Life History and Ecology of the Five-lined Skink, Eumeces fasciatus Part 9

Movements

Data obtained concerning the movements of these skinks demonstrated that individuals tend to limit their activities to small areas thoroughly familiar to them, and wander but little. Although the nature and extent of movements in reptiles in general, and in lizards especially, are poorly known, my findings are perhaps what might be expected from the studies of earlier workers on various other species of reptiles.

Goin and Goin (1951:29) observed that _Eumeces laticeps_ in Florida lives in hollow stumps, each individual excluding other adults from its stump but tolerating young. Movements have not been studied in detail in any member of the Scincidae, however. The observations of Goin and Goin, and those of other authors, seem to indicate that _E. laticeps_ is territorial, and that each individual centers its activities about a tree or snag, regularly using the same hollow as a shelter and home base. In contrast, _E. fasciatus_ is not territorial and has no regular home base.

The iguanid genus _Sceloporus_ is perhaps better known than any other kind of lizard as regards its movements. Studies by Newman and Patterson (1909), Stebbins and Robinson (1946), and Fitch (1940) on three different species have shown that individuals of _Sceloporus_ keep to small individual areas, and that territoriality is well developed, in some species at least.

Among other reptiles, turtles are much better known, as detailed studies of movements have been made on several species, of which the life histories and ecology have been thoroughly investigated (Nichols, 1939; Cagle, 1942 and 1944; Woodbury and Hardy, 1948; Stickel, 1950). They have been found to have well-defined and fairly extensive home ranges, which are not defended as territories. Studies of movements in several different kinds of snakes, by Blanchard and Finster (1933), Stickel and Cope (1947), Fitch (1949), Lowe and Norris (1950), and Carpenter (1952) have shown that these reptiles usually have definite home ranges, which may be several or many acres in extent. Their home ranges are not defended as territories against other members of the species. In general, turtles and snakes have been found to occupy home ranges that are much larger than those of lizards.

Most information concerning movements of _Eumeces fasciatus_ has been obtained from the recapture of marked individuals. Actual distances of travel, and the time, frequency and motivation of movement was uncertain. A skink marked, recorded, and subsequently recaptured at a second location may have wandered widely in the meantime, visiting points relatively remote from either location of capture. The two points of capture may be within a home range regularly or occasionally covered by the individual in the course of its routine activities; or the second point may have been recorded only after a permanent shift of activities away from the area within which the original point was located. Various types of movements probably were involved.

Interpretation of the records is difficult because of the paucity of direct observations on the behavior and movements of skinks under natural conditions. Often when one is alarmed, it will run as much as 30 feet, in a fairly direct course, to a tree or bush or rock where it can find refuge. Undisturbed individuals move about slowly and circuitously.

It is difficult to keep one under observation for any length of time because of the secretive habits causing it to keep under cover, as much as possible while moving about, and to hide in response to any slight disturbance.

It is obvious that individuals shift their activities from time to time, occupying new areas either abruptly or by gradual stages. Even though a successful skink has a life span of several or many years, the populations on the small study areas were found to be much altered from one year to the next. Presumably this change was brought about largely by shifts in home ranges. Several shifts of hundreds of feet were recorded, but the chances of recovering marked individuals that moved so far were relatively poor because their movements generally took them beyond the limits of the study area to locations where recapture was unlikely. Skinks often were caught at their hiding places beneath rocks or other sheltering objects. In many of these instances it was evident from the position, temperature and state of activity of the lizard that it had been in the open but had become alarmed as the collector drew near and had retreated unnoticed to its shelter just before capture, whereas in other instances it was obviously at rest in its chosen shelter. Except for females in their nest burrows individuals were not ordinarily recaptured regularly at the same hiding places. They may seek new hiding places after each period of activity.

However many of the skinks captured were taken again, after long intervals, near the same places. Time elapsed between successive captures for different individuals ranged from one day to 47 months. Of the total of 323 recaptured by September, 1952, approximately half, 162, were taken after intervals including one or more hibernation periods. In appraising home ranges and detecting the occasional shifts over a relatively long time span, chronology of the records needs to be taken into account. Records cl.u.s.tering about the same center seem to indicate continued occupancy of an established home range. However, when one or more early records are well separated from one or more later records, a shift in range seems probable. In some instances successive records were progressively farther from the starting point suggesting two or more shifts in the same direction from an original home range.

Although recorded movements varied from a few inches to hundreds of yards, the most noteworthy feature in general was the short distance between points of capture (considered in relation to the potential mobility of the lizards) after days, weeks, months or years. In many instances no movement was demonstrable, even though successive points of capture were not exactly the same. Named natural landmarks, mostly trees, boulders and logs, well distributed over the study area, were used as a basis for locating points on the map. Direction and distance in feet to the nearest landmark was recorded for each site of capture, but for distances of more than 25 feet estimates were made to the nearest ten feet. Usually at least one landmark was available within a 50-foot radius from any point where a capture was made. Occasional estimates made for distances of more than 50 feet, or even more than 100 feet, in the absence of suitable landmarks nearby, were sources of inaccuracy. For such estimates errors of up to ten feet were common, and some errors of greater magnitude were made.

For most individuals successive sites of capture tended to cl.u.s.ter within a small area, but the occasional outlying capture sites indicate that each individual does range outside the area in which its activities are concentrated. These occasional excursions cannot be consistently attributed to any one ecologic requirement, nor are they limited to any particular time within the season of activity. Adult males, however, tend to make longer movements in the brief period of concentrated s.e.xual activity, thereby increasing their chances of finding mates. Similarly, adult females may wander beyond their usual ranges in search of suitable nesting sites. The home range may be thought of as consisting of a small central portion where activities are largely concentrated, and an outer area several times as large, familiar to the animal but used to a lesser extent by it. The activities gradually become more diffuse farther from the central part of the home range. In the five-lined skink, home ranges are unlikely to approximate the circular shape because they are molded with respect to environmental features that are not uniformly distributed. A rotting log, an old tree with decayed hollow base and nearby fallen slabs of bark and dead limbs, a rock outcrop with numerous deep holes and crevices, or a group of flat rocks in a forest glade fulfill requirements not met in the surrounding habitat with the result that home ranges are built around them. Consequently a home range may be long and narrow, with maximum diameter several times the minimum diameter.

The usual concept of home range, as a finite area with well defined boundaries is not entirely satisfactory for an animal with the habits of the five-lined skink. The skink spends much of its time in inactivity underground or otherwise concealed and sheltered, and when it does move about it takes advantage of natural travel-ways over rock surfaces, tree trunks, and logs. If a log happens to be the home range center, the skink may travel the length of the log many times without making a comparable trip at right angles to this axis of travel, although it may make short side dashes to secure food. On more extended forays, the directional sequence of movements is largely controlled by the distribution of suitable cover and travel routes, as the skink avoids both open areas and dense vegetation. Outlying portions of the home range probably are not uniformly covered but are reached only occasionally as the lizard is led along some natural travel route, or after it has visited, in succession, a series of locations attractive in providing shelter or food.

Marked skinks were recaptured at distances up to 680 feet from points of original capture. Considering only the most remote points of capture for those individuals recaptured more than once, the average recorded movement for the entire group of 323 recaptured skinks was 58 feet. This figure provides a basis for comparing vagility of this species with others. Eliminating some individuals of indefinite status, the average movement for 75 adult males was 69 feet; for 102 adult females, 45 feet; and for 112 young, 61 feet. For the adult females, home range data are biased by the fact that many were caught repeatedly at or near their nests. It is not clear whether females that do not have nests range less widely than males.

Only 15 individuals, less than five per cent, had moved more than 250 feet. These longest movements were: 680 feet, adult female, 26 months; 680 feet, adult female, 10 months; 680 feet, subadult male, one year; 650 feet, young to adult male, 22 months; 640 feet, subadult to adult female, two years; 535 feet, young male, 11 months; 510 feet, adult male, 11 months; 490 feet, young (s.e.x undetermined), 10 months; 450 feet, young male, 13 months; 350 feet, young (s.e.x undetermined), 10-1/2 months; 335 feet, adult female 13-1/2 months; 275 feet, adult male, 35 months; 275 feet, adult male, 24 months; 270 feet, young to adult male, 12-1/2 months.

For those skinks caught on only two occasions, at different places, the single movement record provides some clue as to the location and size of the home range. No evidence was obtained to indicate that the activities of these lizards center at fixed home bases. It may be a.s.sumed that any two successive captures of the same individual separated by a substantial time interval, will be distributed at random to each other within the area to which the animal's activities are confined. The varied techniques of capture, by hand and with different types of traps, would help to secure random distribution of capture sites. If the home range were covered uniformly by the animal in the course of its activities, any two random capture sites would be on the average separated by a distance equal to half the home range diameter.

If the animal tends to concentrate its activities in the central part of the home range, as seems to be the case, the capture sites will be correspondingly closer together. For the 196 skinks that were caught on only two occasions, average movement was 62 feet. Within this group the 42 adult males that were recaptured only once had averaged movements of 58 feet. One had made an exceptionally long movement of 510 feet, which obviously was not entirely within its home range. Excluding this one long movement, the remaining 41 had moved on the average, approximately 47 feet (Table 15). Among the other skinks caught only twice one of 61 females and 8 of 93 young had likewise made such long shifts that it seemed inadvisable to include them in computing the size of the home range.

Distance between points of capture showed little correlation with elapsed time. For 24 of the adult males that were recaptured in the same year they were originally marked, the average distance was 49 feet, whereas in the 17 others recaptured after one or more hibernations the average movement was 45 feet. For adult females, the corresponding figures were, respectively, 22 feet and 29 feet; and for young, 33 feet and 66 feet.

For those individuals recaptured twice, at different locations, the three points of capture show to a greater or lesser degree the position, and, in part, the extent of the home range. Of course, all three points may be concentrated near the center of the home range, or they all may be scattered along its edges. In general, however, each point will lie somewhere between the center and edge of the home range, separated from each of the other two points by a distance of, on the average, approximately a home range radius.

Table 15 shows that adult males and young tend to range more widely than adult females, and that young tend to shift to new areas more frequently than do adults. Many of the recorded movements (in addition to the long ones that were excluded from the home range computations) may have involved short shifts in ranges. If all such shifts could be definitely identified and eliminated from the computations, actual home ranges might be considerably smaller than those indicated by the present set of data. Home ranges approximately 90 feet across for adult males and young, and a little more than 30 feet across for females are indicated.

Actual area of a home range would amount to only a fraction of an acre--from about one-seventh to less than one-fiftieth. The dash of an alarmed skink to a place of refuge, though involving at most only a few seconds, may traverse a large part of its home range. Through long a.s.sociation the lizard is thoroughly familiar with the terrain, so that it can take full advantage of the peculiar features in escaping, hunting, traveling or resting.

Table 15. Distances Between Successive Sites of Capture for Marked Five-lined Skinks on Study Areas, Indicating Home Range Sizes.

=======================+============+==========+=========== Number Average of skinks maximum discarded distance Number from in feet of sample between skinks because of Age, s.e.x and Number points included relatively of Captures of in long capture, sample movements, and indicative extremes of shifts of range -----------------------+------------+----------+----------- Adult males Individuals captured just twice 47 (225-0) 41 1 Individuals captured just three times 47 (130-0) 18 0 Individuals captured four or more times 91 (200-0) 17 2 Adult females Individuals captured just twice 16 (90-0) 56 4 Individuals captured just three times 25 (90-0) 25 3 Individuals captured four or more times 28 (90-0) 15 1 Young Individuals captured just twice 45 (160-0) 85 8 Individuals captured just three times 46 (150-0) 14 0 Individuals captured four or more times 82 (175-0) 14 2 -----------------------+------------+----------+-----------

Relatively few marked individuals were caught four or more times at different sites. For these individuals listed below the distribution of the sites is more or less indicative of shape and size of the home range in some instances. For some of them successive locations of capture are shown and possible home ranges are outlined in Figures 21-25.

[Ill.u.s.tration: FIG. 21. Map of Skink Woods study-area, showing chief physiographic features and landmarks, and showing also successive sites and dates of capture of a marked male skink and two marked females, suggesting extent of home ranges.]

ADULT MALES

No. 1: Seven captures in two years, on May 13, 1950, May 12, 1951, and in 1952 on April 28, May 1, 2, 4 and 6, these seven locations well distributed over a stretch of rocky slope 275 feet in greatest diameter.

The fifth location was only 20 feet from the original, whereas the last, only four days later, was the most remote, suggesting that the whole area covered may have been within a home range.

No. 2: Seven captures in 46 months, skink not fully grown when first captured on June 22, 1949; 275 feet south on May 4, 1950; had moved from this second location 150 feet west northwest on June 17, 1950, and this third location together with the last four, on May 15, 1951, and May 13 and 15, 1952, and April 6, 1953, were all within a 20 foot diameter.

Evidently two shifts in range were involved.

[Ill.u.s.tration: FIG. 22. Sites of successive captures of two marked adult males in the Skink Woods study-area.]

No. 3: Six captures, all at different locations, in 22 months, on July 5 and 28, 1950, May 3 and 23, and June 21, 1951, and May 1, 1952. The 190-foot-wide area was probably all within a home range, as the fourth and fifth sites were those most remote from each other.

No. 4: Six captures in 21 months, in 1950 on August 14 and September 3, in 1951 on April 27 and August 21, and in 1952, on May 28 and 30. The four 1950 and 1951 locations were within a 30-foot diameter, whereas the two 1952 locations were 150 feet farther east, and even nearer together, suggesting a shift in range.

No. 5: Five captures in five months, all within a 40-foot diameter, on April 24, May 7 and 28, June 14, and September 22, 1951. The first and third locations were at almost the same spot.

No. 6: Five captures all at different locations, in 23 months; in 1950 on July 27, in 1951 on April 30 and May 25, and in 1952, on May 1 and June 28. The second, third and fourth locations were all within 45 feet of each other and of the first, but the last was 110 feet from the first, possibly representing a shift.

[Ill.u.s.tration: FIG. 23. Sites of successive captures of three marked adult males in the Skink Woods study-area.]

No. 7: Four captures in two months, at approximately the same place on May 1 and 5, 1950; on May 30 had moved 35 feet farther north along ledge, and on July 1, 25 feet farther in the same direction.

No. 8: Four captures in one year, all at approximately the same place along rock ledge, on June 17, 1949, and April 21, May 3 and June 15, 1950; trapped three times and once caught by hand.

No. 9: Four captures in one year, on April 7 and 11, and July 27, 1950, and April 14, 1951, the four different locations all within a 30-foot diameter.

No. 10: Four captures in 22 months, in 1950 on July 7, and again on July 23, 175 feet farther north; on May 25, 1951, 200 feet east of second location, and on May 2, 1952, 30 feet from third location. At least one shift in range probably occurred, from 1950 to 1951.

No. 11: Four captures in 36 days, in 1951 on April 30, May 8 and 15, and June 5. The last two captures were made in the same trap and were only 15 feet from the original location, but the second location was 130 feet from both. Because the time span was short and the lizard returned from the most remote point, it seems probable that all four records were within its home range.

[Ill.u.s.tration: FIG. 24. Sites of successive captures of marked skinks, a male and two females, in the Skink Woods study-area.]

No. 12: Four captures in 11 months, all within a 50-foot diameter, in 1951, on June 1 and 26, and August 27, and in 1952, on April 29.

No. 13: Four captures in 15 days, all in July 1949 within a 10-foot diameter.

No. 14: Four captures in 22 months, July 22, 1950 (as subadult), in 1951, on May 8 and June 5, and on May 13, 1952. Second location 295 feet southwest of first, third 30 feet north of second, and fourth 650 feet east of second and third. Probably two shifts of range were involved.

ADULT FEMALES

No. 1: Six captures in 26 months; in 1950 at the same place on June 4 and 13, in 1951 on May 26 it had moved from the original quarry ledge location 680 feet south southeast down the slope to the pond rock pile, where recaptured on June 9, and in 1952 on May 21 and July 22.

No. 2: Six captures at four locations all within a 25-foot diameter, in 13 months; June 5, 1950, and May 25, June 18, 26 and 29, 1952. On each occasion this female was hiding in a nest burrow, but she shifted to new nest sites as a result of disturbance by the investigator or flooding when there were unusually heavy rains.

[Ill.u.s.tration: FIG. 25. Sites of successive captures of a marked male and a marked female, each taken in three different years in the Skink Woods study-area.]

No. 3: Five captures in 34 months, all within a radius of a few yards, at the pond rock pile, on August 8, 1949, June 5 and July 23, 1951, and May 15 and June 4, 1952.

No. 4: Four captures in 34 months, all within a radius of a few yards at the pond rock pile, on August 8, 1949, June 7, 1950, May 30, 1951 and May 21, 1952. It is notable that this female was taken only once in each of four different years, her occupancy of this rock pile seemingly continuing throughout the duration of the study.