Life History and Ecology of the Five-lined Skink, Eumeces fasciatus Part 4

n.o.ble and Bradley (1933:77) mention frequent h.o.m.os.e.xual matings between captive males. However, I observed no h.o.m.os.e.xual matings, either under natural conditions or in confinement. The pugnacious behavior of males that are in breeding condition ordinarily would prevent h.o.m.os.e.xual mating. Males in such weakened condition as to be unable to defend themselves effectively might evoke s.e.xual 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 s.e.x 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 s.e.x 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 male stickleback, it facilitates s.e.x recognition and evokes hostile behavior on the part of other males. Courtship, mating, and fighting reactions however, seem to be evoked by the interaction of a complex of social releasers. Whereas males and females are strikingly different in appearance in the breeding season, visual s.e.x recognition is complicated by ontogenetic changes. The body stripes characteristic of the female pattern, become dull or even disappear in some old females, which then approximate the typical male pattern. On the other hand newly matured males in their first breeding season retain distinct body stripes of the female pattern. Their s.e.x is evidenced mainly by their reddish facial suffusion, which is not quite so extensively developed as it is in older individuals. Also, in these newly matured males the temporal region is not so swollen as it is in old males.

The male whose dormancy was terminated in early winter by bringing him into a warm room causing him to a.s.sume 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 s.e.xual 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 s.e.x 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 s.e.xual behavior of lizards has not been appreciated, although n.o.ble 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 summer climate in the southern states, dates of laying and hatching may be several weeks advanced. On April 12, 1952, Dr. Wilfred T. Neill showed me several live _E. fasciatus_, collected a few days before along the Trinity River in southeastern Texas, which appeared to be at the height of breeding condition. In northeastern Kansas on that date, general emergence had not yet occurred, and it was not until about May 10 that the population attained the peak of breeding condition. On May 8, 1948, near Burr Ferry, Vernon Parish, Louisiana, I caught an adult female in her nest burrow, and she contained eggs ready to be laid. Data with which Mr. Robert Gordon kindly provided me for specimens from southern Louisiana and southeastern Texas, in the Tulane University collection, indicate gravid females on June 4, 1952, and June 17, 1948 (3), and females with their egg clutches on June 16, 1948, June 17, 1948, June 23, 1950; and hatching dates in captivity of July 19, 1949, July 19, 1950, July 25-26, 1949. These dates correspond well with those for specimens obtained in northeastern Kansas in the same years. In the northern part of the range, Ruthven (1911:264) recorded that in the Saginaw Bay region, females taken on June 19 had eggs nearly ready to be laid, and after July 2 clutches were found frequently; young of the year were first observed on July 31. A juvenal specimen in the University of Minnesota Natural History Museum, collected on August 11, 1938, at Dresser Junction, Wisconsin, is 30-1/2 mm. in snout-vent length--approximately the size of juveniles in northeastern Kansas at the same season. Evans and Roecker (1951:6) record hatching as occurring in the first week of September at Arden, Ontario, indicating that at the northern edge of the range hatching may be delayed as much as two months. With such delayed hatching, but little time remains for the young to grow before they are forced into retirement for hibernation.

FIGHTING

Territoriality 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 encompa.s.s only a small fraction of the entire home range. The senses are inadequate to inform one lizard of the presence of another until the two are only a few yards, or even a few inches apart. Usually the lizard is on the ground, where even small objects obstruct its view, and vision is probably effective for only a few yards. Hearing is probably effective for about the same radius in detecting animals of approximately its own size. Scent is effective in detecting prey near at hand or on contact, but probably does not serve for detection of other lizards that are not in the immediate vicinity. Therefore, the area covered by one in the course of its normal activities may harbor many others, and individuals most of the time are unaware of the others on their home ranges.

Under most circ.u.mstances 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 s.e.x 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 they dodged and sparred.

As one broke away to run, the other seized it by the tail. They were on an exposed tree root about an inch in diameter. The skink that was caught twisted its body around underneath the root and seized its adversary by the tail likewise, so that their linked bodies encircled the root, each squirming to disengage itself from the other's jaws.

After a few seconds they did break apart, and then maneuvered briefly menacing each other at close quarters, but they gradually moved away and lost contact.

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.

[Ill.u.s.tration: 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 other's jaws. The one caught in the attacker's grip was usually unable to flex its body sharply enough to reach its opponent at all, or could barely reach it at such an oblique angle that its jaws slipped off the smooth body. Sometimes the one held did succeed in catching the other's front foot. The one caught in the other's jaws always succeeded in tearing loose after a short time. In the interval while the attacker rested with jaws partly relaxed, the victim had an opportunity to break away. Even when both were free, they did not obtain grips easily, but often made several unsuccessful lunges and bites, the jaws of each slipping off the firm, smooth sides of its opponent. Sometimes the attacker seized a fold of skin, or sometimes obtained a wide grip on its body. One which had obtained a grip sometimes rolled rapidly, spinning the other around and dashing it against the ground. As these rotations stopped, the victim might come to rest on its back in such a position that it was temporarily helpless, but always broke loose after further struggles. Neither showed any inclination to retreat until finally, when they were interlocked, rolling about almost at my feet, I attempted to catch them. Then they instantly disengaged and rushed away, and one escaped. The one caught had suffered but little injury in the fight.

Numerous tooth marks were discernible as minute abrasions on the surface of the scales, but the bony dermal armor had not been perceptibly penetrated during the prolonged and violent struggle.

EGGS

The 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 sh.e.l.ls. These sh.e.l.ls are thin and easily punctured. As incubation proceeds, the egg enlarges by gradual absorption of moisture and the somewhat elastic sh.e.l.l 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 sh.e.l.l. 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 hatch from them. Both in the field, and in laboratory experiments, eggs were found to have remarkable tolerance for excess moisture. After heavy rains of summer thunderstorms, nests were sometimes found to have water trickling through them, and on occasion eggs were found to be partly submerged in water in the nest cavity. Exposed rocks at the heads of small gullies often were chosen by the female skinks as the shelter for their nests.

In these situations the nests were exposed to run-off water. In July, 1951, especially, unusually heavy precipitation resulted in the flooding of many nests. In some instances desertion by the females and destruction of the eggs seemed to have been caused by this flooding, even in the well-drained hillside situations where this study was made.

Table 5.--Measurements in Millimeters and Weights in Grams of Eggs in the Same Clutch at Different Stages During Their Incubation, Showing Gradual Increase in Size.

===============+=========+========+========+========+========+========+========+=========== June 17 June June June July July July July 30 (laid) 18 24 28 17 20 28 (hatched) ---------------+---------+--------+--------+--------+--------+--------+--------+----------- Average length (for 7) ..... .... .... .... 13.7 14.3 14.7 .......

Average width (for 7) ..... .... .... .... 10.5 10.9 11.1 .......

Typical length ..... 11.1 .... 12.5 14.0 14.3 14.8 .......

Typical width ..... 7.5 .... 9.9 11.0 11.2 11.0 .......

Maximum length ..... 11.5 .... .... 14.5 15.0 15.5 .......

Maximum width ..... 7.5 .... .... 10.9 11.1 11.4 .......

Minimum length ..... 10.5 .... .... 12.5 12.8 13.5 .......

Minimum width ..... 7.0 .... .... 9.9 10.0 10.5 .......

Average weight ..... .38^10 .58^5 .63^9 .82^8 .90^7 1.0^7 .......

Typical weight ..... .4 .... .... .... .9 1.0 .......

Maximum weight ..... .... .... .... .... 1.0 1.1 .......

Minimum weight ..... .... .... .... .... .7 .7 .......

Superior number indicates the number of individuals averaged.

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, which was a week short of hatching. On July 22 another egg of the same clutch was immersed and left in water for 23 hours. On July 30 it was ruptured in handling and found to contain a living fetus. On July 31 two eggs were placed in a dish of water in a refrigerator. On August 5 they were removed and opened. Fetuses were dead and were not appreciably larger than the one of the same clutch in the egg opened on July 31. On August 5 two of the remaining eggs of this clutch were placed in a Petri dish, partly immersed in water, with approximately one-fourth of the surface of each protruding and exposed to the air. Forty-eight hours later it was found that both eggs had hatched. Evaporation had reduced the water in the dish to an amount sufficient to cover only about the lower one-third of each egg. One hatchling was missing, evidently having climbed out of the shallow dish and escaped to the floor. The other was found still standing in the water with its head protruding, and it was lively and in good condition.

The remaining four eggs in this clutch, which had been kept in a container of damp earth, were also hatching on this date. On July 10, 1952, an egg in a late stage of incubation was immersed in water in the laboratory. On July 14 when removed, it had fungus growing on it, and was found to have a dead fetus, nearly full-sized.

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.

1. July 8, 1952 Egg transferred from natural nest to jar of damp soil in refrigerator at 13.8C.

July 14, 1952 Seems to be in good condition.

July 19, 1952 Partly collapsed. Weight and measurements same as on July 8; opened and found to contain a dead fetus.

Snout-vent length 23 mm., forehead bulging, skin delicate and membranous. Colors somewhat dull, indicating that it was not quite fully developed, although it had attained the minimum hatching size.

2. July 10, 1952 Egg from natural nest (15.0 10.5 mm., .95 gm.) put in refrigerator at 11.6C. Control (14.5 10.6 mm., .8 gm.) from the same clutch kept in hatching medium in laboratory.

July 13, 1952 Control egg hatching; refrigerated egg shows no indication of hatching.

July 14, 1952 Experimental egg 15.8 10.8 mm., 1.0 gm., seems to be in good condition. Nest from which it was taken found to have all remaining eggs hatching today.

July 19, 1952 Experimental egg 15.0 10.0 mm., 1.0 gm., removed from refrigerator and transferred to container in damp rotten wood in laboratory. Seems to be in good condition.

July 23, 1952 Experimental egg found to be hatched this morning, and hatching must have occurred either in the night or late yesterday. Eggsh.e.l.l still damp and pliable.

3. July 10, 1952 Egg from natural nest (14.0 10.5 mm., .8 gm.) put in refrigerator at 11.2C., in container with damp decayed wood. Control egg (14.2 10.1 mm., .8 gm.) from the same clutch kept in the same hatching medium in the laboratory.

July 12, 1952 Nest from which experimental and control eggs were taken has started to hatch, and two hatchlings were seen there.

July 13, 1952 Control egg hatched.

July 14, 1952 Experimental egg 14.2 10.1 mm., .8 gm., seems to be in good condition. Nest in field examined and all eggs were hatched, with only three of the hatchlings remaining, the others having dispersed.

July 19, 1952 Experimental egg 14.0 10.0 mm., .95 gm., still appears to be in good condition; removed from refrigerator and kept in laboratory.

July 23, 1952 Experimental egg found to be hatched, and hatchling active although still in hiding beneath rotten wood.

Probably it hatched early in the day of July 22; the empty sh.e.l.l is still moist.

These experiments seem to show that, in the later stages of incubation at least, lowering of temperature to 11 or 12C. 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.