Life History and Ecology of the Five-Lined Skink, Eumeces fasciatus

The five-lined skink is confined to a region where summer rains are frequent. It is evident that a regular supply of drinking water is one of the most critical ecological requirements. Bogert and Cowles (1947:19) found that an E. inexpectatus experimentally kept at high temperature lost moisture at a more rapid rate than any other reptile tested (including two other kinds of lizards, four kinds of turtles, an alligator, and three kinds of snakes). They remarked that this rapid moisture loss presumably accounts for the inability of skinks to survive in containers when no moisture is readily available, and also accounts for their absence in truly arid habitats. The Natural History Reservation is situated near the western edge of the species’ range in a climate that may be near the limit of its range of tolerance. However, on most summer mornings low woodland vegetation is copiously laden with dew, and this evidently fulfills the need for drinking water. Diminution of surface activity and retirement to underground retreats seem to be closely correlated with cessation of rains in late summer. After rainless periods in August and September, when morning dew is no longer available these skinks, especially the adults, are no longer regularly seen in the open. They have retreated to underground shelters where they spend nearly all their time. The time of disappearance varies from year to year and the correlation with varying weather conditions seems obvious. While no actual experiments were performed to determine the moisture requirements, it is evident that the need for moisture rises sharply with increased temperature. Skinks that are dormant in hibernation survive for periods of months without drinking, with but little loss of weight. In their underground shelters temperature is low and presumably relative humidity is high. At temperatures above their optimum of approximately 34°C. the skinks are especially subject to rapid moisture loss, since evaporation of body moisture is resorted to as a device to keep the temperature below the lethal level. The skinks subjected to extremes of temperature in an experimental terrarium were seen to lap up condensed moisture on the cooled metal plate at intervals of a few minutes. After an hour or more in the experimental terrarium they seemed somewhat debilitated. Skinks brought from the study areas to the laboratory for weighing and other records, were ordinarily returned on the following day. When circumstances prevented adherence to this schedule in hot summer weather, mortality could be expected in the skinks kept in cloth bags or glass containers, unless water was provided. Dramatic weight loss of up to more than 30 per cent was recorded in some individuals, kept at the high temperatures which usually prevailed in the laboratory, over periods of days in the summer. Skinks having access to drinking water often ingest amounts far beyond their immediate requirements, which may be stored in the bladder and drawn upon over periods of days as it is needed, or may be utilized to dampen the soil of the underground shelter and raise the humidity, as incubating females seem to do.

Geographic Range and the Deciduous Forest Habitat

Eumeces fasciatus corresponds in its distribution with the original hardwood forests of eastern North America, as mapped by Braun (1950:cover folder) and the “Oak-Wild Turkey Biome” of Shelford (1945:240). Few species of vertebrate animals have ranges that coincide more closely with this extensive area (exclusive of the northern edge, that part characterized by Braun as the Hemlock-White Pine-Northern Hardwoods). This latter is a mixed forest which actually is transitional between the more typical deciduous forest farther south and the Taiga Biome (or Formation) to the north, which is dominated entirely by conifers. At the northern edge of its range Eumeces fasciatus is much less generally distributed than it is farther south. Although it is well established and even may be locally numerous in South Dakota, Minnesota, Wisconsin, northern Michigan, Ontario, northern New York, and Connecticut, the locality records from these states are few, and seemingly represent isolated and widely separated colonies that are able to persist because of favorable combinations of environmental factors not of general occurrence in the surrounding regions. Figure 6 shows the extent of the hardwood forests as mapped by Braun (excluding the transitional Hemlock-White Pine-Northern Hardwoods Association) with specific locality records of E. fasciatus included in all outlying portions of the range. The locality records are those published by Taylor (1936:206-212) supplemented by other marginal records, more recently published, by Hamilton (1947:64) for New York, Breckenridge (1944:97) for Minnesota, Hudson (1942:42) for Nebraska, Smith (1950:185) for Kansas, Brown (1950:116) for Texas, Neill (1948:156) for Georgia, and Neill and Allen (1950:156) for Florida. Along the northern edge of its range, the skink invades the Hemlock-White Pine-Northern Hardwoods Association, in Massachusetts, New York, Pennsylvania, Ontario, Michigan, and Wisconsin, but does not penetrate far into it anywhere. Correspondence of its northern limits with those of the Oak-Chestnut, Maple-Basswood, Beech-Maple and Oak-Hickory associations is remarkably close, considering the fact that the boundaries of these climax associations are not sharply defined; rather they merge by gradual stages into the northern coniferous forests, with outlying peninsulas and islands where conditions are favorable.

The outlying northern localities where E. fasciatus occurs within the Hemlock-White Pine-Northern Hardwoods Association are all within the region of Pleistocene glaciation, which 20,000 years ago, or even more recently, were covered with the continental ice mass during Wisconsinan time. Yet the localized northern populations of skinks evidently are relicts from a time when favorable conditions were more widespread in the general region. Braun (op. cit.:464-465) indicates five successive postglacial stages in the trends of climate up to the present, as revealed by bog pollen profiles: (1) Cool and moist; (2) warm and dry; (3) warm and humid; (4) warm and dry; (5) cool and moist. Stages 2 and 4 would have been most favorable for encroachment of the skink into glaciated regions, whereas stages 3 and 5 might have caused retrenchment of its populations. In view of the localized habits of individuals, and the lack of any mechanism for rapid dispersal, the time available seems no more than adequate for the distance of 200 miles or more northward that the skinks must have moved since the final retreat of the ice sheet. This northward movement involved crossing of formidable barriers such as the Great Lakes. Even minor barriers such us small rivers and creeks, might be expected to halt population movements for long periods.

The over-all geographic range is approximately square, roughly a thousand miles across, from north to south and from east to west. On the east and south it is limited by the Atlantic Ocean and the Gulf of Mexico. On the north and west its limits correspond with those of the hardwood forests. On the northwest, it reaches southwestern Minnesota and the southeastern corner of South Dakota, extending far out into peninsular extensions of the Oak-Hickory Association which penetrate westward into the prairies along the main river valleys.

In Kansas it occurs over the eastern one-fourth, west to the Flint Hills, and a little farther west in peninsular extensions of the forest along some of the main river valleys. In Braun’s map the Deciduous Forest Biome is shown to reach only the eastern edge of Kansas along the Kaw River and Missouri River at and near their junction, the Osage (or Marais des Cygnes) River valley near the Missouri border, and the southeastern corner of Kansas. However, for almost 100 miles farther west from the Missouri border, the country has the aspect of a savannah with scattered groves of trees on hillsides and along streams, providing suitable habitat. The distribution of the five-lined skink in eastern Kansas corresponds well with certain “Biotic Districts” as mapped by Cockrum (1952:12), namely the Kaw River, Osage Savannah (southern part), and Cherokee Prairie. Conversely the skink is excluded from the Short Grass Plains and Mixed Grass Plains Biotic Districts which occupy nearly all of the western three-fourths of the state. There are two specimens in the University of Kansas Natural History Museum, labelled Ranson, Ness County. This locality, in the western third of the state, more than 150 miles from any other recorded station, may represent an isolated colony; however Smith (1950:185) states that the record needs verification, and it is not included in the map, Figure 6.

In Oklahoma the distribution records fit fairly well the portion of the state mapped by Braun as the Oak-Hickory Association of the Deciduous Forest, but extends a little farther west in the northeastern part of the state. A game type map published by the Oklahoma Game and Fish Department, Division of Wildlife Restoration, in 1943 shows in more detail distribution of the main vegetation types within the state. The locality records for the skink fall almost entirely within three of the fifteen vegetation types mapped, namely, the oak-pine, and oak-hickory forest of the state’s eastern edge and the post oak-blackjack oak type of the eastern and central parts. The locality records extend almost throughout the area occupied by these three types but not in attenuate westward extensions of the post oak-blackjack type that occur along several of the main stream courses. In Texas likewise the recorded localities fall mainly within the area mapped as deciduous forest, but with several slightly beyond its boundaries. In a detailed map of the “game regions” of Texas (Anonymous, 1945:1), some of these outlying localities fall into the coastal prairie area, and the remainder into the post oak and blackland prairie belts, which grade into each other and the oak-hickory forest.

The former distribution of the five-lined skink may be postulated on the basis of the fossil record of its community associates since it is a primitive and conservative type. Taylor (1936:56) explained the present discontinuous distribution of the genus on opposite sides of the world on the basis of a former northern connection of the continents. He wrote: “I regard migration from North America to Asia as having taken place via land bridges joining the Alaskan peninsula with Asia either at Bering Straits or via the Aleutian Island arc to Kamchatka, or both. One would need postulate but slight climatic changes since the present climate of this coastal region is probably no more rigorous than that of southern Canada which has three species of the genus.” However, such former northward distribution, while entirely probable, would have been possible only in a climate much milder than that which prevails at present. In Asia, tunganus on the mainland and latiscutatus on the island of Hokkaido extend north to about latitude 43°, and in North America, fasciatus extends slightly farther north. In order to have crossed between Alaska and Asia on presumed land bridges these skinks would have had to extend their ranges about 20 degrees north of their present limits, into what is now a cool climate. The winter climate of the Bering Sea is perhaps not much beyond the range of tolerance of the more cold-adapted forms of Eumeces, but the cold, cloudy, wet, and changeable summer climate is far beyond the range of tolerance of Eumeces or any other lizard.

It is highly improbable that the fossil record will yield direct evidence for the existence of a northern ancestral Eumeces of the fasciatus group. The characters by which the various forms are recognized are to be found mainly in details of pattern and scalation; the skeleton is so conservative that specific characters are ill defined or lacking even in well preserved fossil material. This hypothetical ancestor probably was a member of a deciduous forest community having components in common with the modern forests where the American and Asiatic species occur, along with types now extinct, and others which, though existing at the present time, have become separated from their original associates and occur in other regions.

Hollick (1936:11) has described a rich early Tertiary Alaskan flora strikingly different from that of the same region at the present time. Composed of genera now characteristic of warm-temperate to subtropical climates, it was remarkable in having many types of plants that are now most characteristic of the North American hardwood forests in the southeastern part of the continent. Besides such widespread genera as Fagus, Betula, Ulmus, Platanus, Castanea, Corylus, Carpinus, Crataegus, Spiraea, Myrica, Smilax, Pinus, Picea, and Abies, this flora included others now characteristic of both warm-temperate southeastern North America and Eastern Asia, as Magnolia, Nyssa, Sassafras, Persea, Benzoin, Hamamelis, Liquidambar, Celastrus, Nelumbo, and Onoclea. It included genera Carya, Taxodium and Comptonia that now are limited to SE North America, Sequoia, now limited to western North America, and also included several genera which at present are limited to southeastern Asia: Ginkgo, Glyptostrobus, Cinnamomum, Hausmannia, Artocarpus, Dillenia and Koelreuteria. This fossil flora provides strong evidence that in the early Tertiary climatic and habitat conditions as far north as Alaska were favorable for the existence of an ancestral Eumeces similar to the modern E. fasciatus, which might have given rise to both North American and Asiatic members of the fasciatus group.

There is abundant evidence for the existence of an Eocene land connection between Alaska and northeastern Siberia, permitting free interchange of faunas between the two continents, as shown by the almost simultaneous appearance of various mammalian groups in the fossil records of Asia and North America. Simpson (1947:627) has summarized the evidence that such intermigrations were occurring throughout most of the Tertiary, with occasional interruptions as in middle Eocene, and in middle and late Oligocene, and with increasing selectivity, chiefly a progressive tendency toward screening out of the groups less tolerant of cold (judged on the basis of their modern representatives). In the late Tertiary, and especially in the Pleistocene, animals known to have made migrations between North America and Asia were types now characteristic of boreal climates (e. g. pika, hare, vole, lemmings, marmot, jumping mouse, fox, wolverine, bear, moose, caribou, sheep, bison, camels, mammoth). Simpson believes that there was fairly strong climatic selectivity even in the Miocene interchanges, and he indicates several important groups that were non-migrants in the Miocene, most of them remaining so through the Pliocene and Pleistocene—the primates, Rhizomyidae, Gliridae, Viverridae, Hyaenidae, Dicerorhininae, Suidae, late Anthracotheriidae, Hippopotamidae, Tragulidae, Muntiacinae, Lagomerycidae, Giraffidae, and Bovidae. He states that there is good evidence that these are all mainly warm-climate animals which are not likely to have ranged in any force into a cold-temperate or boreal environment. In view of these conclusions it seems doubtful whether Eumeces or other reptiles could have crossed the Alaskan-Siberian land connection so late as the Miocene.

On the contrary, the climate and habitat conditions with which Eumeces might have been associated, although present as far north as Alaska in the Eocene, evidently had shifted far to the south by mid-Tertiary time. Axelrod (1950:230) has described a Miocene forest of the Columbia Plateau and northern Great Basin indicative of a uniform temperate climate and an average rainfall of thirty-five to sixty inches. This forest included: (a) various genera now characteristic of the southeastern hardwood forest or confined to it—Carya, Castanea, Comptonia, Fagus, Liquidambar, Nyssa, Taxodium; (b) other genera at present more characteristic of the western United States—Sequoia, Lithocarpus, Pseudotsuga, Mahonia, Thuja, Gaultheria, Amelanchier; (c) wide-ranging genera including Alnus, Acer, Betula, Populus, Quercus, Picea, Pinus, Tsuga, Cornus, Ribes, Rosa, Hydrangea; (d) modern east Asian genera, including Ginkgo, Ailanthus, Glyptostrobus, Keteleria, Koelreuteria, Metasequoia, Pseudolarix, Pterocarya, Zelkova, which were eliminated from the North American flora in the latter part of the Tertiary. In short, this western Miocene forest was remarkably similar in many respects both to the presumably ancestral early Tertiary Alaskan forest and the modern southeastern hardwood forest. The extent of this Miocene forest is unknown but judging from the sites where it has been recorded, it had progressed about halfway, both in latitude and in actual distance, from Alaska to the area occupied by the modern southeastern deciduous forests. Several other reptilian genera have distributions similar to that of the fasciatus group, with representatives in southeastern Asia and southeastern North America that probably have parallel histories of distributional divergence from early Tertiary northern ancestors similar to contemporary species (Schmidt, 1946:148-150). Alligator, Natrix, Ancistrodon, Scincella, Elaphe, Opheodrys, and within the genus Eumeces, the obsoletus group, all provide excellent examples.

Effect of Climatic Factors

Accounts of the habits and habitat, by various authors, indicate versatility in behavior, and adaptation to a variety of habitat conditions in different climates and plant associations. Some of the differences evidently result from the skink’s tendency to maintain itself in surroundings of favorable temperature and humidity, which obviously are to be found in different types of situations at different extremes of the range. Hence even though the skink itself may remain unchanged, it tends to behave somewhat differently under diverse environmental conditions. Such environmentally enforced differences in habits would be difficult to distinguish from those having a genetic basis. Although no subspecies of Eumeces fasciatus have been recognized, local populations undoubtedly differ somewhat in size and other characters that have a genetic basis.

At the northern edge of its geographic range, fasciatus occurs in isolated colonies and seems to be restricted to open, rocky situations which receive the maximum amount of sunlight. Breckenridge (1944:96) wrote that at the two Minnesota localities representing the northwestern corner of the known range, the skinks were found at granite outcrops, and he mentions one found in western Wisconsin, at Taylor Falls, under an 18-inch slab of a basalt outcrop in sparse oak woods. Patch (1934:51) described a habitat at Arden, Ontario, among massive granite-gneiss domes, with sparse vegetation. At Point Pelee, Ontario, the species is common in the drier, more sparsely wooded situations, hiding beneath loose bark of stumps and logs.

Ruthven (1911:264) found E. fasciatus in the vicinity of sandy beaches in the Saginaw Bay region of Michigan. Elsewhere in its range it is more characteristically an inhabitant of hardwood forests, preferring the better drained and more rocky situations, according to the testimony of numerous authors. In eastern Illinois, Smith (1947:33) found it confined to the area south of the Shelbyville moraine, and not ranging into a prairie habitat. Near Elkville, Illinois, Cagle found the species abundant in higher and drier areas within sparse stands of oak in second growth woods, but it was absent from the low swampy areas adjacent to streams. Conant (1951:30, 210), describing the habitat in Ohio, stated that the species does not occur in swamps and areas that are subject to spring floods nor on dry hillsides, but is abundant in some areas where there are rotting stumps and logs remaining from former patches of swamp forest, and usually is found in low, moist situations, in wooded valleys or even at the edges of swamps and bogs. Lynn (1936:49) wrote that in Virginia, it is most often seen on steep, boulder-strewn hillsides and old sawdust piles. In the central Ozarks of Missouri, Owen (1949:49) found it abundant and saw it almost daily on rocky ledges, fallen timber, and fence rails, while E. laticeps was seen only once. Taylor (1936:59) wrote that E. fasciatus occurs where there is timber and is often found about fallen trees and rotting stumps, or about old sawmills where wood refuse has accumulated. Smith (1950:187) wrote that in Kansas the species is commonly found in wooded areas in moist situations about stones, leaves and rotten logs. Gloyd (1928:120) wrote that in Franklin County, Kansas, E. fasciatus occurred in upland situations and was the most abundant lizard where there were rocks, brush, or decaying wood. Gloyd (1932:401) also recorded it as abundant in the Pigeon Lake area, Miami County, Kansas, in wooded areas of sufficient elevation to be out of the river flood-plain.

Habitat in Northeastern Kansas

In northeastern Kansas I have collected or observed this skink in several dozen localities, and searched unsuccessfully in numerous other localities. Absence of this skink, in some situations and its presence and relative abundance in others, provided a basis for appraising the environmental factors that are of critical importance. River valleys, of the Kaw and Wakarusa and their tributaries, with deep alluvial soil, alternate with flat or rolling upland some two hundred feet higher in elevation, and having shallow, rocky soil. Where the uplands slope to the valley floors, there are steep hillsides, usually with extensive limestone outcrops along their upper slopes. The alluvial plains formerly supported hardwood forests, while the hill slopes and uplands were largely prairie. At the present time the bottomland forest has been almost completely destroyed, as it grew on the most fertile and potentially productive soil, and has been replaced by cultivated crops. There are still trees along streambanks, and in occasional woodlots, but I have failed to find any skinks in such situations. I doubt that they ever have been numerous in the bottomland woods; lack of rocks for shelter, and periodic flooding are unfavorable factors. In the Kaw flood of June and July, 1951, for instance, the entire valley was inundated, and in smaller tributary valleys such as that of the Wakarusa, flooding is frequent at the season when skinks are incubating their eggs. The uplands, formerly prairie, now are used partly for cultivated crops and partly for pasture. The soil is poor and rocky, and now heavily eroded. The pastures mostly have a weedy type of vegetation indicative of overgrazing. Five-lined skinks are absent from most of this upland.

The steep slopes from the upland to the valley floor are now mostly wooded, and the population of skinks is chiefly in this band of woodland. Some of the hillsides that have relatively gentle slopes are treeless and are used for pasture, or are even under cultivation. Where second growth forest is present its aspect differs depending upon slope, exposure, and past treatment. Osage orange and honey locust are aggressive invaders on some dry hillside pastures, and in this type of woods the skinks are scarce or absent. Some hillside areas, especially on moist north slopes have thick second-growth woods, in which elm is usually the principal tree, with several oaks and hickories, walnut, hackberry, coffee tree, locust and osage orange, and with a dense understory vegetation of dogwood, gooseberry and coralberry, with vine tangles of grape, poison oak, and greenbrier. Such woodlands provide little food for livestock, and are often fenced off from adjacent pastures. The shading creates conditions unfavorable for skinks and they are relatively scarce in the denser woods. They are much more numerous in woodlands that are fenced in with pastures heavily grazed by cattle or horses, with understory vegetation kept cropped back, and with more open ground and patches of sunlight. However, they are absent or scarce in woods that have been subjected over periods of years to browsing, by sheep or goats, so heavily that hardly any herbaceous vegetation remains and so heavily that the soil is packed from trampling. Along the upper slopes, especially about heads of gullies, in areas strewn with flat rocks, in fairly open mixed woods, with some decaying wood on the ground, habitat conditions are most nearly optimum for the skinks. Artificial habitat features, such as rock piles, stone walls, wood piles, rail fences, or old deserted buildings and sheds, with loose boards lying about on the ground may support unusually high concentrations of skinks when the surrounding habitat is favorable.

Study Areas

The University of Kansas Natural History Reservation where most of the field work for this study was done, has been described in a recent publication (Fitch 1952:8). While records were obtained from scattered points throughout the 590-acre Reservation and elsewhere in northeastern Kansas, field study of this skink was concentrated on four relatively small areas totalling only about ten acres in extent (Figure 26). These areas were selected on the basis of abundance and availability of the skinks, and of variety of habitat conditions represented.

One of these sites was a deserted quarry on a southward projecting spur of the plateau-like cuesta top, where the upper layers of the Oread limestone are prominently exposed. In the course of operations, begun about 1937, the area was denuded of trees and shrubs, and the upper layers of limestone were removed from a strip about 50 feet wide and more than 100 yards long. The exposed outcrop presented a vertical rock face five to ten feet high, with south and southeast exposure. Numerous jagged seams and fissures in the rock hastened its disintegration. Quarrying had been discontinued several years before the present study was begun in 1948. At that time there were talus-like accumulations of rock and soil several feet wide along the base of the rock face, supporting a luxuriant pioneer vegetation especially, sweet clover, stickleaf, ragweed and elm seedlings.

The habitat conditions provided by the exposed rock outcrop at the border of woods and open land, proved unusually favorable for reptiles in general, and it was one of the most productive sites on the Reservation for Sonoran skinks, collared lizards, racerunners, ring-necked snakes, blue-racers, bull snakes, pilot blacksnakes, scarlet king snakes, slender tantillas, copperheads, and timber rattlesnakes. For the five-lined skink, however, this disturbed area was marginal, and supported only a sparse population. Several decaying two-inch boards were preferred hiding places where the skinks were found most frequently, and remains of collapsed rock walls, one in the center of the area and one at the edge of the woods, were also occupied. Skinks may have tended to wander away to more favorable situations or may have been more subject to predation than those elsewhere, since the incidence of recaptures was relatively low. Most of the records from this general area were from a ledge in adjacent woods rather than from the quarry itself. Another site was a rock fill in a ravine below a pond made in 1937. This rock fill was 70 feet long, up to 30 feet wide, and three feet deep. East and north of the rock pile was a grassy dike, and beyond it the pond. On the west open grassland extended approximately 200 feet to the edge of the woods, with a diversion ditch at its border. On the south end, the rock pile was adjacent to woodland at the base of a steep slope with north exposure. On this slope the dense stand of second growth oak and hickory with an almost continuous leaf canopy was a poor habitat. The rock pile was thus partly isolated and surrounded by areas that were either uninhabitable to the skinks or supported only sparse populations of them. By 1948 the rock pile was partly covered by grape vines. Dead leaves and other debris had accumulated in the deeper interstices between the rocks. Spiders, beetles, snails and other small animals were extremely numerous in the vicinity of the rock pile and provided an abundant food supply. A large sycamore on the west side of the rocks provided some afternoon shade. This rock pile provided shelter for reptiles other than the five-lined skink—especially the garter snake, water snake, copperhead, and brown skink. Another area of about two and a fourth acres (“Skink Woods,” Figure 21) was the one most productive of skinks. It is a wooded upper slope adjacent to a hilltop pasture. Along the hilltop rim the upper stratum of the Oread limestone presents a rock face as much as four feet high at the north end, but less exposed at the south end where it was partly covered by deposited soil. Approximately 100 feet down the slope a second outcropping is present, with many loose rocks and boulders throughout the whole area. Soil is light and loamy. The slope has a west exposure. The stand of trees is fairly open, with several large elms, walnuts, and yellow oaks, and occasional hackberries, ailanthus and red haws. This area was included in a narrow strip of woodland fenced about 1940 as a runway connecting a hilltop pasture with a valley pasture where water was available at a time when both pastures were heavily grazed by horses and cattle. As a result of trampling, browsing and grazing by livestock, understory vegetation of this area presented a different aspect from that in most other parts of the woodland. Saplings of the dominant tree species and shrubs, notably dogwood, gooseberry and crabapple, were relatively scarce. Herbaceous vegetation, especially muhly grass, was conspicuous. By 1953 in the fifth growing season after livestock were removed, the area still contrasted with other parts of the woodland in sparseness of shrubby vegetation. Old stock trails were still discernible, and some sheet erosion and gullying had occurred. The effect of livestock in holding back woody undergrowth seemed to be an important factor in improving the habitat as the skinks were much scarcer in adjacent woodlands on either side that were similar in species composition, size, and numbers of the larger trees, but different in having much thicker underbrush. These adjacent woodlands were not entirely comparable, however, because they had more north-facing exposures. Reptile associates in the Skink Woods area include the brown skink, Sonoran skink, glass-snake, worm snake, ring-necked snake, blue-racer, garter snake, pilot blacksnake, copperhead and timber rattlesnake, but only the worm snake and ring-necked snake were abundant.

Rat Woods, an area of approximately four acres, was like Skink Woods, formerly the upper part of a connecting strip between hilltop and valley pastures and was altered by the effect of concentrated trampling and browsing by livestock. It is V-shaped, with the apex at the north end, and the slope exposures southwest and southeast. The area is bisected from north to south by a small gully, and remains of an old rock wall. To the east of this gully the lower outcrop is prominent but west of the gully, it is but little developed. As compared with other wooded areas, this one was relatively dry. Trees, and other vegetation in general, are somewhat more xeric in aspect than are those in Skink Woods. Along the upper ledge are elms and hackberries, with many thick clumps of fragrant sumac. The trees are mainly elm, walnut, honey locust, and osage orange with hardly any oaks or hickories and, with shrubby undergrowth of dogwood, gooseberry, and coralberry sparser than in adjacent woodlands. Herbaceous vegetation consists largely of muhly grass, geum, and avens. On the hilltop edge above the ledge are many flat rocks of varying sizes, and the slope is thickly strewn with rocks, some of the larger ones deeply embedded in the soil. The population of five-lined skinks was relatively sparser than in Skink Woods. Other reptiles including the Sonoran skink, racerunner, glass-snake, worm snake, ring-necked snake, blue-racer, bull snake, pilot blacksnake, garter snake, scarlet king snake, slender tantilla, and copperhead, were more numerous in this area than in most other parts of the Reservation. The comparatively scarce prairie skink was found only in this area, and the scarlet king snake and slender tantilla were found only here and at the quarry.

The Annual Cycle of Reproduction and Growth

Seasonal Occurrence

Collectors and other observers have often noted that reptiles, in general, are not found in equal abundance throughout the entire season of their activity. Many kinds are most in evidence within a period of weeks after emergence from hibernation, which corresponds with the time of breeding and later they become much scarcer. In skinks of the genus Eumeces this tendency is perhaps even more pronounced than in most other kinds of reptiles. By midsummer or considerably earlier their period of greatest activity is passed, and in some kinds, adults, or individuals of any size can rarely be found in the latter half of the growing season, even by a skilled collector familiar with their habitats and habits. Thus, Taylor (1936:5) in the preface of his revision of Eumeces, describing the difficulties involved in assembling needed series of the many Mexican species by collecting on summer field trips, wrote: "In 1934 in western Mexico … I met with most disheartening results … (although more than 1500 specimens were collected) only a single specimen of Eumeces was taken. Hobart Smith, in 1934, accompanied by David Dunkle, made a journey into northwestern Mexico … and while generally successful, likewise obtained only a single specimen of Eumeces."

In the present study the tendency of E. fasciatus to concentrate its surface activity in early spring was clearly shown. In unseasonably warm weather in early spring, even in February in one instance, individual skinks have been found active on the surface or beneath flat rocks warmed by the sun; but general emergence ordinarily does not occur until sometime in April, depending on the weather. Unless the weather is much warmer than the seasonal norm, the skinks spend much of April in a torpid condition, either not becoming fully active until late in the month, or lapsing into torpidity with the return of cool weather after their first emergence from hibernation. During warm periods in April, however, activity is at or near its annual maximum for all individuals regardless of sex or age.

In May, with the advent of much warmer weather, daytime temperatures are usually high enough for the skinks to be active. Adult males travel about more actively and persistently than females or young, and as a result they are found so much more frequently that the numbers taken approximate those for adult females and young combined. Many of the adult males recorded in May were taken in funnel traps or pitfalls. Active males in the open were difficult to catch, and a high percentage of them escaped. To the casual collector or observer, these skinks are much more in evidence in May than at any other time of year, and most of those seen are adult males. By June, the numbers of skinks seen in the open decline abruptly. The adult males become relatively scarce, with reduction from more than half to about one-sixth of the total, and the young, about half-grown at that season, make up approximately half of the total. The adult females make up approximately one-third of the total June sample, but few of them were found active on the ground surface. Most were found in nest burrows beneath flat rocks. Under such conditions they tended to be sluggish in behavior, and were caught much more easily than were males and young. July was characterized by progressive decrease in the numbers of adult males, adult females, and second year young, whereby the numbers of each group were little more than half of those for June; and by appearance of a new crop of hatchlings which made up about one-third of the month’s sample. Hatchlings first appeared from early July to late July in different years; few were recorded in July in some years. Females were much less commonly found in nests in July than in June because many nesting attempts were terminated before the beginning of July or early in the month, and probably because those that remained were often more deeply buried and better concealed. By August the adult males, and the second year young (by then approaching adult size) were found in still smaller numbers, but the number of hatchlings and of adult females approximated those recorded in July. In the females there is evidently some resumption of activity after the incubation period is terminated. The females are then hungry and sometimes emaciated, weighing less, on the average, than the year-old young of shorter snout-vent length. The numbers of hatchlings are augmented through early August in some years, as late broods continue to hatch. By early September few skinks except hatchlings are to be found, and activity continues to wane throughout the month. In October skinks of any age or sex group are a rarity, even though temperature is about the optimum for their activity. Little is known concerning where and how they spend the fall months. Probably they are not actually dormant, but retreat underground where temperature is moderate and humidity is high. Individuals kept in captivity at this season were listless showing but little inclination to feed. The only five-lined skink taken on the Reservation in November was found in a funnel trap after a rain at the end of a long drought. It may have been attracted to the surface by moisture.

The following table shows the dates on which various events of the annual cycle were observed in each of five different years. Owing, to the secretive habits of the skinks, these events generally were not observed until somewhat after their earliest occurrence in any one season. The lag was greater in some instances than in others.

Table 3. Phenology of the Annual Cycle in Five Different Years.

Reynolds (1943:370 and 1947:191) studied the histological and gross seasonal changes in the reproductive organs of the adult male Eumeces fasciatus. There is a well defined annual cycle. "Early seasonal increase in seminiferous epithelial heights and in diameter of lumina and tubules reached a maximum in April followed by regression reaching complete involution by August. Late seasonal revival of activity results, by November, in size of testicular elements comparable to those seen in January. Primary spermatocytes predominate in the germinal epithelium in January, secondary spermatocytes and spermatids in February, with spermatids and metamorphosing sperm dominating from March until late June when the germinal material of the current season is exhausted." Fifty-three adult males were used as a basis for his study. These were of diverse origins from Arkansas, Florida, Missouri, Tennessee, and Indiana. Since sexual cycles in such widely ranging species tend to be synchronized with local phenology, and change somewhat from one region to another, the seasonal cycle may have been somewhat obscured by the diverse origins of the material. The Florida specimens may have been of the species E. inexpectatus. Apparently Reynolds’ experimental skinks were kept in captivity for varying lengths of time before their reproductive organs were examined. The normal cycle would almost certainly be altered in captivity, especially in skinks kept at high temperatures during the time that they would normally be hibernating.

The seasonal change in gross appearance of the testes is not great. In the breeding season the testes are slightly enlarged and are firm and engorged, with pinkish or orange tinge. In immature males, and adults that are not in breeding condition, the testes are smaller, attenuate, paler colored, and flaccid. Sizes of testes in some males killed in the breeding season are recorded in Table 4.

Table 4. Sizes of Testes in Spring and Early Summer in Sexually Mature and Juvenal Males.

From the time of emergence in spring, males show some tendency to seek out females, and frequently a pair may be found together under the same rock, weeks before the onset of the breeding season. There is no satisfactory evidence that such associations have any permanence. At the time of emergence from hibernation the males rarely have even a trace of reddish coloration on their heads, and more than a month normally elapses before attainment of breeding coloration. Each year that observations were made activity of the skinks was interrupted by cold weather in April, so that the lizards were fully active for only part of the time between their earliest emergence and their attainment of breeding condition five to eight weeks later. The reddish suffusion of the breeding season, hardly showing in the first few weeks after emergence, appears suddenly within a few days in all adult males of the population. The best indication of the time necessary to attain breeding condition was provided by an adult male whose hibernation was interrupted on December 15 by bringing him into a warm room where he was kept at 80° F. or more in the daytime, and approximately 70° F. at night. Thirteen days later, on December 28, the male had developed a noticeable reddish suffusion. On January 3, nineteen days after hibernation terminated, the suffusion was near its maximum. When an adult female was placed with the male on this date, he showed sexual interest but the courtship was not consummated. On January 6, the 22nd day, the male’s colors had reached their maximum, and when the female was placed with him, pursuit and copulation occurred promptly.

In the spring of 1952, the first skink of the season was found on March 29, still only partly activated, and under a large flat rock. Skinks were not caught or seen in any numbers until April 17, however, and general emergence probably occurred only a day or two earlier than this. On May 10, 1952, breeding activity was estimated to be at its peak. By May 28, the reddish suffusion was conspicuously faded in several males taken. By June 10 it was no longer discernible.

In the immature female the oviducts are small and threadlike, and the ovaries have grapelike clusters of pale whitish eggs, which are minute, often less than .5 mm. in diameter (Figure 8A). In sexually mature females ova enlarge rapidly after emergence from hibernation in the spring. While eggs are still in the ovary, they are approximately spherical. In late April and early May the developing ova enlarge rapidly. Approximate average sizes (dimensions in mm.) of developing ovarian ova in each of 22 mature females on different dates were as follows: April 17, 1949: 2.6, 2.3, 2.2, 2.2, 1.9, 1.9; April 18, 1949: 2.2, 1.9, 1.8, 1.1, 1.1; April 24, 1949: 4.6, 3.2, 2.5, 2.3; May 6, 1951: 2.5, 2.3; May 20, 1951: 7.0, 6.2; May 25, 1951: 8.0; June 3, 1951: 6.0, 5.5.

The two females containing ovarian eggs on June 3, 1951, were retarded individuals, taken along with several others that had already ovulated. Copulation takes place in early May before the ova have grown to their full size. In the following weeks both the ova and the oviducts enlarge rapidly. Upon passing into the oviducts, the ova assume an oval shape and are approximately 9 by 6 mm. before the albumen and shell are added. Deposition of a clutch of eggs probably extends over only a day or two at most, as clutches appear abruptly in the nest cavities. On only a few occasions were the females found in nest cavities with their clutches partly laid.