidermal laminae (as opposed to regular exfoliation of the older parts of scutes) occurs in all box turtles, in several other groups of terrestrial emyids, and in testudinids. The phenomenon is here considered to be a specialization of scute shedding—developed in terrestrial and semiterrestrial chelonians—that provides additional protection to the shell against wear and minor injuries.
General shortening of digits—the result of reduction in number of phalanges as well as in their length, and to a lesser degree the shortening of metapodial elements—has occurred in several groups of chelonians with terrestrial tendencies (the opposite—lengthening of phalanges and metapodials, and hyperphalangy—has occurred in certain groups that are highly aquatic). The pes of box turtles has remained relatively unchanged in this respect; a fewphalanges on the lateral digit have been lost (especially in three-toed forms), but little reduction in length has occurred. The chief modification of the pes is a general narrowing brought about by the tendency of the digits to be crowded together, one on top of the other, rather than spread in a horizontal plane. Considerably more modification is seen in the manus ofTerrapene. Phalangeal formulae (expressing the number of phalanges from the first digit outward) range from 2-3-3-3-2 (primitive inTerrapene) to 2-3-3-2-2 in the races ofcarolinaand have the same range in the species of eastern Mexico. Extreme reduction in number (2-2-2-2-2) as well as general shortening of phalanges occurs inT. ornata. The formula is the same in the one specimen ofT. klauberithat has been skeletonized. This modification of the forelimb inT. ornatahas produced a more rigid, stronger manus that is well adapted to the requirements of burrowing and to locomotion over unyielding substrata. Shortening of the manus (and, to a lesser extent, the pes) has been accompanied by reduction and loss of interdigital webbing. It is noteworthy thatT. ornatahas achieved the same reduction in number of phalanges asGopherus, which displays the extreme of specialization in this respect among North American turtles. The manus inT. ornatais not shortened so much as inGopherus.
The first toe in males ofT. ornatais uniquely widened, thickened, and inturned. Males of some other species ofTerrapenehave greatly enlarged rear claws, some of which turn slightly inward, but none has the flexed first toe hooklike as it is inornata(a modified first toe, resembling that described forT. ornata, has been observed in a live male ofT. klauberi[now KU 51430] since the preparation of this manuscript). In males ofT. ornatathe penultimate phalanx of the first toe has a normal, vertical articular surface on its proximal end. However, the distal articular surface (when viewed from the distal end of the phalanx) has its axis rotated away from the vertical plane approximately 45 degrees in a counterclockwise direction. As the foot is pronated and extended, and as the digits are flexed, there is a concomitant inward rotation of the first metatarsal at its proximal joint; this rotation, combined with the divergent planes of the articulating surfaces on the penultimate phalanx, cause the ungual phalanx to be flexed at right angles to the inner side of foot, in a plane perpendicular to that of the other toes (Fig. 21).
The precise function of the modified first toe of males is unknown,although it is reasonably safe to assume that the modification is closely associated with clasping during coition. In the matings that I observed, the inturned first claw of the male secured a hold on the female's rump or just beneath her legs, whereas the remaining three toes gripped the edge of her plastron. The combined hold, on shell and skin, clearly affords the male a more secure position during coitus (whether the female clasps his legs with hers or not) than would a hold on skin or shell alone. Possibly intromission can be maintained in this position even when the female is attempting to escape. In males the plastron is less concave inT. ornatathan inT. carolina. Furthermore, males ofT. ornataare, on the average, smaller than females, whereas the reverse is true inT. carolina. Possibly the ability of the male to secure an especially firm grip on the female enhances the probability of small males mounting and inseminating larger females, whereas successful matings might otherwise be limited to pairs in which the male was the larger member.
It is worthy of note that turtles of the genusTerrapeneare seemingly the only North American emyids that carry out the entire process of mating on land; other, semiterrestrial emyids (for example,Clemmys insculptaandEmydoidea blandingi) return to water for actual coition, although the precoital behavior sometimes occurs on land.
Nearly all gradations from a fully developed zygomatic arch to a greatly reduced arch can be observed in skulls of the various species ofTerrapene(Fig. 2) (Taylor, 1895:586, Figs. 2-7). The highest degree of reduction is achieved inT. ornataandT. klauberi, both of which lack the quadratojugal bone and have no zygomatic arch whatever (except for an occasional, poorly defined anterior vestige formed by the postfrontal, the jugal, or both). Reduction of the zygoma clearly represents modification of a more generalized, complete arch. As yet there is no clear evidence that reduction of the zygomatic arch is of adaptive value. It is noteworthy, however, that similar reduction of the arch has occurred independently in a number of emyid and testudinid groups, nearly all of which have terrestrial or semiterrestrial habits. Although discussion of phyletic lines inTerrapeneis beyond the scope of this report, I tentatively suggest that reduced zygomatic arches have arisen independently in more than one group ofTerrapeneand that similar reduction of the arch in two species of the genus does not necessarily indicate an especially close relationship of such species.
In a recent survey of cloacal bursae in chelonians, Smith and James (1958:88) reportedT. ornataandT. mexicanato be among the few emyids that lacked these structures; in the opinion of the authors (op. cit.:94) cloacal bursae evolved in chelonians that required an accessory respiratory organ for long periods of quiescence (hibernation or aestivation) under water, and were secondarily lost in terrestrial forms that hibernated on land. The assumption is a reasonable one, at least in regard to emyids and testudinids. Lack of cloacal bursae inT. ornataand in all testudinids, can be correlated with the completely terrestrial habits of those turtles. Cloacal bursae seem to be vestigial in the species ofTerrapenepossessing them and to be of little or no use as respiratory structures (except perhaps inT. coahuila).
In most of the species ofTerrapenethe carapace has a pattern of pale markings on a darker background; however, unicolored individuals are the rule in certain populations (for example, at the western edge of the range ofT. carolinaand inT. ornata luteola) and occur as occasional variations in other populations (inT. yucatana,T. mexicana, and, throughout the range ofT. carolina, albeit more commonly in the southeastern part of the range). Personal observation of interspecific and ontogenetic variation of color patterns of box turtles has convinced me that a basic pattern of more or less linear radiations is the one from which all other patterns (including spots, blotches, rosettes, and the unicolored condition) can be derived, and, that the radial pattern is generalized and primitive forTerrapene(possibly for all emyids and testudinids as well). In the light of this conclusion, the radial pattern ofT. ornatamay be considered generalized. I suspect, however, that the pattern of a living species most closely approaching that of the primitive ancestral stock ofTerrapeneis the pattern of fine, wavy, dark radiations (on a paler background) present in young examples ofT. coahuila.
Box turtles in general have lower reproductive potentials (as indicated by fewer eggs and longer prepuberal period) than do most aquatic emyids. This low potential seems to be compensated for by a lower rate of postnatal mortality (especially in adults) due to the protection afforded by the closable shell and the ability to recover from serious injury.Terrapene o. ornataandT. c. carolinaare the only box turtles the life histories of which are known well enough to permit significant comparison. The reproductive potentials ofT. o. ornataandT. c. carolinaseem to be much the same.