  Fig. 1. Old adult male, year-old young and hatchling in July, showing differences in size and pattern. Fig. 2. Adult female skink in a natural nest, with her clutch of eggs late in incubation. The nest cavity is excavated in loose soil beneath a flat rock, which was raised momentarily to expose the nest to view. Fig. 3. The same female and nest, with eggs in process of hatching. Noble and Bradley (1933:77) mention frequent homosexual matings between captive males. However, I observed no homosexual matings, either under natural conditions or in confinement. The pugnacious behavior of males that are in breeding condition ordinarily would prevent homosexual mating. Males in such weakened condition as to be unable to defend themselves effectively might evoke sexual attack, instead of the usual fighting response in other males. Although no actual experiments were performed in the present study in connection with the courtship and mating behavior, accounts of some workers seem misleading. My own observations indicate that the capacity for sex discrimination in this particular kind of lizard, and probably in others, has been underrated. For example, it has been stated that the male rushes with open mouth at the neck of any other skink that happens to be around, and he identifies it as a male if it fights back, or as a female if it does not. On the contrary my observations indicate that sex recognition occurs almost as soon as the male is aware of another skink’s presence. The red head of the breeding male is an excellent example of a social releaser in the sense that this term was used by Tinbergen (1948:8). Like the red belly of the breeding The male whose dormancy was terminated in early winter by bringing him into a warm room causing him to assume breeding coloration and to breed some four months earlier than those under natural conditions has already been mentioned. By the time the regular breeding season arrived, this male had long since undergone sexual regression and retained no trace of the red suffusion. In this condition, placed in a terrarium with a mixed group of breeding adults, his social status was of unusual interest. He exhibited no interest in the females and was less pugnacious toward other males than were the individuals in breeding condition. Although he seemed somewhat more nervous and timid, his hostile behavior was not entirely suppressed, as from time to time he moved up to other males and bit them viciously. His color pattern resembled those of certain old adult females in which the body stripes have been suppressed, but the breeding males evidenced no uncertainty as to his sex and were uniformly hostile. Their reactions were not noticeably different toward him than they were toward breeding males. The importance of an olfactory stimulus as a social releaser in sexual behavior of lizards has not been appreciated, although Noble and Mason (1933:10) did demonstrate its importance in the behavior of the female toward her eggs. It is evident from published accounts, and from my own limited experience with fasciatus in parts of its range other than northeastern Kansas, that the phenology of the breeding cycle is subject to geographic variation, synchronizing with the somewhat different climatic conditions under which the species occurs. However, the difference is less than might be expected, in view of the species’ extensive range. As a result of the early spring, and the warm FightingTerritoriality in the usual sense is lacking in the five-lined skink, and could scarcely exist in an animal of its habits. To defend a definite area (territory) against intruders of its own species, the animal would have to detect such intruders promptly. The skink, however, is so secretive in habits that at any given time the individual is likely to be hiding and inactive, even when conditions are favorable for it to be in the open, and other individuals therefore can then wander onto its home range unopposed. Even when an individual is active, it lacks the ability to detect others, except within a radius which would encompass only a small fraction of Under most circumstances these skinks behave toward each other with tolerance or indifference, but during the breeding season adult males become hostile, and fight on sight. Their reddish facial suffusion serves as a social releaser which elicits hostile behavior and facilitates sex recognition. As the breeding season wanes, the reddish suffusion fades rapidly and male hostility, probably controlled by the same hormonal complex, is likewise suppressed. Hostile behavior is rare in adult females or young at any time. Combats and pursuits have been observed most frequently the last week of April and especially in the first two weeks of May. At this season funnel traps set along rock ledges often caught two adult male skinks together. In almost every instance one of the two confined males was mutilated, with pieces of skin and flesh bitten from the tail and with chin, snout, and neck scarred; most serious wounds were usually in the sacral region or base of the tail or both. Often the wounds were so severe that the skink died in a short time in captivity and presumably others that were released died also. On April 28, 1949, a large adult male skink, chased by another, ran out in the middle of a trail and stopped. The pursuer stopped a few inches from it, then after a long pause, retreated in the direction from which it had come. For the five minutes that the pursued skink was watched, it lay motionless, partly hidden by dry leaves, evidently seeking to avoid further pursuit by concealment. I caught it without difficulty, and it seemed weak and dazed, as if injured in the fight. Its reddish suffusion was conspicuous, but not fully developed. On May 3, 1949, an adult male having bright red facial suffusion was observed searching persistently in ground litter; he was seen to find and pursue a female, and to copulate. A few minutes after mating was completed and the pair separated, a second male also searching in the vicinity came within sight of the first one. The two noticed each other at a distance of about 18 inches, indicating their awareness by their more alert, jerky movements, and spasmodic vibrating of their tails. The newcomer darted at the other, and for a moment On May 10, 1949, two adult males were seen to approach each other slowly, pausing for perhaps a minute when they were a little more than one foot apart. Then one edged up to the other, and with a sudden lunge seized it by the head. The one seized broke away with a vigorous jerk, and promptly retaliated by biting the first one’s head. After a few seconds of rapid sparring and thrashing, they broke apart, and one chased the other for several feet until it eluded further pursuit by dodging and hiding. Fig. 9. Adult male skinks fighting. A. Menacing approach. B. One has lunged and secured a grip on the other’s side, holding it at right angle. The one caught is unable to flex its body and neck enough to secure a retaliatory grip on the attacker, and must break away by violent thrashing. On May 12, 1950, my attention was attracted by a rustling in dry leaves. Within a few inches of my foot two adult males were struggling fiercely with jaws interlocked. Sudden violent twisting and thrashing alternated with quiet periods of a few seconds duration, in which the lizards scarcely moved except for heavy panting and twitching of their tails. After perhaps two minutes of fighting, one broke away and ran. For a distance of several feet it was closely pursued by the other, which, however, soon lost contact with it in the rough terrain and surface litter. On May 12, 1951, rustling in dry leaves attracted my attention to two large adult males fighting. For about fifteen minutes that they were observed, they struggled, with neither yielding ground, though they thrashed and rolled about over an area of several square feet. Sometimes they were disengaged for short intervals. Then facing in opposite directions, with their heads side by side, they would snap at each other’s necks and shoulders (Figure 9). Part of the time both males had grips and were biting each other simultaneously, but more frequently one or the other had a temporary advantage. When one secured a grip it would strain to the utmost, biting as hard as it could and lunging forward with frequent short jerks, meanwhile striving to keep out of reach of the EggsThe eggs of Eumeces fasciatus are like diminutive chicken eggs in appearance. They are white when first laid, slightly translucent when held to the light. Within a day or two after they are laid, these eggs are soiled to a dull tan color, somewhat mottled, as a result of being rolled and dragged about in contact with the floor and wall of the nest burrow. Like the eggs of most other reptiles, those of Eumeces fasciatus have parchmentlike shells. These shells are thin and easily punctured. As incubation proceeds, the egg enlarges by gradual absorption of moisture and the somewhat elastic shell is stretched. An egg left in water for as much as a day does not gain in weight appreciably. Except for occasional abnormal ones, the eggs of any one clutch are notably uniform in size and shape at the time they are laid. As incubation proceeds, some eggs enlarge more rapidly than others, and attain larger ultimate size. Differences in shape also appear, some eggs becoming relatively elongate and thin, while others are thick and blunt. Some become distorted to asymmetrical shapes. In nests that have been deserted by the females, eggs of irregular shape are especially noticeable. It seems probable that the frequent shifting of the eggs by the female prevents unequal drying or stretching in different areas of the shell. Normal young were observed to hatch from grossly misshapen eggs. Under conditions of drought, the eggs may not enlarge normally during the latter part of incubation, and may become indented or partly collapsed, and yet apparently normal young
The extent of tolerance to immersion in water probably depends on the stage of development, the temperature, the oxygen content of the water and other factors. One egg was fully immersed for ten minutes on July 20, 1951, then returned to a container with damp soil in the laboratory, where it seemed to develop normally. On July 30 it was opened and found to have a living fetus, The range of temperature tolerance of the embryo is wide, probably comparable to that of the adult. Time required for incubation is dependent on temperature. Persistently wet and cloudy weather in the summer of 1951, keeping temperatures relatively low in nests, was a contributing cause to late hatching that summer. As compared with 1952, hatching was about one month delayed in 1951, but later emergence and breeding accounts for part of the difference. The extent to which low temperature may delay incubation was indicated by the effect of refrigeration on several experimental eggs, as recorded below.
These experiments seem to show that, in the later stages of incubation at least, lowering of temperature to 11° or 12°C. almost halts development of the fetus. Harm does not necessarily result, however, and when again warmed to normal incubation temperatures, the eggs eventually hatch, the incubation period being lengthened by a time approximately equivalent to the interval of refrigeration. Under natural conditions the time required for incubation probably varies within wide limits, controlled mainly by temperature. No two clutches receive the same amount of heat, as sites differ greatly in extent of insulation, and exposure to sunlight. Each year, earliest appearance of hatchlings is in a warm, sunny situation, and in cooler, well shaded places hatchlings appear somewhat later. Their incubation is evidently somewhat protracted, although later emergence from hibernation and later breeding of adults in these situations might also contribute to the delay. Widely different incubation periods have been recorded in the literature and the variation probably is not due to temperature alone. Noble and Mason (1933:4) recorded incubation periods for six females from the same locality, and evidently kept under the same laboratory conditions, as 47, 41, 36, 29, 29, and 27 days. Despite In the present study no incubation periods so short as those recorded by Noble and Mason, and Cagle, were observed. Incubation times were recorded for clutches both in the laboratory and in the field, but for most of the clutches only approximate incubation periods were recorded. Failure to record the exact date of laying or of hatching, or both resulted from attempts to avoid frequent disturbance of females in their nests, which might have caused them to desert. One clutch of eggs laid in a terrarium probably on June 17, 1951—possibly a day or two earlier—hatched on July 30, after an incubation of about 44 days. Another clutch, found in a terrarium on July 17, 1951, was estimated to have been laid about a week earlier, judging from the average length (11.8 mm.) and average weight (.55 gm.) of the eggs. These eggs hatched on August 9, a little more than three weeks after their discovery. A clutch found in the field on June 25, 1951, evidently recently laid (average length 12 mm., weight .45 gm.), hatched 41 days later, on August 5. Another clutch found in a terrarium on July 17, 1951, was estimated to have been laid ten days or two weeks before, as the average length was 12.7 mm. The eggs hatched on August 7, three weeks after their discovery. On June 25, 1951, an incomplete clutch of three eggs was found with a female which still had an unlaid egg. The three eggs probably had been laid the same day or the day before. They were kept in the laboratory and weighed and measured at intervals until July 28, 33 days after their discovery when both those that remained were accidentally punctured On June 21, 1951, a natural nest was found with eggs already somewhat enlarged (12.5 × 8 mm.) and mud stained. This nest was checked from time to time in the next few weeks, and after 39 days, on July 30, it was found that all the eggs had recently hatched, but six young were still in the nest cavity. Another nest was found on June 24, 1951, with the eggs already markedly enlarged (14 × 8 mm.) indicating that laying must have been several days earlier—probably well over a week. Hatching occurred approximately 34 days later, probably on July 28, since on July 26 there was no sign that hatching was imminent, and on July 30 only the empty dried eggshells remained in the nest. The incubation time approximated six weeks for those nests with most complete records. Under wet and stormy weather conditions such as prevailed in 1951, this may have been the normal incubation period, but in warmer and drier years incubation time is shortened. In the five-lined skink each adult female normally produces one clutch of eggs annually. The size of the clutch produced is subject to individual variation, and is influenced by the age, size and condition of the female. Geographic variation in clutch size might also be expected. Data were obtained from breeding females killed and dissected, from counts of eggs found in natural nests in the field, and from clutches of eggs laid by females kept in captivity. For the total of 115 recorded clutches represented by the combined data from all these sources, the average number of eggs per clutch was 9.5. In many females dissected for the purpose of obtaining egg counts, ovulation had not yet occurred. The ovarian eggs present in each of these females included two main size groups, the larger ones in process of maturing and evidently destined for deposition in the current season, and minute, immature ones. A few of intermediate size were always present, however, resulting in uncertainty as to the size of the clutch being produced, especially when development had not proceeded far. Even when the larger eggs formed a fairly distinct size group, some usually were well below maximum size. Relatively high counts of clutches were obtained from these examinations of enlarged ovarian eggs. Evidently development frequently is arrested, and resorption may occur before ovulation. As a result the numbers of ovarian eggs developing are a poor indication of actual clutch size. A series of gravid females were obtained For different sets of data on clutch size, numbers were as follows:
On the average, larger females produce more eggs per clutch than do smaller females. Of 49 females for which measurements were recorded, and which had uterine or large ovarian eggs, 31 were 70 mm. or more in snout-vent length. These 31, mostly or entirely old adults, averaged 9.9 eggs per clutch, whereas 18 others that were 69 mm. or less in snout-vent length, and that must have been mainly or entirely newly matured adults in their first breeding season, averaged only 7.8 eggs per clutch. Smith (1946:350) states that in the northern part of the range of this skink there is some indication of decrease in size of clutches. This is not well shown by published records. For the southern states, most of the published records of clutch size are by authors who did not clearly distinguish between the three kinds of five-lined skinks, and there is some doubt as to which species is involved in each record. For 56 clutches reported upon from north of approximately latitude 37°, I obtain a slightly higher figure than for 11 clutches from south of this line. Geographic trends are, of In Table 7, the figures marked with asterisks pertain to clutches that might have belonged to skinks of the species E. laticeps or E. inexpectatus since they were recorded in regions where laticeps and in some cases, inexpectatus also, occurs along with fasciatus. If these questionable clutches are excluded the remaining 55, definitely of fasciatus, average 8.48 eggs per clutch, whereas the 12 questionable clutches average 8.42. Both figures are close to the average of 8.82 ± .32 eggs for the 34 natural nests recorded in the present study. For the total of 1661 eggs of 182 clutches, from the combined sample of all available records for clutches found in the present study or reported upon in the literature, the average egg number is 9.13. Fig. 10. Correlation between size of female and number of eggs in clutch; females in their first breeding season, mostly less than 72 mm. in snout-vent length, produce smaller clutches, on the average, than do larger and older females, but there is extensive overlap. To sum up the available information on clutch size, the number of eggs is most typically 9, 10, or 11 and is more in large old females, than in small, newly matured females. In natural nests, even in those that are successful, there is often some loss of eggs, which are eaten by predators, or by the female herself, with the
BroodingLizards and snakes of several different families, are known to brood their clutches of eggs, although the great majority of oviparous forms do not do so. The brooding habit is perhaps best known in Eumeces fasciatus, and has been described by many authors. By far the most thorough account is that of Noble and Mason (1933) who observed and experimented upon seven females that laid clutches of eggs in captivity. These females, kept in separate terraria, excavated nest burrows for reception of their clutches, and remained with them throughout the time of incubation. There were three characteristic brooding postures; curved in a semicircle around the clutch, in an S-shaped figure extending among them, or lying straight, either over or among the eggs. The brooding females, taken quietly from their nests without disturbing them, were found to have temperatures averaging .4°C. higher than the nests. Evidently normal room temperatures were maintained in the laboratory where the terraria were kept. The females occasionally left their nests, especially in late afternoon, to wander about the terraria, and to bask in sunlight. While basking, their temperatures averaged 2.7°C. higher than the nest temperatures. The authors suggested that an important function of the brooding female was to transfer warmth from absorbed sunlight to the eggs. They state: “In nature the importance of the mother’s body heat in the incubation of the eggs probably varies greatly with the type of nesting site selected.” They suggest that in clutches deposited in logs or stumps beneath a thin layer of bark exposed to direct sunlight the need for warming by the female would be less. My own observations do not support the idea that brooding by the female serves to hasten the development of the eggs. Both in the laboratory and in natural nests, clutches deserted by disturbed females hatched and the hatching was not unduly delayed. In the field, females were never observed to bask in the sun beside their nest burrows, and seemingly left them infrequently even to feed. When a female was caught in her nest burrow, her temperature nearly always approximated that of the surrounding earth with which she was in contact. The temperature in each nest depends primarily upon its situation. When the immediate vicinity of the nest receives direct sunlight, the eggs are warmed without the aid of the female, but when there is no sunlight the temperature is much lower. In order to maintain an appreciably higher nest temperature the female would have to make frequent trips to spots perhaps several feet or several yards away to find sunlight. Upon Noble and Mason state (op. cit.:9) that while in some non-brooding kinds of lizards the eggs are actually damaged by turning, the female fasciatus frequently turns her eggs and moves the whole clutch about in the nest cavity. On returning to their nests the experimental females each invariably touched one or more eggs with their tongues as an olfactory test. Eggs of other kinds of lizards not of the genus Eumeces, and shellacked eggs of fasciatus, or paraffin models of them, ordinarily were discarded immediately after a single touch of the tongue. Eggs of other individuals of the species, and even the eggs of Eumeces laticeps were accepted as part of the brood. Any of the experimental females would quickly retrieve one of her eggs moved a short distance outside the nest cavity. Even if the whole clutch of eggs were scattered about, the female would, over a period of hours, gather the eggs and return them to the nest cavity. This movement of the eggs is accomplished by rolling or pushing them in a loop of the body or tail, or, less frequently, by grasping an egg in the jaws, lifting it, and gently placing it in a new position. Even if the females were blindfolded, they were still able to retrieve scattered eggs, but one in which the tongue tip was experimentally removed showed no further interest in its eggs, presumably having lost the capacity to recognize them by olfactory test. In the present study clutches unattended by females were observed to sustain heavy losses, both in the laboratory and in the field, and no doubt the attending female performs important functions other than that of warming the eggs. In the damp or wet nest cavity, the eggs tend to adhere to each other and to the earth walls and floor, and become sealed to such surfaces as a result of partial drying, reducing the amount of surface exposed to the air and probably hindering respiration. An eggshell sealed in prolonged contact with the soil tends to rot with the result that it is easily ruptured, and even if it is not broken there is the likelihood of fungi or microorganisms gaining entry and killing the embryo. In many of the eggs that were handled to obtain measurements and Another important function of the brooding female seems to be that of altering the nest burrow and shifting the eggs so that the effects of unfavorable weather are minimized. The usual response to warm and dry weather is deepening of the nest burrow. A cavity originally in loose soil on the underside of a flat rock, having the eggs in contact with the rock surface, may be displaced downward. The female excavates loose soil from the floor of the burrow and packs it on the top and sides, until the eggs are two or even three inches underground, in a cavity different in position and shape from the original one, although derived from it by gradual stages. In many instances, however, no such response to drying was observed. Probably extensive alteration of the nest burrow no longer is possible after drying of the soil has progressed beyond a certain stage as these skinks are not strong diggers. In some nests that were examined frequently, with resulting desertions by the attending females, the outlines of the cavities became indistinct and the soil around them became dry and packed. In heavy rains, when nest burrows are partly flooded, the females move the eggs to avoid their being submerged. The extent of the female’s activity within the nest burrow is suggested by the glazed condition of the earth walls and floor, and by the mottled appearance which the eggshells soon acquire as a result of being slid and dragged about in the nest cavity. Still another important function of the female is to dampen the nest burrow to prevent desiccation of the eggs. Even in dry weather, females taken from nests almost invariably voided water in relatively large quantities. They drink dew or other available water, and may void the contents of the bladder to moisten the nest cavity, as on numerous occasions, when nests were exposed by raising Noble and Mason (op. cit.:16-19) found that brooding females, in the laboratory, would vigorously defend their eggs against small enemies, including mice and lizards and the smaller kinds of snakes that were tested. The female watched alertly as the intruder approached, and attempted to bite it if it came too near or touched an egg. The females failed to defend their nests against persons and against a large blacksnake; when confronted with such a threat, the female would run from her nest cavity to hide. Cagle (1940:228) stated that the brooding females found by him stayed in the nests even when the logs in which they were situated were chopped open with an ax, and that the skinks would attempt to bite when touched with the finger. In the present study, females whose nests were exposed never made any active attempt to defend them. Many darted away and hid as soon as they were exposed. In other instances, especially when the nest cavity was only partly exposed, from one side, the female cowered back against the inner wall, opening her mouth in threat if closely approached. If further molested she might then attempt to escape. In brooding females a tendency to sluggishness, and an affinity for the eggs delayed the usually speedy escape reactions. The temperature of the female was ordinarily lower than it would have been in the open or on the underside of a flat rock, and this also tended to slow her reactions. Gravid females when exposed in nest cavities that still contain no eggs are similarly sluggish and reluctant to leave differing little or none in behavior from those that have laid their clutches. Usually the female was found with her body encircling the eggs, holding them together in a compact cluster in the center of the nest cavity. The eggs rest in contact with the loose soil on the floor of the cavity, with each other, and with the female’s body in the case of the outer ones of the cluster. Normal brooding habits proved to be difficult to follow because the females were easily disturbed. In many instances those that had excavated nest burrows, but had not yet laid, deserted the nests after the disturbance involved in raising the sheltering rock. Females that had already laid before discovery of their nests were somewhat less inclined to desert, but many did so. On numerous occasions, at the time of year when most females are gravid and are staying in nest burrows, I have discovered well formed nest burrows empty and seemingly deserted, with no female As compared with other North American lizards in general, Eumeces fasciatus is notable for the relatively exposed and superficial situations chosen as nesting sites. However, it occurs in a climate of high humidity; in contrast, the great majority of our lizards live in arid climates where the eggs are in much greater danger of desiccation, and require better shelter to maintain the humidity at a sufficiently high level. Accounts in the literature and observations in the present study indicate that these skinks exercise a wide range of choice of nesting sites. Ruthven (1911:264) stated that in northern Michigan nests were usually in decaying logs; occasional nests were found in burrows in sand, but invariably decaying wood was present in or around at least part of the nest. Blanchard (1922) mentions a nest in Tennessee that may have been made by either this species or E. laticeps “in a hollow in a dead willow tree about fifteen feet from the ground buried in the loose, damp, rotted wood.” Noble and Mason (op. cit.:16) quote Blanchard (in litt.) that in northern Michigan fasciatus nests in logs that are exposed to sunlight. Conant (1951:31) stated that several clutches of eggs found in Ohio were an inch to six inches beneath the upper surface of the log or stump which sheltered them. Evans and Roecker (1951:70) record finding two incubating females inside rotten pine logs, in Ontario. Cagle, studying this species near Elkville, Illinois, in oak-hickory woods, found 25 natural nests of In the present study, more than one hundred natural nests were found, of which just one (containing two clutches of eggs) was in decaying wood beneath the bark of an old log. All other nests were beneath rocks. On the University of Kansas Natural History Reservation, where most of the nests were found, the policy is not to tear apart decaying logs; therefore the nests probably present in such situations were not ordinarily found. On several occasions groups of hatchlings were seen on logs within which they probably had hatched. In the area of the study, however, decaying logs are scarce. The hardwood forests consist mostly of young trees that are second growth on cutover areas or pioneer on areas that were previously grassland. Because of frequent cutting there are few old mature trees, and logs have not accumulated on the forest floor. In northeastern Kansas, nesting in logs is comparatively rare. On wooded slopes and the edges of level hilltops, the flat limestone rocks that are often abundant provide preferred nesting sites. Even on collecting trips off the Reservation, where stumps and logs could be torn apart and searched, flat rocks were found to provide the main source of nesting sites. These nest rocks varied from less than an inch in thickness to nine inches or more, and from a few inches in diameter to three feet or more. Some were resting loosely on the surface of the soil and others were deeply sunken, on one side. Some were in situations exposing them to nearly the maximum amount of sunshine whereas others were in sites nearly always shaded. The varied character of the nesting sites chosen demonstrated a wide range of tolerance for temperature, moisture, and other factors, in the gravid and brooding female and in the developing embryo. As already mentioned, Noble and Mason (op. cit.:9-10) noted that females would accept and brood the eggs of other individuals just as readily as their own, and several writers have reported gregarious nesting habits, with two or more females occupying either the same nest cavity, or separate cavities that were in close proximity. For instance, Cagle wrote that among the small logs he found to contain nests, four logs each contained one nest, five each contained two nests, and two each contained three nests, while three other nests were found within an eight inch square area in loose The gregarious nesting habit may be of benefit in permitting maximum utilization of choice nesting sites, where such sites are in short supply in an environment otherwise favorable. Also, the gregarious tendencies make possible more continuous guarding of the eggs against such natural enemies as can be repulsed by the female, since each female occasionally interrupts her brooding to bask or forage. Many of the nests that I found were in close proximity to others. Often two nests, and sometimes even three, were found beneath the same rock, and sometimes a distance of only two or three inches intervened between the separate clutches. It seemed, however, that in almost every instance each female had excavated a separate nest chamber originally. In some instances adjacent nest chambers communicated with each other. On July 13, 1948, a communal nest was discovered beneath loose bark of a decaying elm log. There were 22 eggs in the combined clutch, and there were two females in the vicinity. The bark was raised on several different days to examine the eggs, and one or both females always were found with the eggs. On June 10, 1949, at the pond rock pile, a flat rock was turned and an unusual nesting aggregation consisting of a minimum of eight females, and probably more than ten, was found. The nests were somewhat disturbed by movement of the rock. The ground beneath was honeycombed with tunnels connecting the flask-shaped nest cavities, which were in part open to the rock surface on their upper sides. Clutches of eggs numbered 13, 12, 11, 8, and 6 (the last attended by a female which appeared to be still distended with several more unlaid eggs). Of five other females taken, two had laid and three were still gravid. Of the five clutches, two had eggs noticeably larger than those in the other three, and with their shells mottled brown from adhering earth. These nest cavities were about half an inch deep and two to three inches wide. The females were released as soon as they had been examined. One female moved about over the nest areas exposed, and evinced interest in a lone egg which had become separated from the others. She moved up to it, standing high off the ground, with her head turned at right angles to her body as if preparing to push the egg forward in On July 9, 1949, the flat rock covering the nests was raised again. Most of the eggs had hatched. Two broods of hatchlings were still in their respective nest cavities, and one entire clutch had not begun to hatch although its incubation was nearly completed. Three eggs of Scincella laterale were found mixed with the Eumeces eggs. One of these was opened to verify their identity; the other two hatched a few days later in the laboratory. The following selected excerpts from my field notes, setting forth histories of several nests, so far as they were known, give some idea of the types of nesting sites chosen, the behavior of the females, and the hazards to which the eggs are exposed. No. 1. At corner of pond rock pile. June 21, 1951. Female escaped when rock was turned. One egg measured 12.5 × 8 mm., mud-stained. June 22, 1951. Nest not in evidence when rock was turned; digging into loose soil beneath to a depth of about an inch I exposed the eggs but did not disturb them further. July 23, 1951. When rock was turned, female did not attempt to escape, but withdrew to far corner of nest cavity; when caught she voided a large scat which seemed to consist mainly of Ceuthophilus remains. Largest eggs in the clutch were 18 × 10 mm. but two were noticeably smaller, and all were heavily coated with dried mud. July 30, 1951. Six young in the nest cavity, still not fully active; all of them were heavily coated with dried mud. No. 2. At hilltop ledge, under flat rock 13 × 10 × 1 inches, with one edge sunken in soil; exposed to sunshine for most of day. June 24, 1951. Female, snout-vent length 70 mm., tail 27-51, weight 5 gms. Nine eggs, one of which measured 14 × 8 mm. July 18, 1951. Nine eggs still in their original nest cavity, attended by the female; she escaped into crevice behind the rock. The eggs were in slightly damp soil, and in contact with the undersurface of the rock on their upper sides; one egg was 17 × 10 mm. July 26, 1951. Eggs caked with dried mud; still attended by female. July 30, 1951. Dry and empty eggshells in nest cavity, evidently all the eggs had hatched; no other trace of female nor of young; July 28th seems most probable hatching date—if, on the 27th, some of eggs almost certainly would have shown signs of hatching on the 26th when they were examined, and if on the 29th some stragglers almost surely would have remained at the nest on July 30. No. 3. In small gully, on lower slope in hickory woods, beneath rock 9 × 9 × 1 inches, shaded by trees on south side for much of the day, especially during latter part of morning. June 24, 1951. The gravid female was deep in nest burrow. June 29, 1951. When rock was lifted no trace of nest was visible except for slightly disturbed loose soil at the point where it had been. When some of this loose soil was cleared away, nest was revealed, with 11 eggs, mud-stained, approximately 12.5 × 8 mm. The female was cold and sluggish, and did not attempt to escape, but cowered in the back of the nest burrow, with jaws gaping; she was caught and marked. July 20, 1951. Eight eggs remaining in the nest—two were accidentally destroyed in moving them. These two were fertile and contained live embryos, one of which measured 29 mm. in over-all length. One of the remaining eggs was 16.5 × 10 mm. Female was present with the eggs. July 25, 1951. Eggs still present in the nest cavity; female not in evidence, but might have been concealed in corner of nest chamber as it was not disturbed. July 28, 1951. Female was again found with the eggs. One or more of the seven remaining eggs were punctured in moving them during their examination. Eggs about 16 × 10 mm. August 3, 1951. Female was in nest with the eggs some of which are slightly indented from drying. August 6, 1951. When rock was turned, female darted out and ran to cover about ten feet away. The eggs had hatched but two young remained in the nest cavity, still rather slow and feeble in their movements and not yet fully active. When routed from cover a second time, the female ran back to the nest rock and took shelter beneath it. No. 4. On upper slope above ledge, under a rock 18 × 9 inches, in site shaded most of day; burrow nearly concealed beneath rock. June 24, 1951. Nest occupied by a gravid female, apparently ready to lay. June 30, 1951. Rock covering this nest has been undermined by a mole tunnel, and many nearby rocks are undermined also. The eggs were almost certainly destroyed by the mole’s tunneling and may have been eaten by it, since no remains are in evidence. No. 5. At hilltop ledge beside old abandoned road, beneath flat rock nine inches in diameter and about 11/2 inches thick, shaded for first half of morning and most of afternoon, but exposed to mid-day sunshine. June 29, 1951. Standing water in bottom of nest chamber 11/2 inches below underside of the rock. Some of the eggs are more than half submerged. One egg is 14 × 8 mm. July 21, 1951. Entrance of abandoned nest burrow has been enlarged by running water channelled through in run-off during and after heavy rains; shrivelled remains of eggs present at the bottom of the burrow. No. 6. On grassy hilltop a few yards from ledge under flat rock, 9 × 6 × 2 inches. July 23, 1951. Large female (snout-vent length 75 mm.) with three eggs, 16 × 22 mm. July 27, 1951. Female escaped from nest cavity as rock was raised. Three eggs were still in the nest, and a young skink was partly emerged from one. A second egg not yet hatching was somewhat flaccid, 16 mm. long, heavily coated with dried mud. The third egg much shrivelled, was opened and found to have a dead fetus, perhaps a week short of hatching. July 28, 1951. The flat rock which formerly covered the nest cavity was found to have been raised and displaced, and no trace of the female, eggs or young remained. Of possible predators that might have moved the rock and destroyed the nest, skunk and opossum seemed the most likely, but there was no definite clue as to the predator’s identity. No. 7. Two feet northeast of pond rock pile, under rock about one foot square on upper surface with maximum thickness of about eight inches, lying with upper side at 45-degree angle. The nest was under one edge, with approximately three inches of rock over it. The rock was exposed to sunshine throughout the day, except for grass shading its edges. July 23, 1951. When rock was turned, the female darted out of the nest cavity, but in her dash to escape she dropped into a nearby pitfall. When handled, she voided feces which contained the nearly intact shell of a skink egg. Six eggs present in the nest; one selected as typical was 111/2 × 8 mm. The eggs were slightly misshapen and might have been damaged from drying. July 26, 1951. When rock was raised, female darted out and escaped. The six eggs still remained in the nest. August 2, 1951. When rock was raised the female was not in evidence, and only three eggs could be found; they had fallen from the nest cavity to the bottom of the depression where the rock was imbedded and were somewhat dried and indented. No. 8. North slope, beneath rock approximately 18 × 15 × 4 inches, at edge of small gully, where shaded most of the time including mid-day hours. July 20, 1951. Female attempted to escape from the nest. Four eggs visible in nest, one 151/2 × 10 mm. July 25, 1951. When rock was raised the female ran from the nest. July 27, 1951. When rock was raised the female was in the nest with the eggs; she ran and hid beneath a boulder five feet away. After a few minutes she emerged and ran 15 feet to a hickory sapling and climbed it. July 28, 1951. Female was not in the nest but the four eggs were still present. July 30, 1951. Female found dead and partly eaten by ants beside rock one foot from nest; eggs still present in the nest. July 31, 1951. Eggs still present in the nest. August 3, 1951. Eggs still present, including some deep in the nest cavity which apparently were overlooked previously. August 6, 1951. One much indented egg found outside the nest cavity was opened and found to contain a live fetus, seemingly fully developed and normal. The opened egg was placed on damp soil in a shady place near the nest, but two hours later the hatchling had been killed and partly eaten by swarms of ants. August 9, 1951. The remaining eggs had disappeared, evidently taken by a predator as no empty shells remained to indicate that the young had hatched. HatchlingsCagle (1940:229 and 232) has graphically described and illustrated the hatching of the five-lined skink, and numerous observations in the present study have served to corroborate his description. The first indication that the time of hatching is at hand is a twitching Eggs ready to hatch ordinarily weigh somewhat more than one gram, up to at least as much as 1.7 grams, but much of this weight is made up of water absorbed during incubation. The hatchlings usually weigh from .2 to .45 grams. For each of two eggshells recently vacated, that were washed and squeezed dry, weights were approximately .125 grams. Hatchlings of the same brood differ perceptibly in size with several per cent variation in total length, and weight. Some seem to be less fully developed than others. On July 8, 1952, hatching of the last young in a clutch was observed. Upon emergence, it differed in appearance from the others of the brood hatched a few hours earlier. The top of its head bulged slightly as in fetuses. The umbilicus was not yet closed, and the protruding yolk mass hindered the hatchling’s movements and made crawling difficult. In order to progress it had to stand high off the ground to prevent its ventral surface from dragging. Protrusion of the yolk mass has been described in newly emerged hatchlings for the closely related E. anthracinus (Clausen, 1938:3-7) as well as in fasciatus. Cagle (loc. cit.) states that the mass of yolk is at first about 3 mm. in diameter, but is completely used at the end of the third day. A group of young retained by him, without food, died the sixth day after hatching, seemingly from starvation. Three of five recently hatched young were found by Cagle to have eaten ant pupae placed in a box with them on the preceding day, even though the skinks still retained the yolk masses. One hatchling of this group Contrary to the statement by Noble and Mason (1933:5) that in captivity the hatchlings seldom stayed together more than a few hours, litters of young fully active, a day or two after hatching were found in the nests with the females still looped around them on several occasions. On one such occasion, although the brood scattered immediately into surrounding vegetation where they hid, I succeeded in catching the female and six of the young, and put them all together in a nylon bag to carry them back to the laboratory. Several hours after the bag had been placed on a table it was noticed that the family had again gathered into a compact cluster in the bag with the female’s body looped around the young in the characteristic brooding position seen in those with young or eggs in their nest cavities. When hatching is complete, the female may leave before the young have dispersed. On August 5, 1950, a nest under observation was found to have all of the young or most of them still clustered in the cavity, but the female was not in evidence. The young were active, and immediately took alarm as the rock was raised exposing them. Almost instantly, they scattered and vanished. Subsequent search revealed five of the young, each poorly concealed in tufts of grass or under dry leaves or other ground litter at the edges of the depression where the rock had lain. Once hidden, these young were reluctant to run again and depended on concealment. Having once left the nest, the young probably do not return to it, as many nests examined within a few days after hatching were never found occupied either by females or young after their original dispersal. As soon as the dispersal occurs family ties are permanently severed. On July 19, 1950, a group of active hatchlings was observed moving about over a log, on what was probably the first day of activity away from the nest. The log was in the bottom of a steep-walled gully, where it had come to rest the night before. It had been an erect but dead and partly undermined snag on the edge of the gully, and was blown down that night in a violent thunderstorm. Most of the log was held clear of the rushing water in the bottom of the gully by projecting limbs. The little skinks Fig. 11. Sizes on specific dates of young hatched in 1950 and 1952. Approximate size ranges at different times of year, and differences in trend between the two years are brought out. The young were much more active than the female. These and other young observed in the open were almost constantly in motion. Pauses to bask at any one spot were of only a few seconds duration. A certain log in Skink Woods evidently was the site of one or more successful skink nests each year that observations were made, although a nest was actually found in it only in 1948. On July 26, 1950, recently hatched young were active on this log. Temperature was about 22°C. and the young were alternating frequently between shade and sunshine to maintain their body temperature. Collectively they seemed to cover every square inch of the log surface, poking and probing into niches, crevices and insect borings. They had a tendency to seek out the highest points on the log as resting places. In moving about, foraging or sunning, the young often carry the tail arched high, and keep it in motion with slow squirming undulations. These undulations may be continued even when the lizard itself has come to rest momentarily. The movements of the tail together with its vivid blue color serve to attract attention to it. Such behavior has not been observed in adults or partly grown young. Jopson (1938:90) observed an instance in which two dogs cornered a young five-lined skink (either the present species or E. laticeps) but were distracted by the wriggling of its bright blue tail “either dropped by autotomy or knocked off” so that the skink itself was allowed to escape. On another occasion these same two dogs attacking an adult male skink, were not distracted by the wriggling but dull colored broken tail, and they killed the lizard. GrowthThe subject of growth in Eumeces was briefly discussed by Taylor (1936:66) in his revision of the genus. Sorting fairly large series of museum specimens into seeming age-size groups, Taylor concluded that skinks require as much as 9 or 10 years to attain adult size. For fasciatus, for instance, the snout-vent length of 65.7 mm. (small adult size) was considered typical of individuals in their ninth year of life, with yearly gain of only 6 or 7 mm. in length in the young. I have seen the original data on which this conclusion was based, and the age groupings, as assigned by Taylor, seemed plausible. However, in the light of present knowledge, it is certain that the seeming intervals between his assumed age groups would have disappeared with a still larger series of specimens. The eight or nine size groups that Taylor recognized as distinct annual age groups actually comprise only two age groups, each having such Growth in reptiles is now much better understood. Many species have been studied by a variety of methods, including observation of growth in captives, recording of growth in marked individuals living under natural conditions, and sorting of large series into age-size groups. Two species of Eumeces have been studied in some detail. Breckenridge (1943:601-602) marked all the individuals of septentrionalis that could be found in a small colony in Minnesota and he concluded from the growth recorded in several that were recaptured, that these skinks grow to mature size (65 mm. and larger) at the end of their second year of life and are ready to breed the following spring. Rodgers and Memmler (1943:61) plotted the size distribution of a large year-round collection of skiltonianus from near Berkeley, California. They found that in this species hatching occurs in July and August, hatchlings are about 25 mm. in snout-vent length, and grow to about 50 mm. by the time they are one year old, and to about 65 mm. at two years of age, but most of them breed at the end of their third year. Within the genus the species septentrionalis and skiltonianus belong to groups separate from each other and from that including fasciatus. While septentrionalis and skiltonianus resemble each other in their growth pattern and in the time required to reach sexual maturity, fasciatus is notably different in its more rapid growth and the shorter time it requires to reach breeding maturity. This would scarcely be expected, as all three are of similar size. Furthermore, skiltonianus in the region of Rodgers’ and Memmler’s study has a longer growing season than fasciatus in northeastern Kansas, while septentrionalis in Minnesota has a growing season markedly shorter than either. It is noteworthy that each of these three skinks is the northernmost lizard in the section of the country where it occurs. In the present study growth was investigated by measuring and marking large numbers of young, many of which were recaptured for subsequent records, and by sorting into age-size groups all available measurements. An understanding of the latter set of data was facilitated by correlating it with the growth records of marked individuals. Changes in the phenology of growth from year to year according to weather conditions were noted. As already indicated, hatching occurs from early July to mid-August in northeastern Kansas. Unseasonably cool weather with frequent rains may cause cumulative delay in breeding and incubation so that hatching may average several weeks later than it does Newly hatched young average just under an inch in snout-vent length (23-27 mm.) and weigh .2 to .45 grams. Most rapid growth occurs in the period of weeks following hatching. The growth rate during this late summer period cannot be well shown by comparing average size of series taken on successive dates, because each series is likely to include some newly hatched young. In 1949, a series of recently hatched young averaged 26.7 mm. on July 10. By August 26, average length in a series collected was 42.9 mm., indicating an average gain of at least .35 mm. per day. One that may be considered typical was marked on July 23, 1950, soon after hatching, and it had a snout-vent length of 26.5 mm. and weighed .25 grams. It was recaptured just a month later when it had grown to 36 mm. snout-vent length, and weighed .8 grams. Potential growth rate under favorable conditions is shown by the fact that some individuals have attained a snout-vent length of 50 mm. by the third week of August, thus approximately doubling their hatching length. A maximum growth rate of about .5 mm. per day is indicated for these accelerated individuals, but on the average, young are considerably less than 50 mm. in length even when they enter hibernation. At the other extreme, representing retarded growth, is an individual having a snout-vent length of only 34 mm. on May 1. It must have been approximately nine months old on that date, but of course had spent at least six months in hibernation. Even if it made rapid growth subsequently, this yearling could scarcely have attained by midsummer the pre-hibernation length of the most accelerated individuals. During the growing season following their first hibernation period, the young grow to small adult size in most instances. After emerging from a second hibernation they mature sexually and constitute an important part of the breeding population. Many of the skinks marked before their first hibernation, as hatchlings, when they were a few days or a few weeks old, were subsequently recaptured as well-grown yearlings or small adults,
Many other young were not caught and marked until the growing season following their first hibernation, and were recaptured within this second growing season weeks or months after they were originally marked, and after they had made substantial growth. Those recaptured near the end of this second growing season, when they were a year old, or a little more, usually had attained small adult size or were nearing it. Selected records of these yearlings are presented below.
Adult skinks can be found in greatest numbers in the breeding season and many of the young that were marked were recaptured as newly matured breeding adults soon after their second hibernation, often still short of average adult size. Selected records of such individuals are presented below.
Fig. 12. Sizes of immature skinks of successive annual broods, grouped in biweekly or monthly intervals, with mean, standard error, standard deviation, and extremes shown for each group. A certain small percentage fail to attain minimum adult size or breeding maturity by the time of emergence from their second hibernation. Among 77 individuals marked as young either soon after hatching or in spring and early summer, and recaptured the following spring, only one had failed to grow to adult size. It was 46.5 mm. in length when marked on June 13. When recaptured on April 25 of the following year, it had grown to a length of 59 mm., still short of minimum adult length. During the interval between captures it had maintained about the average growth rate. Its failure to attain maturity was obviously the result of its early retardation, and probably late hatching was primarily responsible. Although this is the only individual with known history, which failed to attain breeding maturity after its second hibernation, occasional specimens are taken in spring which are somewhat below adult size but seem too large to be young hatched the preceding summer. Obviously, the incidence of such failure from year to year would be influenced by weather conditions, and an unusually cool summer may result in such delayed laying and hatching that an unusually large proportion of young might fail to attain sexual maturity at the usual time. At more northern localities, the percentage of such failures might be expected to increase. At the northern edge of the range attainment of breeding maturity may normally require more than two years. Such delayed development would result in a drastic reduction of the reproductive potential which might be critically limiting to the species, even in an otherwise favorable environment, as the population would be unable to replace rapidly enough the individuals eliminated by normal mortality factors. In contrast to the delayed development of those that have failed to attain maturity at an age of two years, is the accelerated development of those that have already more than doubled in length before the first hibernation, and continue to grow rapidly after emergence. By late spring they are already approaching adult size, perhaps even before laying has occurred, and while breeding is still in progress. It is certain that in northeastern Kansas there is no breeding by such accelerated individuals approaching adult size at an age of nine or ten months. Farther south in the species’ range with a much longer growing season, there is perhaps some possibility of such early breeding by first-year individuals. This would reduce by more than half the length of time required for a generation, and would tremendously increase the reproductive potential. With such added impetus to its reproduction the species might be able to withstand greatly increased predation pressure, or other mortality factors. Fig. 13. Growth curves of successive annual broods (designated by the year of hatching), superimposed to bring out differences in trends resulting from changes in weather from year to year. Extremes of acceleration or retardation are relatively rare in the population studied. Nevertheless, in April there are some individuals between 50 and 60 mm. in snout-vent length which cannot be classified with certainty as to their age group, and might be either accelerated individuals about nine months old or retarded individuals about 21 months old. The spread in size for any given age group is especially large, if data from different years are combined. A typical individual, having a snout-vent length of 25 mm. at hatching in mid-July may have attained 30 mm. by early August, 35 mm. by late August, and 45 mm. by the time it hibernates late in September. Emerging shortly before the middle of April it may grow to 50 mm. by the end of May, 58 mm. by the end of June, and more than 60 mm. by the end of July when it is a little more than a year old. By the time of its second hibernation it may have attained a length of from 65 mm. to 70 mm., and emerges from this hibernation as a breeding adult. Fig. 14. Records of growth of immature individual skinks, both hatchlings and yearlings, that were marked in one year and recaptured the next. In reptiles in general there is a wide range in adult size, and the extent and rapidity of continued growth after attainment of sexual maturity and minimum adult size is still insufficiently understood. Information bearing on this problem was obtained in the present study from the recapture of marked skinks already measured as adults. It is evident that the growth rate of the young, amounts to as much as 15 mm. per month in snout-vent length in the late summer period from hatching until hibernation, averages perhaps three or four mm. per month in the summer after emergence from the first hibernation, and tapers off rapidly as adult size is approached. One hundred of the skinks marked as adults or subadults and recaptured after intervals of months, including, in most instances, one or more hibernation periods, represent in the aggregate, 87 years of
Opportunity to compare the rapid growth of young during their first year of life with the relatively slow continued growth after attainment of sexual maturity is afforded by the records of skinks caught and marked while yet immature and recaptured in two or more successive years after their attainment of sexual maturity. The records of selected individuals of this group are presented below. With the exception of number three, all in this series are of the 1949 brood, and probably all hatched within a two-week period.
Differences in their growth rates therefore reflect differences in sex, individual vigor, and local situation, in individuals living at the same time and within the same general environment. Changing weather, and other factors that vary from year to year cause marked differences in the dates of important events in the annual cycle, and in the stage of development at any given date. Data are available for five successive annual broods of young, those of 1948, 1949, 1950, 1951, and 1952, and each brood differs from the others to some extent, as shown in Figures 11 to 13. In 1949, for instance, young hatched relatively early, and probably most of them were active by the middle of July. They made rapid growth in August, averaging larger than young hatched in other years on any given date in late summer. However, they retired into dormancy early in the fall. Cool and dry weather in early September ended their activity for the season. In 1950, young hatched, on the average, at least three weeks later, about the first of August, but they remained active until late in September, and by hibernation time had partly caught up to the stage of development attained by the young of 1949. Most young of 1951 hatched late in the first half of Fig. 15. Records of growth in another group of recaptured young that grew less rapidly than those of Fig. 14. The young of 1948, first sampled after their emergence from their first hibernation in mid-April of 1949, were then somewhat intermediate in size as compared with those of 1949 and 1950 at the same times of year. Their subsequent development was rapid; by late May they had caught up and passed the stage reached by the 1949 young at the same time of year. The young of 1950 after having a late start, were further set back by cold weather in April 1951 delaying their emergence from hibernation. As a result they were still unusually small in late April and May. Even though they grew rapidly subsequently, they were consistently smaller than those of other broods on corresponding dates. Favorable fall weather prolonging the 1951 growing season into late September beyond the time of retirement in other years may have permitted many of them to attain adult size. Fig. 16. Records of immature individual skinks marked and recaptured within the same growing season, showing the trend of rapid growth, and differences in growth rate between individuals. The varying fortunes of the several annual broods studied were closely correlated with weather trends, and suggest possible effects of slight changes in climate. An unfavorable sequence of weather might bring about drastic reduction of the population without causing any direct mortality. A late spring in two successive years would have cumulative effect in delaying emergence and breeding of adults the first year, and delaying in the second year emergence of the young, already retarded by the lateness of their hatching. If this sequence were followed by onset of unusually cool and dry weather in early September, or even in late August, the young might Under favorable conditions an adult female produces about ten offspring annually of which about half are females. It is calculated that if all survived, after ten breeding seasons, the progeny of an original female might have increased to a population of more than 97,000, under the climatic conditions of eastern Kansas, permitting attainment of breeding maturity late in the second year of life. In the same ten year period under climatic conditions delaying maturity until late in the third year of life (as seems normally to occur in E. septentrionalis and E. skiltonianus, and probably in E. fasciatus at the northern edge of its range) the original female would have produced a population of somewhat less than 7,800 assuming that all survived. With a long growing season such as occurs in the southern part of the range, it seems theoretically possible (though not probable) that individuals might mature before the end of their first year, in time to participate in the next breeding season. If this should occur the original female might produce a population of more than 120 million by the end of the tenth breeding season. Changes in PatternProgressive alteration of the color pattern is more rapid in males than in females and is synchronized with growth. During the first year of life changes in the pattern are gradual, and consist chiefly of loss in vividness. The blue of the tail is slightly dulled. The light lines become suffused with brown and the dorsolateral dark areas become paler, with light brown areas appearing on the corners of the scales and gradually spreading to replace the original black. In skinks that are in the second year of life the striped pattern although Fig. 17. Records of another group of immature skinks marked and recaptured within the same growing season. Even in hatchlings, the dorsal part of the rostrum and the inter-nasals are of a somewhat neutral brownish color, matching neither the light lines nor the dark interspaces of the striped body pattern. With advancing age this neutral brown color gradually spreads posteriorly on the head so that the striking lyrate marking of the bifurcated dorsal stripe on the head in the juvenile become obscured by the time the skink has grown to small adult size, at 21 months. The top of the head is then dull brown, with a slightly mottled appearance caused by the different intensity of pigmentation in different areas. The stripes though still discernible, are faint and
In tracing the gradual ontogenetic changes in the striped pattern, from the vividly contrasting colors of hatchlings to the dull, patternless coloration of old adult males, five descriptive terms have been applied to the successive stages: “sharp,” “distinct,” “dull,” “faint,” and “absent.” To most individuals below minimum adult size, the term “sharp” is applicable, although there is some loss in vividness in the larger young, as compared with hatchlings. Fading of the original striped pattern proceeds more rapidly on the head than on the body. Upon emergence from their second hibernation at an age of about 21 months, the skinks, mostly grown to adult size, and ready to mature sexually, still show but little sexual difference. They retain the hatchling pattern essentially unchanged, but with colors dulled and contrasts reduced. Within a few weeks the newly matured males undergo relatively rapid color change as the breeding Table 13 refers to adult pattern and coloration as they appear in the breeding season. After the breeding season, in late spring and early summer, when the red suffusion of the head and neck has faded in adult males, the original striped pattern, after having been almost completely suppressed may again become discernible. Individuals of the same size differ in extent of pattern change, and the color descriptions made of individuals were not sufficiently detailed to show fully the changes occurring between successive dates of capture. However, most large adult males taken later than mid-June had at least some trace of the striped body pattern and many of them had become so much like females in appearance that close scrutiny was necessary to determine their sex. They were especially like females in having the dark lateral area extending forward onto the cheek and setting it off sharply from the paler temporal region above it. In breeding males the head has no such dark markings and is suffused with red. Growth and Regeneration of the TailEven among those skinks which have never broken their tails there is a wide range of variation in relative length of tail. This is partly a matter of relative growth since the proportions change during the course of development. Also there may be slight sexual difference and there is much individual variation. In fetuses still well below hatching size, the tail length is less than the snout-vent length. For instance, an egg in a natural nest 12 days short of hatching contained a fetus that had a snout-vent length of 14 mm. and tail length of 12 mm. (Figure 18). In the late stages of fetal development the tail growth is relatively rapid. At hatching, the tail is considerably more than half the total length. In a large series of young with snout-vent lengths from 30 mm. down to hatching size of 25 mm. or less, the tail length averaged 130.8 per cent of snout-vent length. In larger young, up to a snout-vent length of 40 mm. or more, the tail continues to lengthen more rapidly than the body. In skinks that are about two thirds grown, the tails average relatively longer than in either larger or smaller individuals. In the sample representing the size class 50-54 mm. snout-vent length, the tails average 163.3 per cent of the snout-vent lengths, whereas in groups of adults of various sizes and both sexes, the tail length Fig. 18. Diagram showing relative tail-length (as a percentage of snout-vent length) in skinks of different size groups that retain their original tails unbroken; in the early stages of growth the tail becomes relatively longer as size increases, but the trend is reversed before adult size is attained. For each series the mean, standard error, standard deviation, and extremes are shown. When a skink’s tail is broken, there is almost no loss of blood. The fractured surface is rough and irregular, with exposed muscle masses protruding on the detached end and corresponding concavities on the end of the stump tail retained by the lizard. The concavities are soon filled with oozing blood, and a thick scab forms. As healing begins, the broken end presents a flat, slightly irregular surface. When the scab is sloughed off, a slightly convex surface of delicate, pale-colored new skin of the regenerating tail, is exposed. At first, no scale structure is discernible. As growth proceeds, the new tail takes on a bluntly conical shape. During the early stages of growth, it is well set off from the original portion by the abrupt Fig. 19. Relative lengths of original and regenerated portions of tails in skinks which have had their tails broken and regenerated; for each individual, length of each part of the tail is expressed as a percentage of the snout-vent length. Rate of growth in the regenerating tail is controlled by a variety of factors, such as age, condition, and activity of the individual, and Fig. 20. Diagrams showing typical extent of tail regeneration in skinks having tails broken at different points, × approximately 1/2. Original parts of the tails are at the right.
Under favorable conditions regeneration occurs at a relatively rapid rate. After a period of healing the new tail grows with a sudden spurt, making most of its gain in length within a few weeks. Then growth abruptly slows or ceases altogether. In young similarly rapid growth of the regenerating tail occurs, but subsequently Successive records of selected individuals are listed in Table 14 to illustrate trends in regeneration of the tail. In those instances in which the tail is referred to as “newly broken” the separation usually occurred as an accident at the time the lizard was captured, while in those designated as “recently broken” separation had already occurred in some earlier accident but regeneration was not yet perceptibly underway. In the “Tail length” column, plus signs separate the original portion of tail, on the left, from the regenerated portion, on the right. As in many other kinds of lizards, the tail in the five-lined skink serves as a reservoir for fat, which may be drawn upon for nutrition in time of food scarcity. An individual that is in good condition has a plump and rounded tail. Fat comprises much of its bulk. Upon emergence from hibernation this fat supply is not noticeably depleted. Brooding females in the latter part of the incubation period have the supply of caudal fat most noticeably depleted, and their tails may appear emaciated, with kinks on the terminal portion. It is my impression that in adults the capacity for storage of fat is most developed in the females, and that their tails vary in proportions more than do those of males. The capacity to shed the tail easily seems somewhat inconsistent with this function of fat storage. Loss of the tail sometimes involves loss of a large amount of reserve fat. Many detached tails that were broken accidentally at the time of the skinks’ captures were weighed. In those that were broken off near the base and were not previously regenerated, weights were usually 16 to 20 percent of the lizards’ total weights. | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
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