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1-1-2008 A study on the population size and natural history of the Eastern Six‐lined Racerunner, sexlineata, in West Virginia, with notes on other Emmy Johnson

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Recommended Citation Johnson, Emmy, "A study on the population size and natural history of the Eastern Six‐lined Racerunner, Aspidoscelis sexlineata, in West Virginia, with notes on other lizard species" (2008). Theses, Dissertations and Capstones. Paper 674.

This Thesis is brought to you for free and open access by Marshall Digital Scholar. It has been accepted for inclusion in Theses, Dissertations and Capstones by an authorized administrator of Marshall Digital Scholar. For more information, please contact [email protected]. A study on the population size and natural history of the Eastern Six‐lined Racerunner, Aspidoscelis sexlineata, in West Virginia, with notes on other lizard species.

Thesis submitted to the Graduate College of Marshall University

Partial fulfillment of the requirements for the degree of Master of Science in Biology

By

Emmy Johnson

Dr. Thomas Pauley, Ph.D., Committee Chairperson Dr. Dan Evans, Ph.D. Dr. Jeffrey May, Ph.D.

Marshall University

May 2008

Keywords: Lizard activity patterns, Range expansion, Natural history, Distribution, Behavior

ii

ABSTRACT

A Study on the population size and natural history of the Eastern Six‐lined Racerunner, Aspidoscelis sexlineata, in West Virginia, with notes on other lizard species.

By Emmy Johnson

Species will expand ranges through natural or human‐alterations. Such an occurrence has happened in West Virginia with Eastern Six‐lined Racerunner, Aspidsocelis sexlineata, a species that has been found to thrive in areas of high disturbance from humans. While found abundantly in other states, they are thought to occur only in the eastern panhandle of West Virginia (Morgan County) where they are believed to have come from Maryland via a railroad bridge. Without the bridge, the Potomac River forms a natural barrier between these two states. The objectives of my study were to determine the population size and natural history of these . My hypotheses were that these lizards came from Maryland and are well established in a small area of West Virginia. Forty three A. sexlineata were captured over the summer of 2007, including 11 recaptures. Aspidoscelis sexlineata were found to have similar seasonal activity patterns as lizards in other climate regions. They were observed to have a short daily activity pattern of 1100h to 1500h. Gravid females were captured during May, June, and July. A population estimate of 70.2 individuals was found for a closed system and 171 individuals were found for an open system. Both of these estimates indicate a healthy population that is self‐reproducing and could occupy this area for a significant period of time. Aspidoscelis sexlineata were found farther down the railroad than previously recorded; this may indicate an expansion of range. Potential competitors of A. sexlineata are Eumeces fasciatus, E. anthracinus, E. laticeps, Scincella lateralis, and Sceloporus undulatus. Surveys were conducted to find populations of these other lizards. Lizards were found in Cabell, Kanawha, Nicholas, Grant, and Morgan Counties. They were not found in Wayne, Fayette, Pocahontas, and Tucker Counties. More studies need to be conducted on the lizards of West Virginia, because little is known about most of these species. iii

Acknowledgements

I would first like to thank Dr. Pauley for giving me this opportunity to further my education and gain many life experiences along the way. I am glad Dr. Pauley takes time to take his graduate students out in the field with him. It is a very rewarding to have one on one

experience.

My sister, Erin, has been there for me during my frustration of capturing nothing and making me take a look at what I was doing and how crazy it sounds to most people. She also gave me a place to take shelter and rest when I thought I wasn’t going to be able to make it through the summer. I would also like to thank her for always introducing me to her friends as someone who cuts the toes off of lizards, with no other explanation.

I would like to thank Matthew Graham for encouraging me to do this project and also

being really good at convincing me to do things I am not prepared to do (like go on a jump when I had only been snowboarding twice before).

I would like to thank Conor Keitzer for helping me while out in the field, collecting lizards, enduring the summer heat with very little shade, and withstanding the railroad dirt which is impossible to clean off you. He helped me with forming an idea of how to make my traps and fence moveable. He also helped build some traps which are a long and painful task. I

would also like to thank him for reading over my thesis, multiple times, and talking over my ideas with me or rather just listening to me talk about my thesis all the time. Without his help, my thesis would not be what it is.

iv

Table of Contents

Page #

Abstract………………………………………………………………………………………………………..……………………………ii

Acknowledgments…………………………………………………………………………..………………………………………..iii

List of Tables……………………………………………………………………………………..……………………………………….v

List of Figures…………………………………………………………………………………………………………………………….vi

Part 1: Study on Aspidoscelis sexlineata

Chapter 1: Overview………………………………………………………………………………………………………….………1

Chapter 2: Introduction………………………………………………………………………………………………….…..……..4

Chapter 3: Methods…………………………………………………………………………………………………………………12

Chapter 4: Results………………………………………………………………………………………………………….………..17

Chapter 5: Discussion……………………………………………………………………………………………………………….24

Chapter 6: Conclusions………………………………………………………………………………………..…………………..34

Part 2: Notes on other lizard species in West Virginia

Chapter 7: Overview………………………………………………………………………………..………………………………36

Chapter 8: Introduction……………………………………………………………………………..…………………………….39

Chapter 9: Methods…………………………………………………………………………………..…………………………….50

Chapter 10: Results………………………………………………………………………………………………………………….51

Chapter 11: Discussion………………………………………………………………………….…………………………………55

Chapter 12: Conclusions…………………………………………………………………………………………………………..57

Literature Cited………………………………………………………………………………………………..………………………58

v

List of Tables

Table 1. A. sexlineata searches in the state of West Virginia……………………………………………….……11

Table 2. Locations where lizards were found during surveys for the summer of 2007……….…….52

Table 3. Locations searched during the summer of 2007, where no lizards were seen………..…..53

vi

List of Figures

Figure 1. Six dorsal stripes against a black background found on A. sexlineata……………...... 7

Figure 2. Female A. sexlineata with a white/cream ventral side…………………………………………………7

Figure 3. Male A. sexlineata with blue tint on ventral side, used for sex identification………...... 8

Figure 4. Shale barren in Maryland where A. sexlineata were captured…………………………………….8

Figure 5. Railroad in Morgan County where A. sexlineata have been found……………………………….9

Figure 6. Shale barren locations in West Virginia……………..………………………………………………………10

Figure 7. Trapping site along the railroad with drift fences set up……………………………………………15

Figure 8. Side view of funnel trap, placed against drift fence……………………………………………………15

Figure 9. Funnel design…………………………………………………………………………………………………………….16

Figure 10. Funnel trap design with measurements…………………………….…………………………………….16

Figure 11. Clipping the toe of S. undulatus to give it a specific identification number……………...16

Figure 12. Gravid A. sexlineata…………………………………………………………………………………………………19

Figure 13. The relationship between soil temperature and time of day………………………..…………20

Figure 14. The relationship between soil temperature and month………….………………………………20

Figure 15. The relationship between air temperatures for June and July………………………………….21

Figure 16. The relationship between air temperatures for June and July and time of day…….….21

Figure 17. Average air and soil temperatures for June……………………………………………………………..22

Figure 18. Average air and soil temperatures for July………………………………………………………………22

Figure 19. The relationship between number of A. sexlineata caught and month……….….……….23

Figure 20. The relationship between number of A. sexlineata caught and time of day….………..23

Figure 21. Average July temperatures for the summer of 2007 for the United States…………………………………………………………………………………………………………………………..31 vii

Figure 22. The relationship between soil temperature and number of A. sexlineata captured

and time of day…………………………………………………………………………………………….………………31

Figure 23. Google map of the Potomac River and CSX Railroad between Maryland and West Virginia…………………………………………………………………………………………………………………..…….32

Figure 24. Heterodon platirhinos caught near trapping area….………………………………...... 32

Figure 25. Re‐growth of A. sexlineata tail…………………………………………………………………………………33

Figure 26. Aspidoscelis sexlineata taking refuge in a burrow..………………………………………………….33

Figure 27. Known locations of S. undualatus from the West Virginia Biological Survey…………….43

Figure 28. Sceloporus undulatus with a gray body and keeled scales.……………………………….……..44

Figure 29. Known locations of E. fasciatus from the West Virginia Biological Survey………………..45

Figure 30. Adult E. fasciatus with orange head and olive brown body, caught in trap at the railroad………………………………………………………………………………………………………………………..46

Figure 31. Known locations of S. lateralis from the West Virginia Biological Survey…………………47

Figure 32. Known locations of E. laticeps from the West Virginia Biological Survey………………….48

Figure 33. Known locations of E. a. anthracinus from the West Virginia Biological Survey……….49

Figure 34. West Virginia counties where A. sexlineata were found and not found during the summer of 2007……………………………………………………………………………………………………………54

PART 1 CHAPTER 1 OVERVIEW

Species have been known to expand their ranges due to human development.

Construction of new roads, bridges, and railroads are a few examples of human development that aid species expansion. Not only can human development allow ranges to expand, but it can also alter .

When railroads are built, forests and mountain sides are opened up by cutting down trees, thus changing the amount of sunlight that reaches the soil. They also change soil content by grease and railroad debris that are deposited along the tracks. Development, such as railroads and bridges, opens up new areas for species to expand into, create corridors for species to disperse, and cross natural barriers.

Roads change habitats by altering soil density, temperature, moisture content, light and dust levels, surface waters, runoff, sedimentation, and heavy metal content (Trombulak and

Frissell 2000). They alter an ’s behavior by putting stress on its reproduction, escape responses, and physiological state. Roads cause mortalities from construction and animal collisions with vehicles. They alter the chemical environment, aid in the spread of exotics, and increase human use of the area (Trombulak and Frissell 2000).

Introduction through pet trade and “stowaways” are also unnatural ways species can expand their ranges. The curly tailed lizard, Leiocephalus carinatus armouri, in Florida is an example of a species that expanded its range due to human alteration. L.carinatus is an introduced species native to the islands of the Little Bahama Bank. It was first reported in 1958

1 in Florida and its introduction was attributed to 20 pairs being released in the 1940’s. This lizard occupies habitats that have been degraded by human development, such as buildings, pavement, and parking lots. It has followed the urban and suburban development of Florida

(Smith et al. 2004). This species was first introduced by humans and with the aid of human development; its range has expanded 46.3 km south and 34.1 km north.

As previously stated, human development can facilitate an animal’s expansion into an area previously blocked by natural barriers. Nine‐banded armadillo, Dasypus novemcinctus, has expanded its range as a result of human influence. In the past, the Rio Grande and fire‐ maintained grasslands were natural barriers that inhibited this animal’s range expansion. In the

1850’s, southern Texas was settled and roads and bridges were constructed. As human transportation increased along roads and bridges, armadillo’s range increased as well.

Armadillos were also found on‐board railroad shipments. Humphrey (1974) reported that since

1954, the range had expanded northward and eastward in Texas, but had contracted in western

Texas. He also reported that in Florida the range was expanding in all directions. The species’ range has expanded 4.3km yearly north and east since Humphrey’s (1974) report, although he expected both the northward and western expansion to decelerate. Some natural processes can continue to limit a species from expanding, such as precipitation and climate. In the case of the armadillo, freezing temperatures limit the availability of soil invertebrates, which limits the amount of fat stored by the armadillo, and therefore their ability to survive starvation periods

(Taulman and Robbins 1996). Armadillos utilize habitats with a vegetative litter layer, porous soil, and varied faunal assemblages, precipitation is needed to maintain these characteristics and so it is limiting their range expansion.

2 The Eastern Six‐lined Racerunner, Aspidoscelis sexlineata, have been shown to benefit from human alterations to the environment. Mitchell (1994) found them in open road embankments in the Piedmont Valley and along railroad cuts in the Shenandoah Valley,

Virginia. Rhoads (1895) also found them along railroad embankments in the suburbs of

Chattanooga in Kansas and Burt (1928) found them along the Union Pacific railroad near the

Kansas River. It is obvious this species has been able to expand its range across natural barriers because of human development. They are also able to adapt to changes from agricultural conditions (Burt 1928). Baker and Stebbins (1965) stated that if a species is going to establish in an area they need to be able to adapt to unfavorable conditions, adapt to new environments, have high reproductive capability, and the ability to disperse rapidly. Aspidoscelis sexlineata are able to survive in unfavorable conditions, are adaptable, and can disperse rapidly, so

according to Baker and Stebbins (1965), this species would be able to establish in new areas.

3 CHAPTER 2 INTRODUCTION

Aspidoscelis sexlineata (also called sand lappers and field streaks), have six light stripes which range in color from white to blue on a black background (Figure 1). Their dorsal scales are tiny and granular while their ventral scales are large and rectangular. The tail scales are larger and distinct from the dorsal scales. Aspidoscelis sexlineata have dull scales which distinguish them from , which have glossy scales. The ventral surface of females is a cream color (Figure 2) and males have a bluish‐tint (Figure 3) (Conant and Collins 1998, Mitchell

1994). Aspidoscelis sexlineata are known to have two clutches of eggs during their active season and mating occurs during June and July, with June being the predominant month for breeding (Hardy 1962, Clark 1976).

Aspidoscelis sexlineata are found from the southern mid‐western states to the east coast and are the only species of this found east of the Mississippi River. In West

Virginia, A. sexlineata are ranked an S1, which means they are rare or imperiled. Since these

lizards are found in one area of West Virginia, this means they have a higher chance of being extirpated from the state.

Aspidoscelis sexlineata are fast‐moving, energetic lizards. They are found in various habitats such as rock outcroppings, old fields, and dry open areas because these areas have

sufficient sunlight to maintain optimal temperatures for activity (Conant and Collins 1998). The need to maintain a certain body temperature limits the lizards’ activity to the warmest months of the year.

4 Aspidoscelis sexlineata have a unique activity pattern and have been studied in many states, except West Virginia. They have a short daily active period with an early afternoon retreat. Many studies have tried to address the question of why they have an early retreat.

Winne and Keck (2004) manipulated possible influences on daily activity such as prey, water availability, and temperatures. They found that only morning soil temperatures affect activity and concluded that these unique activity patterns are due to a circadian clock. Hardy (1962) hypothesized that the reason for early retreat was because lizards had gained enough fat reserves for hibernation. She concluded that as the summer progresses, fat storage increases and daily activity decreases. Milstead (1957) attributed their inactivity to the high afternoon temperatures, concluding that as temperature increases lizards would return underground.

Rose (1981) states that territoriality demands daily activity. Since A. sexlineata are not territorial, they expend less energy establishing a territory and therefore are not as active as territorial species. Lack of territoriality may be a reason for reduced daily activities.

Inactivity can also be part of a lizards’ adaptive strategy. Rose (1981) states that since periods of inactivity occur frequently, activity must have some negative aspects and that there

are some benefits to inactivity. These include lower risk of , less energy loss when food is scarce and decreased dehydration during dry periods. An active Sceloporus undulatus,

Fence Lizard, has been found to have a 10 times higher risk of mortality than an inactive lizard

(Adolph and Porter 1993). It has been shown that Sceloporus virgatus, Striped Plateau Lizard,

can be inactive for a six‐day period and its amount of energy used would be equal to that of one active day. Grant and Dunham (1988) found Sceloporus merriami, Canyon Lizard, to have activity patterns and microclimate use that correlated with daily temperature patterns. They

5 stated that limited availability of operative temperatures, temperatures at which lizards are active, could be a factor in reducing daily activity.

In Maryland, A. sexlineata are found on shale barrens which are hot dry areas with low vegetation. Shale barrens have desert‐like conditions and many plant species found there are

endemic to that (Figure 4). Shale barrens are also unconfirmed habitats for A. sexlineata in Virginia (Mitchell 1994). West Virginia also has shale barrens and these areas are potential habitat however, they are limited in West Virginia (Figure 6) and therefore limit the areas A. sexlineata can inhabit.

Railroads are ideal for A. sexlineata because they mimic the hot, xeric habitats of shale barrens. Railroads have characteristics similar to the desert floor habitat (Ruthven 1907).

Lizards are known to move along railroad tracks into previously uninhabited areas (Mitchell

1994). All known populations in Virginia are found along railroad tracks and they have also been found along railroads in Kansas. In West Virginia, they are found along one railroad near

Paw Paw in Morgan County (Figure 5).

The Potomac River creates a natural barrier between Maryland and West Virginia.

Aspidoscelis sexlineata occur in Maryland across the river from where they are found in West

Virginia. It is believed that A. sexlineata crossed the Potomac River into West Virginia via the railroad bridge (Humphries et al. 1999). Aspidoscelis sexlineata were found at the railroad in the 1990’s and other searches have resulted in no new populations (Table 1).

The objectives of my study were to: (1) investigate if A. sexlineata is expanding its range in West Virginia (2) study aspects of its natural history and (3) compare activity patterns of A. sexlineata in West Virginia to areas with different climates.

6

Figure 1. Six dorsal stripes against a black background found on A. sexlineata.

Figure 2. Female A. sexlineata with a white/cream ventral side.

7

Figure 3. Male A. sexlineata with blue tint on ventral side.

Figure 4. Shale barren in Maryland where A. sexlineata were captured.

8

Figure 5. Railroad in Morgan County where A. sexlineata have been found.

9

Figure 6. Shale barren locations in West Virginia. Picture adapted from West Virginia Wildlife.

10 Date Location Northing Easting Present 6/1/2004 Pendleton 4307793 646162 No 6/1/2004 Grant 4316660 651143 No 6/1/2004 Grant 4316998 651535 No 7/12/2004 Grant 4316660 651143 No 7/12/2004 Grant 4316715 652623 No 7/12/2004 Hardy 4317328 670760 No 7/12/2004 Hampshire 4353723 684380 No 7/12/2004 Hampshire 4363639 694307 No 7/12/2004 Hampshire 4363711 694205 No 7/12/2004 Hampshire 4366345 696444 No 7/12/2004 Hampshire 4367385 697008 No 7/12/2004 Hampshire 4370417 668994 No 7/12/2004 Hampshire 4373009 702981 No 7/12/2004 Hampshire 4379180 705117 No 7/12/2004 Hampshire 4374823 721916 No 7/12/2004 Hampshire 4362457 7211470 No 7/12/2004 Hampshire 4366450 708053 No 7/12/2004 Hampshire 4369438 703630 No 7/20/2004 Pendleton 4281298 631369 No 7/20/2004 Pendleton 4285290 633556 No 7/20/2004 Pendleton 4296400 639238 No 7/20/2004 Pendleton 4294142 638529 No 7/20/2004 Pendleton 4293246 640807 No 7/20/2004 Pendleton 4309376 646922 No 7/20/2004 Randolph 4307822 612335 No 9/14/2007 Grant 4308027 0649095 No 9/15/2007 Grant 4308027 0649095 No 9/22/2007 Mineral No 9/22/2007 Grant No 9/22/2007 Pendleton No 9/22/2007 Pendleton No 9/22/2007 Pendleton No 9/29/2007 Mineral 689670 4381303 No 9/29/2007 Mineral 690380 4377740 No 9/29/2007 Mineral No 9/30/2007 Grant 4308066 0649086 No Table 1. Aspidoscelis sexlineata searches in the state of West Virginia.

11 CHAPTER 3

METHODS

A mark‐recapture study was conducted from May until August of 2007 to estimate the size of a population of A. sexlineata in West Virginia. Lizards were trapped during the day and traps were checked every two hours to allow for normal activity to resume. No trapping was conducted later than the 1st of August; instead, visual encounter surveys were conducted until

September 28, 2007.

Trapping

I used two 5m and one 8m drift fences, which were anchored to the ground at the beginning of each trapping period and removed at the end. Traps were covered with cover boards to protect lizards from excessive sunlight, and they were not baited. They were placed in an area where lizards were known to be active (Figure 7). I used 6 funnel traps per drift fence, adapted from the National Park Service (Busteed 2006). I used 6 double‐sided funnel traps and 12 one‐sided funnel traps. Funnel traps were made out of hardware cloth and tied together with zip cords (Figure 8). Funnels were cut from a 61 cm X 61 cm square of hardware cloth and attached to the traps with binder clips (Figure 9). The body of the trap measured 45 cm X 91.4 cm (Figure 10).

Downes and Borges (1998) used sticky traps and claimed them to be effective, but they recommended that traps be placed in the shade. Unfortunately, there were no areas of shade where the lizards were active at my study site. While attempting to use these traps, lizard mortality occurred due to excess sunlight. Sticky traps were also difficult to transport and

12 proved to be inadequate for field work. Sticky traps were initially used for trapping, but were abandoned.

Data recorded

The following measurements were taken with calipers to the nearest 0.1mm on all captured lizards: snout‐to‐vent length (SVL), tail length (TL), head width (HW), head length (HL), weight (W), and sex was determined. Weight was measured by placing the lizard in a Ziploc® bag and hanging it by a My Weigh® 100g spring scale. Lizard coloration was used to determine sex; males are characterized by a bluish color on their ventral surface (Figure 3). Any other unique characteristics were recorded such as tail re‐growth and reproductive status. Soil temperature was taken with a TruTemp™ thermometer when the traps were set and at the time the traps were checked. Soil temperature was taken at three places to give an average of the trapping area. Air temperature was taken using an Enviro‐Safe® thermometer when traps

were checked, but it was only checked during June and July.

Mark‐recapture

One or two toes were clipped from each lizard to give it a specific number (Figure 11).

These numbers were used to indicate its recapture status. Toe clipping has been shown to have

no effect on the running speeds of lizards (Borges‐Landaez 2003, Dodd 1993). Toes were placed in a 1.5ml eppendorf™ vial with 75% ethanol to use for future DNA analysis.

Data analysis

Program Mark was used to estimate the population size (White 2007). A goodness‐of‐fit test was first performed using the program RELEASE (White 2007). The recapture test best suited to the data was the Cormack‐Jolly‐Seber (CJS), which is used in open systems. Using the

13 program Statistix 7.0, a Kruskal‐Wallis test was performed to compare May and June soil temperatures. A Friedman two‐way ANOVA, was performed to compare air temperatures versus time of day for June and July. To compare number of lizards caught per month a

Kruskal‐Wallis one‐way ANOVA was performed.

14

Figure 7. Trapping site along the railroad with drift fences.

Figure 8. Side view of funnel trap, placed against drift fence.

15

Figure 9. Funnel design. (adapted from Busteed 2006)

Figure 10. Funnel trap design with measurements. (adapted from Busteed 2006)

Figure 11. Clipping the toe of S. undulatus to give it a specific identification number

16 CHAPTER 4 RESULTS

Lizards were trapped for three separate weeks during May, June, and July of 2007. Of

35 lizards captured, 32 were A. sexlineata. Of these 32, 9 were males and 10 females, the rest were juveniles. Six of the 35 lizards escaped before they were able to be sexed or measured.

There were 11 recaptures. Four lizards were recaptured once, 2 were recaptured twice, and 1

was recaptured three times. Four gravid females were captured during all months of trapping

(Figure 12). A population estimate of 171 individuals was found for an open system. For a closed capture system, a population estimate of 70.2 was found.

Average soil temperature increased during the day. The highest average soil temperature for a specific time of day was found at 1500 h (Figure 13). The month with the highest average soil temperature of 32.6 °C was June (Figure 14). Soil temperatures recorded in

May and June were significantly different. The lowest soil temperature recorded was 17.7 °C and lizards were not active at this temperature. The lowest soil temperature at which lizards were active was 21.4 °C.

Air temperature for time of day of June and July was not significant. For June and July, air temperatures were significantly different (Figure 15). July had a higher average air temperature of 29.4 °C and also had higher daily air temperatures than June. Air temperatures showed a gradual increase throughout the day with a decrease around 1700h (Figure 16). Soil temperature was always higher than air temperature, but in June (Figure 17) there was a larger difference between these values than in July (Figure 18).

17 Numbers of lizards caught per month were not significantly different. The majority of lizards were captured during June (Figure 19). Lizards were most active between 1100h and

1500h (Figure 20).

On a day when the highest number of lizards was captured the highest average soil temperature of 36.5 °C was recorded, but the air temperature was not at its highest. At the lowest air temperature of 19.6°C and soil temperature of 26.1°C, no lizards were seen or heard.

Other reptilian species caught in traps or found around the trapping area were Common

Five‐lined Skinks, Eumeces fasciatus, Eastern Fence Lizards, Sceloporus undulatus, Black

Ratsnakes, Elaphe obsoleta, Eastern Hog‐nosed , Heterodon platirhinos, Eastern Garter

Snakes, Thamnophis s. sirtalis, and Eastern Box Turtles, Terrapene carolina.

18

Figure 12. Gravid A. sexlineata.

19 35

30 °C

25

20

15 Temperatures

10 Soil 5

0 900 1100 1300 1500 1700 Time

Figure 13. The relationship between soil temperature and time of day.

35

(°C) 30

25

20

15

10 Temperatures 5

Soil 0 May MoJunneths July

Figure 14. The relationship between soil temperature and month.

20 * 30

C) * ( ( 25

20

15

10 Temperatures Temperatures

5 Air 0 June July Months

Figure 15. The relationship between air temperatures for June and July

35

30

(°C) 25

20

15 Temperatures 10 Air 5

0 900 1100 1300 1500 1700 Time of Day

Figure 16. The relationship between air temperatures for June and July and time of day.

21 40

35

C) 30 (

25

20 Air Temperature 15 Soil Temperature 10

Temperatures 5

0 6/11 6/12 6/13 6/14 6/15 Date

Figure 17. Average air and soil temperatures for June.

40 35 C) (

30 25 20 Air Temperature 15 Soil Temperature 10 5 Temperature 0 7/30 7/31 8/1 Date

Figure 18. Average air and soil temperatures for July.

22 6

captured 5

4 3 lizards

of 2

1 0 May June July Number Months

Figure 19. The relationship between the number of A. sexlineata caught and month.

s

d 14

zar 12 li li d

10 f f

o 8

6 er

b 4 captured 2 um 0 Nb N 900 1100 1300 1500 1700 Time of Day

Figure 20. The relationship between number of A. sexlineata caught and time of day.

23 CHAPTER 5 DISCUSSION

Activity Patterns

A goal of this study was to determine if activity patterns in a cold climate region were the same as patterns for studies performed in warmer climate areas. Aspidoscelis sexlineata were active from the 18th of May until the 28th of September in West Virginia, when the last survey was conducted. Studies in northeastern Kansas found adults were active from May until

September, but juvenile activity was not recorded (Fitch 1958). Aspidoscelis sexlineata were active from May until September in Indiana (Minton 1944), April until October in Texas (Clark

1976) and from mid‐April to mid‐November in Alabama and Georgia (Etheridge et al 1983).

Aspidoscelis sexlineata in West Virginia had a similar activity patterns to lizards in Kansas and

Indiana, but had a shorter activity period compared to Texas, Alabama, and Georgia. This may be due to the differences in climate between these states and West Virginia. Using 2007 summer climate data from NOAA, a difference of about 10 C° was found between the air temperature highs of West Virginia and Texas (Figure 21). The air temperature lows had a difference of about 16 C° with West Virginia always having the lower temperatures (Figure 21).

Soil most likely warms faster and stays warmer longer in Texas than in West Virginia and this may be the reason for the Texas population having a longer activity period. Kansas summer air temperatures were about 5 C° higher than West Virginia temperatures (NOAA). Activity

patterns for Kansas and Indiana are similar to those in West Virginia, which may be attributed to the similarity in climate data (Figure 21). Pianka (1970) found southern Texas populations of

24 Coastal , tigris, to have a longer activity period than more northern lizards in Colorado. These differences could be attributed to the ecological challenges that northern populations face, such as colder temperatures and therefore a shorter period of seasonal activity. This may be why lizards in Kansas and Indiana have a shorter activity period than those found in Texas, Alabama, and Georgia.

There was a significant difference between soil temperatures in May and June, with the latter having higher temperatures. Lizard activity has been found to be correlated with soil temperatures (Paulissen 1988, Winne and Keck 2004). Aspidoscelis sexlineata need soil

temperature to be above a certain temperature before they begin their daily activities. In my study, soil temperatures at the beginning of May might not have been high enough to initiate lizard activity. However, there was no relationship between soil temperature and the number of lizards captured (Figure 22). Once soil temperatures are adequate for activity, other factors will influence lizard activity, such as air temperatures, breeding habits, and foraging behaviors.

There was a decrease in the number of lizards captured in July when temperatures were still high (Figure 19). This may be due to the high amount of rainfall that occurred during the

July trapping period. When the moisture content of the soil is high, lizards have difficulty reaching their temperature threshold for activity and so they are not seen on the surface and therefore are not captured (Hardy 1962). Due to high soil moisture, lizards cannot reach optimal body temperatures, which then decrease the chance of survival during hibernation by limiting the amount of fat they can store. It also leads to an increase in likelihood of predation and causes eggs to rot or prolongs the incubation period. High moisture content also produces more vegetation, which will limit the areas of basking and decreases the amount of suitable

25 habitat (Hardy 1962).

Daily air temperatures for July were higher than those of June (Figure 16), but more lizards were captured in June. Along with the excess soil moisture, this could be due to temperatures being above the maximum threshold for lizard activity in July, so lizards were either only active for a short period or not at all. This agrees with the theory that lizards retreat due to high afternoon temperatures (Milstead 1957). However, I ultimately believe the increase in captures in June was due to the breeding season because more lizards are active during this time and so more are captured.

Hardy (1962) found A. sexlineata in Kansas active from morning to early afternoon, but as the summer progressed lizards were only active before 0930. In contrast, I found that most lizards were captured from 1100 to 1500 and that this was maintained through the summer

(Figure 20). Climate seems to be a determining factor in daily activity times. West Virginia could have cooler morning and hotter afternoon temperatures than Kansas. Keeled Earless

Lizard, Holbrookia propinqua, were found to have a later initiation of activity in the spring than the summer in Texas. In each season they were active at similar temperatures (Judd 1975).

Holbrookia propinqua are smaller than A. sexlineata and are found on barrier beaches and sand dunes (Winchester 2001). This lizard emerges at air temperatures and soil temperatures equal to 28 C°. Aspidoscelis sexlineata was found to be active at soil temperatures lower than 28 C°.

This is probably due to adaptation to cooler mornings and cooler temperatures. Damme et al

(1987) found that times of activity changed seasonally for Common Lizard, Lacerta vivipara.

During the spring and late summer they were found in direct sunlight, but during the summer months they were found in the shade. Temperature differences seem to play a role in these

26 lizards’ behavior and activity patterns. Similarly it appears that temperature is playing a role in

A. sexlineata’s behavior and activity patterns.

Aspidoscelis sexlineata have an early retreat of around 1500 h with no reappearance.

Other lizards have been found to be active throughout the entire day, to show a bimodal

pattern, or are active past sunset. Canyon Lizard, Sceloporus merriami, were found active in the

morning, inactive in the afternoon and active again during the evening giving it a bimodal pattern (Grant and Dunham 1988). Desert Horned Lizard, Phyrnosoma platyrhinos, were found active about 2 hours after sunrise and sometimes 2 hours past sunset (Pianka and Parker

1975). Zebra‐tailed Lizard, Callisaurus draconoides, were found to be active slightly after sunset

during the summer (Kay 1972). Some theories given for A. sexlineata’s early retreat are satiation (Hardy 1962), high afternoon temperatures (Milstead 1957), endogenous rhythms

(Winne and Keck 2004), lack of territoriality (Rose 1981), higher benefits of inactivity (Rose

1981), and limited availability of operative temperatures (Grant and Dunham 1988).

Aspidoscelis sexlineata perhaps have a reduced activity time to avoid competition with other lizard species, because resources will be different throughout the day and season (Pianka

1973). Another possible reason for short daily activity is that as activity time increases, metabolic costs will also increase due to energy expenditure pursuing prey, defending territories, and a higher resting metabolic rate (Adolph and Porter 1993). This would result in more energy needed for the day and less fat storage for hibernation.

I found a population estimate of 171 when using an open system analysis and 70.2 individuals when using a closed system analysis. A closed system implies that the size of the population is constant over the study period, no immigration or birth occurs, and no emigration

27 or death occurs. There can also be a geographic barrier to these . Seventy point two individuals depict a better estimate for the area that was surveyed, but it is difficult to determine if no emigration or immigration occurred. Both of these estimates indicate a healthy population of lizards that are self‐reproducing and could occupy this area for a significant period of time.

Expansion

One goal of my study was to determine what local area these lizards came from and how they moved into West Virginia. Aspidoscelis sexlineata were first found in West Virginia in the 1990’s, so it is not known how long they have inhabited this region. It is believed that they came from Maryland into West Virginia via the CSX railroad which runs along the Potomac River and runs across it in several areas. When the railroad was built the habitat was altered to a xeric habitat which allowed lizards to occupy the area. In Mushinksky’s (1985) study on the effects of burning, he found that A. sexlineata were well established on the most intensely disturbed habitat. After surveying both Maryland and West Virginia populations, I believe these

lizards crossed the Potomac River via the railroad.

The thermal activity range for A. sexlineata has been found to be the highest for any species of lizard found in the eastern United States (Mitchell 1994), which restricts them to a xeric habitat. However, their thermal activity range is associated with habitats which are unfavorable to other lizard species, giving them a stronger advantage in these habitats (Hardy

1962).

Another goal of this study was to determine if A. sexlineata are expanding farther into

West Virginia. In the past, A. sexlineata were only reported at one section of the railroad. After

28 surveying more of the railroad, I found A. sexlineata, 5.0km (3.1mi) farther down the railroad into West Virginia than had been previously recorded by past investigators (Figure 23). While this may indicate an expansion of range, it appears that this expansion is limited to areas immediately adjacent to the railroad, because A. sexlineata were not found in forested habitat near the railroad. Forested areas near the railroad may not provide enough sunlight for these lizards to reach their optimal temperature for activity.

Both S. undulatus and E. fasciatus were found occupying the same area as A. sexlineata, so they are possibly competing for basking, nesting, and hibernation sites. Greenburg (2000) found the Southeastern Five‐lined , Eumeces inexpectatus, were negatively correlated with the amount of bar sand in an area, while A. sexlineata were positively correlated. Aspidoscelis sexlineata could possibly out‐compete E. fasciatus for food resources because they have the

same foraging habits. In contrast, Sceloporus undulatus and A. sexlineata differ in their foraging

habits, so are probably not competing in this area of their niche. I therefore believe that if competition is occurring with S. undualatus it is over resources other than food.

Natural history

Eastern Hog‐nosed Snakes, Heterodon platirhinos (Figure 24) and Black Ratsnakes,

Elaphe o. obsoleta, were caught in traps and may be potential predators. The primary diet of

Black Ratsnakes is birds, bird eggs, and rodents, but other prey items include Fence Lizards and

Five‐lined Skinks (Mitchell1994). Hog‐nosed ’s diet consists mostly of toads, but other prey items have been recorded; such as frogs, salamanders, and chipmunks (Mitchell 1994).

Although these snakes show a preference for non‐lizard prey items, they have been shown to

29 consume lizards (Mitchell 1994). Because these snakes were found in traps and in the trapping area, they could prey on A. sexlineata. Some A. sexlineata were found with obvious re‐growth of their tails (Figure 25), which is evidence of possible predation. They are generally not known to autotomize their tails like skinks, because their tails are used for balance (Mitchell 1994).

One of the main reasons a lizard drops its tail is due to predation. Maybe the usefulness of an easily autotomized tail is not as important as balance for running when evading a predator.

Burrows are also a way A. sexlineata evade predators. Burrows have been classified into four types: temporary unexcavated shelters, escape‐burrows, escape tunnels, and inactivity tunnels

(Hardy 1962). Shown below is an escape‐burrow where a lizard disappeared into before the photograph was taken (Figure 26), many lizards were seen to disappear into such tunnels when

checking traps. Judd (1974) states that these burrows are used to protect against colder temperatures and maintain body temperatures for the species Holbrookia propinqua. Pough

(1970) stated that burrowing is not an effective tool for thermoregulation, but a defense mechanism for the Sand Lizard, Uma notata found in California. In a cold climate and high predator areas, burrows would be a key to survival.

30

Figure 21. Average July temperatures for the summer of 2007 for the United States, showing temperature differences between Texas, Kansas, Alabama, Georgia, Indiana, and West Virginia. (cdo.ncdc.noaa.gov)

40.0 16

35.0 14 C)

º 30.0 12

25.0 10

20.0 8

15.0 6 Soil Temperature 10.0 Number of Lizards 4 Soil temperature ( 5.0 2 Number of Lizards Caught

0.0 0 7:12 8:24 9:36 10:48 12:00 13:12 14:24 15:36 16:48 18:00 Time of Day Figure 22. The relationship between soil temperature and number of A. sexlineata captured and time of day.

31

Figure 23. Google map of the Potomac River and CSX Railroad between Maryland and West Virginia. Site A was the main trapping site. Site B is the area where A. sexlineata were found further down the railroad into West Virginia. Site C is the site where A. sexlineata were trapped in Maryland. Notice the railroad crosses the river in several different areas.

Figure 24. Heterodon platirhinos caught near trapping area.

32

Figure 25. Re‐growth of A. sexlineata tail.

Figure 26. Aspidoscelis sexlineata taking refuge in a burrow.

33 CHAPTER 6 CONCLUSION

Habitat degradation, fragmentation, and destruction may cause populations to decline

or may allow them to expand. Aspidoscelis sexlineata are expanding along the railroad into

West Virginia and have been recorded farther into West Virginia than previously documented.

As to whether these lizards will expand into other areas of West Virginia and how far north their range can potentially extend, needs to be studied in more detail. These lizards are a highly‐adaptive species and can survive well in areas of high disturbance (Mitchell 1994). They are observed in cold climate areas, so their range could potentially expand if habitat and soil

temperatures are suitable. These two criteria seem to be the limiting factors for them to establish in an area.

Shale barrens of West Virginia have been found to be uninhabited by A. sexlineata. At this point, railroads do not come in contact with shale barrens and therefore may not be

possible for A. sexlineata to migrate into shale barrens.

Aspidoscelis sexlineata appear to have crossed the railroad bridge from Maryland into

West Virginia. Genetic analysis comparing individuals from each population needs to be done to confirm this. More DNA needs to be collected from the Maryland population before this can

be accomplished. DNA from the West Virginia populations is being kept in the Herpetology

museum at Marshall University for possible further research.

Aspidoscelis sexlineata have different activity patterns in West Virginia compared to other states, perhaps due to differences in climate. Further studies on hibernation and body

34 temperatures would prove useful in determining how this species might differ in other climates.

35 PART 2 CHAPTER 7 OVERVIEW

Most people believe that species need to be saved from extinction, but many of today’s endangered species programs focus on species that include less than 100 individuals (Scott et al. 1987). When species reach this level of abundance it becomes costly and any event could wipe out the population. Scott et al. (1987) stated that this kind of “Emergency Room

Conservation” is unacceptable. The main focus of conservation should be on species diversity through the survival of common species (Scott et al. 1987). We should prevent a species from becoming endangered rather than wait until there are only a few individuals left. Some measures that are taken to prevent a species from becoming extinct are controlling exotics and translocations to new habitats, which prove to be risky and expensive. These tasks should be refined through the use of common species (Hunter and Hutchinson 1994), because these populations can withstand changes.

Warren et al. (2001) stated that habitat degradation and climate change are thought to be altering the distributions and abundances of animals and plants through‐out the world.

These forces can eventually cause habitat specialists to decline while leaving communities dominated by habitat generalists. Habitat specialists are those species with one or a few similar

habitats and generalists are those species that can occupy a broad range of habitats (McPeek

1996). Warren et al. (2001) found that out of 46 species of butterflies, half of the generalist species increased their distribution, while the other half of generalists and 89% of specialist species declined. Determining which species is a habitat specialist or generalist can be an

36 indicator of which species are competing for resources (McPeek 1996). Garrot and White

(1993) stated that populations of native species that are generalists have increased in numbers and distribution over the past century. These species can monopolize resources, introduce or spread parasites, cause local extinctions, and change species composition. Not only do species

have to adapt to habitat and climate changes, but they also have natural processes still occurring such as competition between species. An overabundance of native species will cause more competition between species.

Competition for resources, predation, and nutrient availability are ecological interactions that can limit a species habitat selection. Understanding these interactions and how they influence communities can help predict which species could be impacted and which will be unaffected by alterations in the environment (McPeek 1996).

To reduce competition, species will occupy different niche dimensions. They do this by altering their activity patterns, habitats, and the foods they consume (Pianka 1973). Many species occupy different microhabitats and habitats. Some lizards are fossorial, terrestrial, arboreal, and some are a mixture of these (Pianka 1973). A first step in identifying which species may be influencing another is by identifying any possible overlap in niches. Morphology can sometimes be used to indicate the animal’s niche. For example, a fossorial species will have reduced appendages or none at all, terrestrial species differ in leg size depending on their foraging type, arboreal species are usually long‐tailed and slender (Pianka 1973). Studying morphometrics is a good way to determine an animal’s habitat niche. Hunting for prey, even though more behavioral, can indicate which food niche is occupied. There are two types of foragers, a lizard that actively searches for prey and one that sits and waits for prey. Pianka

37 (1973) stated that in North America, prey items are a major factor that separate out niche and most teids and skinks are active foragers.

A species that can occupy a habitat more efficiently than another will eventually eliminate the other (Volterra 1931, Gause 1934). Gause’s Law or the competitive exclusion

principle states that closely related species that occur in a single habitat will be ecologically separate (Gause 1934). However, Pianka (1973) and Elton (1946) state that competition should occur more frequently between co‐generic species due to their ecological similarity.

There are six species of lizards found in West Virginia (Green and Pauley 1987), including the recent addition of Aspidoscelis sexlineata, Eastern Six‐lined Racerunner (Humphries 1999).

These species occupy similar habitats and could be competing for resources. There have been no studies of lizards in West Virginia, thus little is known about these species. The objectives of my study were to determine where populations existed and where species co‐exist to provide baseline data in regards to the lizards of West Virginia.

38 CHAPTER 8 INTRODUCTION

There are 6 species of lizards found in West Virginia including Eastern Six‐lined

Racerunner, A. sexlineata, Northern Fence Lizard, Sceloporus undulatus, Common Five‐lined

Skink, Eumeces fasciatus, Little Brown Skink, Scincella lateralis, Broad‐head Skink, Eumeces laticeps, and Northern Coal Skink, Eumeces a. anthracinus. They all have an individual state ranks assigned by the West Virginia Division of Natural Resources and are unique in their natural history. Common Five‐lined Skinks and Northern Fence Lizards are ranked S5, which means they are very common and demonstrably secure. Little Brown Skinks are ranked S3, which means there are 21 to 100 occurrences documented. Broad‐head Skinks and Northern

Coal Skinks are ranked S2, which means there are few remaining within the state, approximately 6 to 20 individuals. These rankings help distinguish between species that are common and those that are uncommon.

Northern Fence Lizard

Northern Fence Lizards, Sceloporus undulatus hyacinthinus, are found in most areas of

West Virginia (Figure 27) (Green and Pauley 1987). The body, head, and tail range from gray to brown and they have undulating black cross bands that run the length of the body (Figure 28).

Males have a bluish patch on their throat and along their sides; females may have a slight tinge of blue on their throat (Mitchell 1994). S. undulatus may be considered a habitat generalist and

39 is found in xeric open pine woods, mixed hardwood forests, mixed deciduous forests, stand of

trees in old fields, around barns and houses, and on rock piles (Mitchell 1994). They have also been seen on rail fences and rotting logs or stumps and are generally an arboreal species

(Conant and Collins 1998). They are active from March until November. They will overwinter in

tree holes and crevices in rock piles (Mitchell 1994). They are “sit and wait” foragers” and consume a variety of invertebrates (Mitchell 1994).

Common Five‐lined Skink

Common Five‐lined Skinks, Eumeces fasciatus, are found through‐out West Virginia

(Figure 29)(Green and Pauley 1987). They have five white to cream stripes on a dark‐brown

background, which will fade to an olive‐brown as aging occurs (Figure 30). Juveniles have a blue tail and can be confused with Broad‐head Skinks (Mitchell 1994). Eumeces fasciatus may also be considered a habitat generalist and have been found in a variety of habitats such as; mature hardwood forest, bottomland forest, and second‐growth hardwood, mixed hardwood‐ pine forest, dry pine woods, cypress swamp forest, urban wood lots, edges of woods and fields, urban and suburban buildings (Mitchell 1994). These skinks are typically found in damp areas,

such as rotting stumps and logs, decaying debris, and abandoned boards or sawdust piles. They are terrestrial, but are occasionally arboreal (Conant and Collins 1998). They are active from

April or May until October and sometimes seen during the winter if it is warm enough.

Little Brown Skink

Little Brown Skinks, Scincella lateralis, are found in a few counties of the state (Figure

40 31)(Green and Pauley 1987). The body color ranges from tan to golden brown with a dark‐ brown stripe that runs laterally. Scincella lateralis is an almost exclusively terrestrial/fossorial species and is often found in the leaf litter of hardwood and mixed‐hardwood forests, some urban woodlots, and some pine forests (Mitchell 1994). These skinks can be found among leaves, decaying wood, and detritus of the woodland floor (Conant and Collins 1998). They are active from March until October and may be active during warm winter months (Mitchell 1994).

Eumeces laticeps

The Broad‐head Skinks, Eumeces laticeps, have only been found in a few counties of the

state (Figure 32) (Green and Pauley 1987). They have five narrow stripes on a dark‐brown to black background and these stripes coalesce as they mature. Males have a reddish orange head during the breeding season. These skinks are the most arboreal of the skinks in West Virginia

(Green and Pauley 1987). Their favored habitat is a forest and it prefers more xeric forests than the Common Five‐lined skink. It can also be found in many other habitats ranging from a swampy forest to parking lots (Mitchell 1994). Eumeces laticeps can be found on tree branches,

under leaf litter, in tree holes, and sometimes rail fences. (Conant and Collins 1998) These skinks are very secretive and flee instantly when humans approach, making it a hard species to find. They are active from April through August (Mitchell 1994).

Eumeces a.anthracinus

The Northern Coal Skink, Eumeces a. anthracinus, are found in the southern and eastern parts of the state (Figure 33) (Green and Pauley 1987). This species has a mid‐dorsal

41 stripe on the tail and body that ranges between a broad tan to brown with light dorsolateral

stripes bordering it (Mitchell 1994). The body is dark brown to black and the ventral side is cream to light blue (Mitchell 1994). E. anthracinus are found in forested areas with rock cover and sunlit openings and can also be found in humid portions of wooded hillsides (Conant and

Collins 1998, Mitchell 1994). In general, skinks are known to be active foragers, but it is unknown whether this is true for E. anthracinus. It also unknown whether this species is a habitat generalist or specialist.

All lizards in West Virginia can be found in forested areas, indicating an overlap in habitat niches. Competition may be reduced because of microhabitat differences. Sceloporus undulatus and E. fasciatus are found through‐out the state of West Virginia (Green and Pauley

1987) and are possible generalists or opportunists. Greenburg (2000) suggests that

Southeastern Five‐lined Skinks, Eumeces inexpectatus, and Little Brown Skinks, Scincella lateralis, are generalist species. These species probably adapt to areas with anthropogenic changes more than the specialist species (Greenburg 2000). Eumeces laticeps and E. a. anthracinus are possible specialists and therefore will not survive in areas with a high amount

of change as well as the other species previously mentioned.

42

Figure 27. Known locations of S. undulatus from the West Virginia Biological Survey

43

Figure 28. Sceloporus undulatus with a gray body and keeled scales.

44

Figure 29. Known locations of E. fasciatus from the West Virginia Biological Survey

45

Figure 30. Adult male E. fasciatus with orange head and olive brown body, caught in trap at the railroad.

46

Figure 31. Known locations of S. lateralis from the West Virginia Biological Survey

47

Figure 32. Known locations of E. laticeps from the West Virginia Biological Survey

48

Figure 33. Known locations of E. a. anthracinus from the West Virginia Biological Survey

49 CHAPTER 9 METHODS

Skink and racerunner surveys were conducted throughout the state of West Virginia to

determine presence of common and uncommon species. Surveys were conducted from May to

October of 2007. Visual encounter surveys were the main search technique used. Surveys consisted of hiking in silence, listening for movement and looking for lizards. Hand collection,

noosing, and trapping were used to capture lizards; the first was the most successful. Sites searched were chosen from museum records. State parks and trails in the state were common survey sites. Sites surveyed consisted of railroads, trails within forests, open areas, and areas of human development.

Data recorded

The following measurements were taken with calipers to the nearest 0.1 mm on all captured lizards: snout‐vent length (SVL), tail length (TL), head width (HW), head length (HL), weight (W), and sex was determined. Weight was measured by placing the lizard in a Ziploc® bag and hanging it by a My Weigh® 100 g spring scale. Any other unique characteristics were recorded such as tail re‐growth and reproductive status.

50 CHAPTER 10

RESULTS

Fayette, Tucker, Nicholas, Kanawha, Cabell, Grant, Morgan, Pocahontas, Lewis, and

Wayne counties were surveyed during the summer of 2007. Lizards were found from mid‐May until the end of September. The majority of lizards observed were Fence Lizards and Five‐lined

Skinks. One Coal Skink, Little Brown Skink, and Broad‐head Skink were seen at Beech Fork State

Park in Cabell County. Fence lizards were found at the CSX railroad in Morgan County,

Summersville Dam in Nicholas County, East Lynn upper dam area in Cabell County, Beech Fork

State Park in Cabell County, and North Fork Mountain in Grant County (Table 2, Figure 28).

Five‐lined skinks were found at the CSX railroad in Morgan County, East Lynn upper dam area in

Cabell County, along Ridgewood Road in Cabell County, along a railroad at Swiss Junction in

Nicholas County, and Kanawha State Park in Kanawha County (Table 2, Figure 34). Lizards were not seen in areas searched in Fayette, Tucker, and Wayne counties (Table 3, Figure 34). Surveys were conducted in forested areas, along hiking trails, maintained walking trails, dammed areas, residential areas, and along railroads.

51 Table 2. Locations where lizards were found during surveys for the summer of 2007.

Date Species Road/ Trail County GPS 5/19 Fence Lizard (2) Near Railroad Morgan 0723337/4385768 5/19 Five‐lined Skink Near Railroad Morgan 0723337/4385768 5/21 Fence Lizard Near Railroad Morgan 0723337/4385768 6/8 Fence Lizard (2) Summersville Dam Nicholas 6/22 Fence Lizard (2) East Lynn, below dam Cabell 0378859/4223037 6/23 Fence Lizard (4) East Lynn, upper dam Cabell 0378859/4223037 6/23 Five‐lined Skink East Lynn, upper dam Cabell 0378859/4223037 6/24 Little Brown Skink Beech Fork, Twin Cove trail Cabell 0375580/4238203 6/25 Five‐lined Skink Ridgewood Rd Cabell 6/26 Coal Skink Beech Fork, Twin Cove trail Cabell 0375580/4238203 6/26 Broad‐head Beech Fork, Twin Cove trail Cabell 0376306/4238685 6/27 Fence Lizard (2) East Lynn Cabell 0378859/4223037 6/27 Five‐lined Skink East Lynn Cabell 0378859/4223037 7/22 Five‐lined Skink Swiss Junction, along RR Nicholas 0489497/4230792 7/22 Five‐lined Skink Kanawha State park Kanawha 0442437/4236155 9/30 Fence Lizard North Fork Mountain Grant 0649086/4308066

52 Date Road/Trail County 5/11 FR‐18, Fire Tower Tucker 5/11 Douglas Road‐27 Tucker 5/14 Babcock State Park Fayette 5/15 Thurmond‐Minden Trail Fayette 5/15 Buffalo Creek Fayette 5/16 Road 19‐25, Railroad Nicholas 6/6 Kanawha State Forest Kanawha 6/7 Kanawha State Forest Kanawha 6/18 FR‐18, Fire Tower Tucker 6/18 Douglas Road‐27 Tucker 6/19 Beech Fork State Park Cabell 6/22 Beech Fork State Park Cabell 6/23 Ridgewood Road, Huntington Cabell 6/27 Beech Fork State Park Cabell 6/28 Barboursville Park Cabell 7/18 Buffalo Creek Fayette 7/18 Thurmond‐Minden Trail Fayette 7/19 Cranberry River Trail Pocahontas 7/20 Seneca Rocks Pocahontas 7/20 Cheat Bridge Pocahontas 7/21 Stonewall Jackson State Park Lewis Table 3. Locations searched during the summer of 2007, where no lizards were seen.

53

Figure 34. West Virginia counties where lizards were found and not found during the summer of 2007.

54 CHAPTER 11 DISCUSSION

A species that is a habitat specialist has more limitations than a habitat generalist species. Eumeces fasciatus and S. undulatus may be considered habitat generalists, while E. laticeps and E. a. anthracinus, and S. lateralis may be considered habitat specialists. E. fasciatus and S. undulatus were found in a variety of habitats, including forested and human developed areas. These lizards could be considered generalists because they can adapt to new situations, including human effects on their habitat (Smith and Remington 1996). Sceloporus undulatus have been shown to be flexible in their habitat use (Smith 1946). Eumeces a. anthracinus, E. laticeps, and S. lateralis were only found once during the survey period in forested habitats.

Even though S. lateralis have been shown to be habitat generalists (Greenburg 2000), only one was found during this survey. Some possible reasons for this could be they are secretive and difficult to find, other species are out‐competing them, or they are at the top edge of their range and so only a few occupy this area.

Another way these lizards may be in competition is through their prey items. All skinks in West Virginia are active foragers and therefore consume similar prey items. Their time and

spatial niche may be the indicator of which species are in the most competition. S. undulatus are “sit and wait” foragers, so their prey items differ from the Skinks.

Sceloporus undulatus and S. lateralis are first active in March, while E. fasciatus and E. laticeps are first active in April (Mitchell 1994), so this may indicate a brief period of decreased competition. Sceloporus undulatus is active the longest, until November, E. fasciatus and

55 Scincella lateralis are active until October, and E. laticeps is active the shortest, until August

(Mitchell 1994). These differences in times of activity could be a way these lizards have evolved to limit competition.

Sceloporus undulatus and E. fasciatus are found abundantly throughout the state of

West Virginia and can adapt well to human alterations, they are possibly out‐competing other

lizard species who do not survive well in altered habitats.

56 CHAPTER 12 CONCLUSION

Lizards were found in Morgan, Nicholas, Cabell, and Grant counties. Lizards were not found in Pocahontas, Lewis, Wayne, Fayette, and Tucker. However, museum records indicate

lizards could be in these counties. Eumeces fasciatus and S. undulatus were the most commonly found lizards, in both human disturbed habitat and natural areas, indicating possible

habitat generalists. Lizards are secretive and difficult to find, so they could have easily have been missed during this survey period. More surveys need to be accomplished so that natural history studies can be conducted and habitat can be defined and protected.

57 LITERATURE CITED

Adolph, Stephen C., and Warren P. Porter. 1993. Temperature, activity, and lizard histories. The American Naturalist 142(2): 273‐295.

Borges‐Landaez, P.A. and R. Shine. 2003. Influence of toe‐clipping on running speed in quoyii, an Australian Scincid lizard. Journal of Herpetology 37(3): 592‐595.

Baker H. G. and G. L. Stebbins. 1965. The genetics of colonizing species. Academic Press, New York, New York, USA.

Busteed, G., J.L. Cameron, S. Riley, L. Lee, A. Compton, E. Berbeo & T. Duffield. 2006. Monitoring of Terrestrial & Amphibians by Pitfall Trapping in the Mediterranean Coast Network: Santa Monica Mountains National Recreation Area & Cabrillo National Monument. Version 1.1. Natural Resources Report NPS/MEDN/NRR—2006/00?, 99 pp.

Burt, Charles E., 1928. The lizards of Kansas. Transactions of the Academy of Science of St. Louis.

Clark, Donald R. Jr., 1976.Ecological observations on a Texas population of six‐lined racerunners, Cnemidophorus sexlineatus. Journal of Herpetology. 10(2): 133‐138.

Cooch, Evan and Gary White. 2008. Program Mark: A gentle introduction. 6th edition.

Conant, R and J.T. Collins. 1998. Reptiles and Amphibians: Eastern/Central North America. 3rd Edition. Houghton Mifflin Company, New York, NY.

Damme, Raoul Van., Dirk Bauwens, and Rudolf F. Verheyen. 1987. Thermoregulatory responses to environmental seasonality by the lizard Lacerta vivipara. Herpetologica. 43(4): 405‐ 415.

Dodd, Kenneth C., 1993. The effects of toe‐clipping on sprint performance of the lizard Cnemidophorus sexlineatus. Journal of Herpetology. 27(2): 209‐213.

Downes, Sharon and Pedro Borges. 1998. Sticky traps: An effective way to capture small terrestrial lizards. Herpetological Review 29(2): 94‐95.

Etheridge, Kay., Lawrence C. Wit, and Jeffrey C. Sellers. 1983. Hibernation in the Lizard Cnemidophorus sexlineatus (Lacertilia:Teiidae). Copeia. 1983(1): 206‐214.

Fitch, H.S., 1958. Natural history of the six‐lined racerunner (Cnemidophorus sexlineatus). Univ. Kansas Publ., Mus. Nat. Hist. 11: 11‐62.

Gause, G.F., 1934. The struggle for existence. Williams and Wilkins Company, Baltimore, USA.

58

Garrott, Robert A., P.J. White, and Callie A. Vanderbilt White. 1993. Overabundance: An issue for conservation Biologists? Conservation Biology. 7(4):946‐949.

Grant, Bruce W. and Arthur E. Dunham. 1988. Thermally imposed time constraints on the activity of the desert lizard Sceloporus merriami. Ecology. 69(11): 167‐176.

Greenburg, Cathryn H., 2000. Fire, Habitat Structure and Herpetofauna in the Southeast. The role of fire in nongame wildlife management and community restoration: traditional uses and new directions, proceedings of a special worskshop. USDA Forest Service and Northeastern Research Station. pp 91‐99.

Green, N. Bayard., and Thomas K. Pauley. 1987. Amphibians and Reptiles in West Virginia. University of Pittsburgh Press.

Hardy, Donna F., 1962. Ecology and behavior of the six‐lined racerunner, Cnemidophorus sexlineatus. The University of Kansas Science Bulletin. 45: 1‐73.

Humphrey, Stephen R., 1974. Zoogeography of the Nine‐Banded Armadillo (Dasypus Novemcinctus) in the United States. BioScience, 24(8): 457‐462.

Humphries, Jeffrey W., Zachary I. Felix, W. H. Martin, and Thomas K. Pauley. 1999. A new lizard species, Cnemidophorus sexlineatus, in West Virginia. Proceedings of the West Virginia Academy of Science. 71(2): 1‐3

Hunter, Malcom L. Jr., and Alan Hutchinson. The virtues and shortcomings of parochialism: conserving species that are locally rare, but globally common. Conservation Biology. 8(4): 1163‐1165.

Judd, Frank W., 1975. Activity and thermal ecology of the keeled earless lizard, Holbrookia propinqua. Herpetologica. 31(2): 137‐150.

Kay, Fenton R., 1972. Activity patterns of Callisaurus draconoides at Saratoga Springs, Death Valley, California. Herpetologica. 28(1): 65‐69

McPeek, Mark A., 1996. Trade‐offs, food web structure, and the coexistence of habitat specialists and generalists. The American Naturalist. 148(Supplement):124‐138.

Minton, Minton Jr., 1944. Introduction to the study of the reptiles of Indiana. American Midland Naturalist. 32(2): 438‐477.

Milstead, WM W., 1957. Observations on the natural history of four species of whiptail lizard, Cnemidophorus (Sauria, Teiidae) in Trans‐pecos Texas. The Southwestern Naturalist. 2: 105‐121.

59

Mitchell, Joseph C., 1994. The Reptiles of Virginia. Smithsonian Institution Press.

Mushinksky, Henry R., 1985. Fire and Florida sandhill herpetofaunal community: with special attention to responses of Cnemidophorus sexlineatus. Herpetologica. 41(3): 333‐342.

NOAA Satellite and Information Service. 2008. National Climatic Data Center. cdo.ncdc.noaa.gov

Paulissen, Mark A., 1988. Ontogenic and seasonal comparisons of daily activity patters of the Six‐lined Racerunner, Cnemidophorus sexlineatus (Sauria: Teiidae). The American Midland Naturalist. 120(2): 355‐361.

Pianka, Eric R., 1970. Comparative autecology of the lizard Cnemidophorus tigris in different parts of its geographic range. Ecology. 51(4): 703‐720.

Pianka, Eric R., 1973. The structure of lizard communities. Annual Review of Ecology and Systematics. 4: 53‐74.

Pianka, Eric R., 1974. Niche overlap and diffuse competition. Proceedings of the National Academy of Sciences of the United States of America. 71(5): 2141‐2145.

Pianka, Eric R., and William S. Paker. 1975. Ecology of Horned Lizards: A review with special reference to Phrynosoma platyrhinos. Copeia. 1: 141‐162.

Pough, Harvey F., 1970. The Burrowing Ecology of the Sand Lizard, Uma notata. Copeia. 1970(1): 145‐157.

Rhoads, Samuel N., 1895. Contributions to the zoology of Tennessee. No. 1, Reptiles and Amphibians. Proceedings of the Academy of Natural Sciences of Philadelphia. 47: 376‐ 407.

Rose, Barbara. 1981. Factors affecting activity in Sceloporus virgatus. Ecology. 62(3): 706‐716.

Ruthven, Alexander G., 1907. A collection of reptiles and amphibians from Southern New Mexico and Arizona. Bulletin American Museum of Natural History. 23: 570‐573.

Scott, Michael J., Blair Csuti, James D. Jacobi, and John E. Estes. 1987. Species Richness. BioScience 37(11): 782‐788

Smith, Harvey R., and Charles Lee Remington. 1996, Food Specificity in Interspecies Competition. BioScience. 46(6): 436‐447.

Smith, H.M., 1946. Handbook of Lizards. Comstock, Ithaca, N.Y.

60 Smith, Michaella M., Henry T. Smith, Richard M. Engeman. 2004. Extensive contiguous north‐ south range expansion of the original population of an invasive lizard in Florida. International Biodeterioration & Biodegradation 54: 261‐264.

Statistix 7. Version 7.0. 1985, 2000. Analytical Software.

Taulman, James F., and Lynn W. Robbins. 1996. Recent range expansion and distributional limits of the nine‐banded armadillo (Dasypus novemcinctus) in the United States. Journal of Biogeography 23(5): 635‐648.

Trombulak, Stephen C., and Christopher A. Frissell. 2000. Review of ecological effects of roads on terrestrial and aquatic communities. Conservation Biology 14(1): 18‐30.

Warner, Daniel A., 2000. Ecological observations on the six‐lined racerunner (Cnemidophorus sexlineatus) in Northwestern Illinois. Transactions of the Illinois State Academy of Science 93: 239‐248.

Warren, M.S., et al. 2001. Rapid responses of British butterflies to opposing forces of climate and habitat change. Nature. 414: 65‐68.

White, Gary C., 2007. Program Mark. Version 4.3.

Winne, C.T., and M. B. Keck. 2004. Daily activity patterns of Whiptail Lizards (: Teiidae: Aspidoscelis): a proximate response to environmental conditions or an endogenous rhythm? Functional Ecology 18(3): 314‐321.

Winchester, M. 2001. “Holbrookia propinqua” (On‐line), . Accessed March 16, 2008. http://animaldiversity.ummmz.umich.edu/site/accounts/information/Holbrookia_propi nqua.html.

Volterra, Vito. 1939. Variations and fluctuations of the number of individuals in animal species living together. Animal ecology, with special reference to insects. pp. 409‐448.

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