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Nest-Site Selection in the Eastern Box Turtle,Terrapene carolina carolina, in a Population inCentral IllinoisBeth FlitzEastern Illinois UniversityThis research is a product of the graduate program in Biological Sciences at Eastern Illinois University. Findout more about the program.

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Recommended CitationFlitz, Beth, "Nest-Site Selection in the Eastern Box Turtle, Terrapene carolina carolina, in a Population in Central Illinois" (2003).Masters Theses. 1389.https://thekeep.eiu.edu/theses/1389

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NEST-SITE SELECTION IN THE EASTERN BOX TURTLE, TERRAPENE

CAROLINA CAROLINA, IN A POPULATION IN CENTRAL ILLINOIS

BY

BETHFLITZ

THESIS

SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS

FOR THE DEGREE OF

MASTER'S OF BIOLOGICAL SCIENCES

IN THE GRADUATE SCHOOL, EASTERN ILLINOIS UNIVERSITY

CHARLESTON, ILLINOIS

2003

I HEREBY RECOMMEND THAT THIS THESIS BE ACCEPTED AS FULFILLING

THIS PART OF THE GRADUATE DEGREE CITED ABOVE

DATE

DATE

Abstract

Nest site selection in turtles might allow females to increase the survival of their

offspring but little is known about the variables that influence that choice. I examined the

microhabitat variables at nest sites of the eastern box turtle, Terrapene carolina carolina,

at a Boy Scout camp in Shelby County, Illinois, to determine if females chose nest sites

that differed from random sites within the camp. Nest sites differed from random sites in

vegetation height and composition, percentage ground and canopy cover, and light

intensity. All females placed their nests in open habitat within the camp or a pasture

bordering the camp. I suggest that, due to the importance of incubation temperature on

hatchling development, open sites were selected due to the higher incubation

temperatures they provide.

11

Acknowledgments

I would like to acknowledge the following people for their help in the completion

of this project: Dr. Stephen Mullin, Dr. Robert Fischer, Dr. Eric Bollinger, Theron

Tinker, Nelda Foster, the Boy Scouts of America, Lincoln Trails Council, Jacques Nuzzo,

Jane Seitz, Cora Allard, Crystal Flitz, and Sara Denham. Partial funding of this project

was provided by Eastern Illinois University, the Illinois Department of Natural

Resources, and the Chicago Herpetological Society.

1ll

Table of Contents

List of Tables v

List of Figures VI

Introduction 1

Literature Review 2

Materials and Methods 4

Results 8

Discussion 10

Conclusions 16

Bibliography 19

Appendix I 26

IV

List of Tables

Table 1. Mean and standard deviation values for carapace length (CL), plastron length

(PL), carapace depth (CD), and mass of eastern box turtles (Terrapene c. carolina)

measured at Rhodes-France Boy Scout Camp, Shelby County, Illinois, during Summer

2001.

Table 2. Kolmogorov-Smimov test results for features of nest sites (NS; n = 24) and

random sites (RS; n = 75) at Rhodes-France Boy Scout Camp, Shelby County, Illinois,

during Summer 2001.

v

List of Figures

Fig. 1. Occurrence of nesting in eastern box turtles (Terrapene carolina carolina) with

precipitation events at Rhodes-France Boy Scout Camp, Shelby County, Illinois in

Summer 2001.

Vl

Errata

1. The sentence beginning at the bottom of page 8 of this thesis should read as follows:

Half of the females (50 % ) constructed nests during or within 24 hours of a

precipitation event but there was no relationship between the occurrence of

rainfall and nesting activity <-x2 = 2.06, p = 0.72).

2. Figure I shown on page 11 of this thesis should be replaced with the following:

I precipitation (mm) • nest events x 100 6

• x ...5

1 75-

x ... 4 -a j = 50- x x xx ... 3 ~ it

·f • x ...2 i !lo •

25- • x • • ...1

• • 0 -- - • - il 0

I I I I - I I I I - I

150 155 160 165 170 175 180 185 190 195 200

Julian date

Fig. 1. Occurrence of nesting in eastern box turtles (Terrapene c. carolina) with

precipitation events at Rhodes-France Boy Scout Camp, Shelby County, Illinois in

Summer 2001. Each dot in the figure, even those near the origin, represent

precipitation having occurred.

3. The reader should interpret with caution, statements in the Discussion section concerning

the relationship between precipitation and nesting activity in eastern box turtles.

Introduction

Patterns of habitat selection are an important aspect of an animal's natural history

because of the potential impacts on individual fitness (offspring production, survivorship,

etc.). One critical component of habitat selection among oviparous taxa is nest site

selection. Developing embryos at selected sites often have a greater chance of survival

than those at random sites (Wilson 1998); thus, natural selection should favor the

evolution of nest-site selection (Schwarzkopf and Brooks 1987, Temple 1987, Shine &

Harlow 1996, Kolbe & Janzen 2002). Research has shown that many species of turtles

choose non-random sites for egg placement (e.g., Malaclemys terrapin [Burger and

Montevecchi 1975], Kinosternon baurii [Wilson 1998], Chelydra serpentina [Kolbe &

Janzen 2002]). The early life history stages of turtles are typically characterized by the

absence of post nesting parental care and high egg and hatchling mortality rates (Keller et

al. 1997). The choice of a nest site is important because it will influence the likelihood of

predation of both eggs and hatchlings as well as affect the temperature and moisture

content at which the eggs develop (Dodd 2001 ). The microhabitat (primarily soil

moisture content and temperature) surrounding the nest site has been shown to affect the

developmental rate (Cagle et al. 1993), the sex of offspring (Wilhoft et al. 1983), and

some fitness characteristics of the hatchlings like size and growth rate (Burger 1976,

Packard et al. 1985, Janzen 1993).

Gathering a sufficient amount of data on turtles to test life history theory has been

relatively slow and difficult, in part due to their longevity and secretive nature, and the

logistical difficulties when sampling during some or most of their life stages (Iverson

1990). However, box turtles have the potential to provide researchers with a large amount

1

of data on how reproductive traits affect the life histories oflong-lived organisms (Dodd

2001). With rare exception (Congello 1978, Nieuwolt 1996), the current understanding

about these aspects of turtle ecology is based on aquatic or semi-aquatic species (Tinkle

et al. 1981, Walker and Parmenter 1990, Brewster and Brewster 1991, Congdon et al.

1994, Wilson 1998). In this study I sought answers to the following questions: 1) Do

female eastern box turtles, Terrapene c. carolina place their eggs at sites at different

proportions than the available habitat? 2) What are the microhabitat variables of the nest

that differ from random sites?

Literature Review

Terrapene c. carolina is a small terrestrial turtle with a hinged plastron and a

keeled, high-domed carapace. Females can be distinguished from males by the usual lack

of red pigmentation in the eye, flat plastron (the male plastron is concave), and shorter,

more slender tails. Terrapene c. caro/ina is predominantly an open woodland species,

ranging from New England south to Georgia and west to Michigan, Illinois, Tennessee,

and northern Alabama. It is omnivorous, eating a wide variety of plants, small

invertebrates, and carrion (Ernst et al. 1994).

Box turtles are long-lived, sexual maturity is late, and annual reproduction is low.

Sexual maturity is reached in both sexes at 7-10 years and females usually produce only

one clutch of eggs a year, but may produce more, especially in captivity and in some

southern states (Dodd 2001). Courtship and mating usually begins in May and extends

through the summer into October (Ernst et al. 1994). Females are capable of storing

sperm in seminal receptacles, and have been known to produce fertile offspring up to four

2

years after copulation with a male (Ewing 1943). Nesting occurs from early May to

August depending on the latitude but generally occurs in June.

The exact criteria by which female turtles choose nest sites are unknown but

studies have shown that in some species, females will travel great distances from their

home ranges to nesting sites (Stickel 1950, 1989, Legler 1960, Landers et al. 1980,

Congdon et al. 1983, Diaz-Paniagua et al. 1995, Hall et al. 1999). Females also show

nest site fidelity, choosing locations within a few meters of previous years' sites

(Congdon et al. 1983, Stickel 1989). Nesting activity by female Terrapene begins in the

evening hours (1700-1900 h) and takes two to six hours to complete (Congello 1978).

Clutch size ranges from one to eleven eggs but averages four to five (Congello 1978,

Warner 1982, Ernst et al. 1994). Incubation times reported in the literature range widely

(e.g., 57-136 days; Allard 1948, Iverson 1977, Congello 1978) due to the fact that

developmental rate is temperature dependent (Ernst et al. 1994, Dodd 2001).

There are a number of studies that examined either habitat selection or

reproduction in Terrapene. Reagan (1974) studied habitat selection in the three-toed box

turtle (T. c. triunguis) but, except for the brief mention of an observation of a nesting

female, did not examine the microenvironment of nest sites. Nieuwolt (1996) examined

microhabitat selection in the western box turtle (T. ornata luteola), but did not include

nesting activity or nest-site selection. The same author later used reproduction in T.

ornata luteola to test optimal egg size theory (Nieuwolt-Dacanay 1997) but nest site

variables were not examined in this study.

Due to the relative difficulty in locating nesting females, most of the studies on

Terrapene nesting behavior have focused on captive populations (Bissett 1968, Riemer

3

1981, Messinger and Patton 1995). A five year study of nesting in captive T c. triungis

(Messinger and Patton 1995) described nesting behavior and nest site selection. Most

clutches were laid in open areas in direct sunlight but a few were laid in areas that never

received sunlight. However, the authors noted that the small size of the study area (31 x

34 m) limited the variety of nesting sites. Cahn (193 7), Allard (193 5), Bissett (1968),

Riemer (1981), and Ernst et al. (1994) all provide mostly anecdotal data on nesting

behavior and nest site characteristics. Congello (1978) provides a detailed description of

nesting and egg laying behavior in both wild and captive T c. carolina. The study found

that selection took place from 1700-1900 h and occurred in a cleared area where sunlight

could penetrate the canopy. Nesting activity always corresponded with overcast or rainy

days and the author also suggested a correlation of nesting activity to the lunar cycle

(Congello 1978).

Materials and Methods

Study site -The study was conducted at the Rhodes-France Scout Camp (RFSC), in

western Shelby County, Illinois. RFSC is 200 ha of mainly oak-hickory forest, 40 ha of

which are used by the scouts, who have placed some permanent structures and created

various open areas within the forest for campsites. The area is not open to the public and

has minimal human activity except for the months of June, July, and August, when

summer camps are held. Sections of the camp have been periodically logged as recently

as 30 years ago. The camp is bordered on the north and west by agricultural fields (com,

soybean, alfalfa) and on the south and east by a grazed cattle pasture. Because of the size

of the camp and limited manpower, I restricted my searching to the eastern third of the

4

camp, which contained all of RFSC camping sites and was therefore more accessible by

vehicle.

Methods - Beginning in April 2001, I located turtles by walking through RFSC and

searching. For each turtle encountered I recorded sex, mass(± 1 g), carapace length (CL,

± 0.01 mm), plastron length (PL,± 0.01 mm), shell depth (measured at plastral hinge to

top of carapace,± 0.01 mm), any unique features (bum scars, bite marks, old wounds,

etc.), time of day, and location where found. I then filed a set of unique notches into the

marginal scutes of the carapace (each scute being assigned a numeric value) for future

identification. Sex was determined based on eye color, plastron concavity, and relative

tail length and hind limb thickness as described by Cahn (1937). Any female found that

was greater than 120 mm CL had a radio transmitter glued to the right posterior of the

carapace using rubber silicone sealant (Advanced Telemetry Systems model 7PN). I then

released each turtle at its site of capture.

Based on records in the literature (Ernst et al. 1994, Phillips et al. 1999), I

assumed that oviposition would begin anywhere from late May to early June. Therefore,

I chose the first week of May 2001 to relocate all females carrying transmitters and took

them to a veterinarian's office to be radiographed to determine gravidity. I removed the

transmitters from females lacking eggs and released them at their site of relocation. For

gravid females, I recorded clutch size and then attached a thread trailer with rubber

silicone sealant on their carapace on the side opposite the transmitter. The thread trailer

was constructed from a short length of polyvinyl chloride (PVC) pipe with caps at each

end. A small internally wound thread bobbin was placed inside the pipe. A small hole

was drilled in the middle of one of the caps for the thread to pass through. The trailer

5

was glued to the right posterior side of the turtle's shell with the cap free, to allow the cap

to be removed for replacement of the thread bobbin. The total mass of the thread trailer

with bobbin and the transmitter was 4 7 g (about 10% of the average body weight of an

adult female). Studies have shown that turtles can accommodate an attached load of this

magnitude without significant impairment of their locomotion (Zani and Claussen 1995,

Marvin and Lutterschmidt 1997) so it is unlikely that thread trailers altered behavior or

movement. Once the trailer was attached, I again recorded the turtle's mass and then

released her at point of relocation. I also flagged the location and tied the string from the

trailer to the flag.

Turtles were relocated daily. For each relocation, I recorded the position, activity

when found, and mass. Locations were described using visual landmarks and a flag was

placed at the site (it is unlikely the flag attracted predators to the turtle as no instances of

predation on an marked adult turtle were recorded in this study). Latitude and longitude

were later recorded for each relocation site using a global positioning system (GPS) unit

(accuracy of 15 m). Descriptions of activity when found were based on Dodd et al.

( 1994) and were recorded as either walking, feeding, resting while exposed, resting while

buried under cover, or engaging in nesting behaviors (digging, oviposition, covering nest,

etc.). Box turtles generally nest in the evening hours (Allard 1935, Legler 1960, Dodd

2001) so I always located females between 1700 and 2100 h to increase the chance of

locating a turtle while nesting. I also searched both forested and cleared areas for nesting

activity by unmarked females. After 1700 h, a female was either searching for a nest, in

one of the stages of nesting, or in a "form," a shallow depression in the soil or leaf litter

6

in which the turtle spends the night (Stickel 1950). Locating females in the evening

hours made it relatively easy to determine if a female would lay eggs that night.

If a female was active when found, she was observed from a distance (to prevent

her spotting me and potentially abandoning a nesting attempt) every half hour until

nightfall to determine if she was nesting. When possible I continued observing the

female until oviposition was completed. If this was not possible, I marked the nest site

with a coin and returned to the site the next morning. If the female was found in a form, I

checked the thread trailer and then recorded her mass to make sure she had not laid eggs

earlier in the day. Because each egg weighs approximately 10 g, and there are several per

clutch, abrupt weight reductions (> 30 g) would indicate if eggs have been laid (Doroff

and Keith 1990). If a female had laid eggs without my observation of the occurrence, I

followed the path left by the thread trailer to search for the nest. Nesting dates were

recorded and later compared with precipitation data for Pana (approximately 14 km away

from RFSC), Christian County, Illinois received from the Midwest Climate Center.

If the nest was still intact when located, I recorded the number, size, and mass of

the eggs, and then protected the nest from future predation attempts by placing a circular

cage, 30 cm in diameter, of 1.3-cm hardware cloth over the top and down 15 cm into the

soil around the nest (Graham 1997). Relative clutch mass was also determined for each

female with an intact nest.

On 30-31July2001, I quantified the vegetation structure of the sites using

methods similar to those of Wilson (1998). A one-meter square area centered on each

nest site was visually estimated for percentage bare ground, herbaceous plants, woody

plants, and leaf litter. Vegetation height was measured to the nearest centimeter with a

7

ruler and assigned to a category (0-10 cm, 11-20 cm, 20-30 cm, etc.). I also measured

canopy cover using a spherical densiometer, and light intensity(± 1 lux) with a hand-held

light meter. I then recorded the same measurements for 75 random sites. I located

random sites by first making an east and west border on each side of my study area. I

then started at the western side and made a series of parallel lines, spaced 15 m apart, that

extended from west to east. At the starting point of each of these parallel lines, I then

moved eastward across the site, pacing random distances (determined by pulling two

numbers, 0-9, out of a cup) until I had reached the other side, marking each randomly­

located site with a flag. The categories from nesting and random sites were then analyzed

using a principle component analysis (Ludwig & Reynolds 1988).

Results

A total of 117 turtles of both sexes were observed in this study. Sixty-three males

and 35 females were found and marked, giving a male: female sex ratio of 1.8:1.0. I also

found eight juveniles(< 120 mm CL) and recorded eight hatchlings from three successful

nestings. Morphometric data for these individuals are reported in Table 1.

Of the 35 female turtles observed, ten had transmitters attached to their carapace

and were radiographed, eight of which were gravid. Clutch size was determined from the

radiographs and ranged from one to seven eggs (mean= 3.88 eggs). However, only four

of these turtles nested. The remaining four females were tracked until September and

then radio graphed again. No eggs were visible in any of these individuals.

I located twenty-four nests, all through direct observation of nesting activity, from

both marked and unmarked females. Most of the females (88%) nested within 24 hours,

8

\0

Table 1. Mean and standard deviation values for carapace length (CL), plastron length (PL), carapace depth (CD), and mass of eastern

box turtles (Terrapene c. carolina) measured at Rhodes-France Boy Scout Camp, Shelby County, Illinois, during Summer 2001.

Sex CL (mm)

Males (n = 63) 139.20 ± 10.59

Females (n = 35) 134.81±13.91

Hatchlings (n = 8) 32.02 ± 0.87

PL(mm)

128.49 ± 7.01

128.14 ± 9.00

29.87 ± 1.79

CD(mm)

65.17 ± 8.34

66.78 ± 4.12

14.06 ± 1.82

Mass (g)

468.95 ± 78.21

468.29 ± 108.01

8.00 ± 0.00

before or after, a precipitation event (y_2 = 4.46, p ~ 0.05; Fig. 1), and laid their eggs in

either open areas (campsites or unpaved roadways) within the camp or in grazed

meadows bordering the camp. Each female from the three intact nests I recorded had a

relative clutch mass of 0 .10 (following Shine and Schwarzkopf 1992). The principle

component analysis applied to the measured habitat variables resulted in 98.8% of the

variance being explained by one component. Nest sites differed from random sites in

percent woody vegetation, bare ground, leaf litter, canopy cover, vegetation height, and

light intensity (Kolmogorov-Smirnov tests, p < 0.02; Table 2). When compared to

random sites, nest sites are characterized as having shorter vegetation, less leaf litter,

more bare ground, less woody vegetation, less canopy cover and higher light intensity

than random sites.

Of the twenty-four nests that were observed, 21 were destroyed by predators

within 72 hours of oviposition, with 85.7% (18 nests) of this predation occurring within

24 hours. The three nests I located intact and protected with hardware cloth from

predators each had hatchlings emerge from them in September.

Discussion

Body size - The morphometric data I collected in this study is typical of T. c. carolina.

Males in this taxa are on average larger and reach greater maximum sizes than females

(Ernst et al. 1994, Dodd 2001 ). The best explanation for this difference is that larger

males have an advantage over smaller males in copulation because the plastron concavity

of larger males allows a better fit against the female's carapace (Dodd 2001 ).

10

I precipitation (mm) • nest events x 100- 6

• x 5

75-

i x 4 ....,

~ 3 ;I 50- x x xx 3 t ·i 1 ·g • a x • 2

25-• x • •

• • 0- ·- - • - .. ... 0 150 155 160 165 170 175 180 185 190 195 200

Julian date

Fig. 1. Occurrence of nesting in eastern box turtles (Terrapene c. carolina) with

precipitation events at Rhodes-France Boy Scout Camp, Shelby County, Illinois in

Summer 2001. Each dot in the figure, even those near the origin, represent precipitation

having occurred.

11

Table 2. Kolmogorov-Smimov test results for features of nest sites (NS; n = 24) and random sites (RS; n = 75) at Rhodes-

France Boy Scout Camp, Shelby County, Illinois, during Summer 2001. Means are reported± 1 SD.

%woody % bare % leaf Light intensity % canopy Vegetation Variable vegetation ground litter (lux) cover height (cm)

Mean (NS) 0.0 ± 0.0 0.39 ± 0.36 0.03 + 0.05 656.7 ± 399.3 0.26 ± 0.39 8.3 ±4.8

Mean (RS) 0.11±0.2 0.15±0.24 0.27 ± 0.29 138.7 ± 216.2 0.71±0.38 37.4 ± 46.3

- Chi square 21.734 9.601 15.501 31.202 19.918 18.670 N

P-value <.0001 .0165 .0009 <.0001 <.0001 .0002

Lack of egg laying by gravid females - Four of the eight females I tracked using radio

transmitters never oviposited. All eight females showed eggs in radiographs taken in

May 2001 and were tracked continuously from May until September. During that time

period I observed no nesting activity and did not observe a weight loss of more than 30 g

in any of these four females, so I must assume that the eggs were laid in increments, less

than three at a time, at different sites. One possibility for its cause is that my nightly

inspections may have stressed the turtles enough to either delay or even prevent

oviposition. Female turtles have been known to abandon nesting activity if they suspect

the presence of a predator (Dodd 2001 ). While I cannot rule out my disturbances as a

cause, the fact that the other four females successfully laid eggs makes this possibility

unlikely.

Nest site selection - All nesting females laid their eggs at sites that differed from random

sites in regard to each variable examined except percent herbaceous cover. All females

placed their nests in open areas within RFSC or within a meadow next to the camp's

boundaries. Previous studies suggest that sunnier areas free of debris may be preferred

by nesting females (Cahn 1937, Legler 1960, Congello 1978, Messinger and Patton

1995). The reasons for this preference are many. For one, although ground temperatures

were not recorded, it is logical to assume that areas with a higher light intensity and less

canopy cover would be warmer than sites with lower light levels and high canopy cover.

Incubation temperature is important because it affects the sex of offspring, developmental

rate, and can possibly influence the fitness of the hatchlings (Burger 1976, Packard et al.

1985, Janzen 1993). Besides providing a higher incubation temperature, bare ground that

is free of debris would also provide an area easy to excavate a nest cavity and may play a

13

role in hatchling survival. Studies with hatchling snapping turtles have shown that the

probability of survival increased with decreasing ground vegetation (Kolbe & Janzen

2001). Survival is probably increased because hatchlings emerging in sparsely vegetated

areas move quicker and more continuously than hatchlings emerging in denser

vegetation. In box turtles, bare areas may encourage hatchlings to disperse quickly into

forested habitat and may aid in their movement by providing fewer obstacles to move

through.

Turtles may use soil temperature, soil moisture, slope, vegetation, or even

predation risk when choosing a nest site (Wilson 1998, Weisrock & Janzen 1999, Wood

& Bjomdal 2000, Kolbe & Janzen 2001, Spencer 2002, Morjan 2003). I believe that the

turtles in this study chose this type of micro habitat because of the higher soil

temperatures. However, because the females in this study all began digging from 1800-

2000 h, it is unknown whether they were using light intensity, lack of canopy cover,

density of vegetation, or were sensing changes in soil temperatures when choosing sites.

No nesting activity was observed in the wooded areas of my study site, even

though I spent approximately twice as much time searching in wooded areas as I did open

areas. It could be argued that nesting females were easier to find in the open areas than in

wooded areas and this explains why none were discovered nesting in the forest.

However, the four females I found nesting with radiotransmitters all chose open areas,

and previous studies suggest that an open canopy is a key element in nest site choice and

that females prefer open areas (Cahn 1937, Legler 1960, Congello 1978, Messinger and

Patton 1995). Also, the amount of tree and shrub roots in wooded areas would appear to

make digging a nest cavity difficult if not impossible. For these reasons, I feel I obtained

14

a fairly accurate sample of nesting activity. This idea could be tested in a future study by

including a larger area of the camp. Because of the relative ease in locating and traveling

to sites with turtles, I concentrated my efforts in an area of RFSC that contained

campsites. By observing females in a part of the camp that did not contain campsites one

could compare nest site choice between the two groups. It is possible that the females in

my study readily used the campsites because they were convenient and that females

without these options are forced to choose less favorable sites or travel farther to reach

more favorable sites (see Appendix I).

Most turtles in this study nested within 24 hours of a rain event. This phenomenon

has been recorded by Congello (1978), who suggested that females may use cloud cover

and rising humidity levels to predict favorable nesting conditions several hours before a

rain event. Nesting around a rain event could be favorable in two ways: the precipitation

would soften the ground and make it easier for a female to excavate a nest cavity, and

rain immediately following nesting could wash away the scent of the female from the

nest site, making nest predation less likely. Some authors have reported observing

females voiding their bladders over the nest site (Conant 1938, Legler 1960) but I neither

observed, nor saw evidence of, this activity at any of the nest sites in this study.

Nest predation- Of the 24 nesting events observed, 21 ended in predation of the nest

(most of these within 24 hours). Almost any digging animal will consume turtle eggs,

and predators include raccoons (Procyon lotor), opossums (Didelphis virginiana),

coyotes (Canis /atrans), mink (Mustela vison), skunks (Mephitis mephitis), and foxes

(Vulpes, Urocyon) (Temple 1987). While I could not positively identify the species of

15

animal that destroyed the nests in my study, it is likely that raccoons were responsible for

the majority of predation. Due to their adaptability and lack of natural predators, raccoon

populations in many areas are at an abnormally high abundance (Garrott et al. 1993).

This same problem exists at RFSC (pers. obs.), which has an extremely high raccoon

population (trapping and removal has been attempted with little success; Theron Tinker,

pers. comm.). The problem of nest predation at this site may even be exacerbated by the

fact that young campers often leave food items strewn about their campsites. The

raccoons are drawn to the campsites in search of food left by campers and can easily

encounter nesting females and eggs. Given this pattern of land use, female box turtles at

RFSC are placing their eggs in areas that are less than ideal for survival.

Conclusions

Female box turtles in this study chose nest sites that differed from random sites in

percent woody vegetation, bare ground, leaf litter, light intensity, canopy cover,

vegetation height, and light intensity. Light intensity and ease of digging in the bare

ground are the most likely factors affecting female choice, but it is not known exactly

how nesting females are detecting or comparing these factors. Females appeared to

prefer to nest either shortly before, during, or immediately following a rain event. The

fact that all the females in this study nested in man-made clearings (campsites, roads, or

meadows) is important for two reasons. First, it indicates that the use of artificial nesting

areas or structures may have a potential use in helping increase turtle numbers for

populations threatened by habitat loss. Conversely, nesting in man-made areas may

actually be producing a negative effect on turtle populations by encouraging females to

16

nest in less than ideal sites. Another issue threatening the turtles in this study is nest

predation, and must be considered if creating a management plan for this species at

RFSC.

Populations of many Terrapene species have been steadily declining in the past

50 years (Stickel 1978, Williams and Parker 1987, Doroff and Keith 1990, Schwartz and

Schwartz 1991, Dodd and Franz 1993, Belzer and Steisslinger 1999), mainly due to

collections for the pet trade and the influences of urbanization and agriculture (Butler and

Sowell 1996). The fact that the annual survivorship of either turtle eggs and hatchlings is

lower than those of later life stages (Iverson 1991) makes understanding these early

stages a vital component in understanding and conserving a species. The use of habitats

for nesting that are different from habitats utilized during other parts of the activity

season also has important conservation implications. If box turtles' home ranges are

protected, but not their nesting sites, the loss of quality nest sites may lead to decreased

egg and hatchling survival (Wilson 1998, Tucker and Paukstis 1999). The division of

home ranges and nesting sites by roads or unsuitable habitat may have the added effect of

increasing the mortality of gravid females as they move between sites (Stickel 1978, Hall

et al. 1999). Increasing fragmentation of turtle habitat may also increase rates of nest

predation (Temple 1987).

Terrapene. c. carolina is still considered a common species in North America, yet

little is known on its status in the wild (Dodd and Franz 1993, Hall et al. 1999). A lack of

basic ecological information on many turtle species, including data on nest site selection,

has hindered efforts to interpret population declines or recommend management-related

research (Landers et al. 1980, Dodd and Franz 1993). It is only by establishing

17

population and life history data on a species while it is still common that wildlife

managers will have the information necessary to initiate conservation actions when

declines become apparent.

18

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25

APPENDIX I

26

Daily Movement and Activity of Gravid Female Box Turtles in Central Illinois

Abstract

I examined the daily movements of 5 female eastern box turtles, Terrapene c.

carolina. Turtles were fitted with a radiotransmitter and thread trailer and were relocated

daily. Using thread trailers I measured daily movements and compared distances moved

to daily temperature, rainfall, and relative humidity. In contrast to previous studies, I

found no correlation between daily movements and any of these meteorological factors.

These findings may be due to the small sample size in this study, or the condition of the

females being followed.

Introduction

Box turtles are, with rare exception (Dodd 2001), diurnal or crepuscular. As

ectothermic animals, box turtles are influenced by exogeneous factors that may affect

their movements. Studies have found that box turtles have periods of activity and

inactivity throughout the day, and these may depend on season (i.e., feeding and

reproduction [Dodd 2001]), temperature (Legler 1960, Nieuwolt 1996, Schwartz and

Schwartz 1974), rainfall (Stickel 1950), and relative humidity (Reagan 1974, Stickel

1950). Turtles are most active when warm temperatures (20°-30°C), high humidity (60%

+),and frequent rainfall occur (Stickel 1950, Strang, 1983, Dodd 2001). When studying

an animal's daily activities and movements, knowing its specific path taken provides

information on types of habitat utilized by the animal, the effect of habitat structure or

landscape features on an animal's movements, distance traveled per day, seasonal

27

changes in movement patterns, and the effect of meteorological conditions on movements

(Claussen et al. 1997).

There are at least three ways in which an animal's movements can be studied:

mark-recapture, radio telemetry, and thread trailing. Mark and recapture provides the

cheapest, least labor intensive approach but recapture rates can be low and the interval

between captures is often unpredictable (Claussen et al. 1997). This technique also gives

no information about the path taken between successive capture points. Radiotelemetry

is a high-cost method that can provide many more location points and is useful in

providing information on long-term habitat use and movements but again, it cannot show

the exact pathway the animal took between relocation points. Thread trailing is a more

time consuming method to track animals but, with the exception of extremely

sophisticated satellite-tracking devices that continuously monitor an animal's position, is

the best option to obtain information about the specific path an animal takes.

The turtle's hard carapace makes them excellent subjects for the attachment of

thread trailers and many studies have used this method (Breder 1927, Legler 1960,

Schwartz and Schwartz 1974, Hailey 1989, Diaz-Paniague et al. 1995, Claussen et al.

1997). Studies have shown that turtles can accommodate an attached load without

significant impairment of their locomotion (Zani and Claussen 1995, Marvin and

Lutterschmidt 1997) so it is unlikely that thread trailers alter behavior or movement.

While turtles may take a seemingly direct path when moving to a specific

location, thread-trailing studies have shown that box turtles often leave meandering trails

that tum, move around objects like trees, and often repeatedly cross their own paths and

those of other turtles (Dodd 2001, Stickel 1950). The distance measured between two

28

capture points may be considerably shorter than the actual distance traveled by the turtle.

For example, Stickel (1950) found that a box turtle in Maryland traveled 139 min one

day but was only 52 m from its last point of capture. When examining the effects of

environmental factors on daily distance moved, it is essential that the actual distance

moved is recorded. Knowing the actual distance moved and the path taken by a turtle

reveals valuable information, especially when developing a conservation management

plan, on how much area and what types of habitat the animal is actually using. For

instance, comparisons of paths and movements between differing box turtle populations

might provide insights into the possible effects of human disturbance on the species. In

this study I sought answers to the following questions: 1) How far does a box turtle

move in a day? 2) Is distance traveled have any relationship to temperature, rainfall,

and/or relative humidity?

Materials and Methods

The data in this study were collected at the Rhodes France Scout Camp (RFSC)

located in western Shelby County, Illinois, from gravid female T c. carolina that were

being tracked with both radio transmitters and thread trailers for the purpose of locating

nest sites (see above for descriptions of the trailers and methods). Thread trailers were

used from 16Juneto31 July2001(Juliandays165-215). Irelocatedturtlesdailyand

recorded mass, activity when found, and location where found (location was recorded by

using visual landmarks and later recorded in latitude and longitude using a global

positioning system [GPS]). I recorded the path taken by following the thread trail that

had been laid out by the turtle from that day's movements, and then collected the thread

29

and measured it to the nearest centimeter. In this study, I was more interested in total

distance moved then the exact path taken so I did not measure each tum the turtle had

taken, only the total distance. In the event that a turtle would run out her trailer in a 24

hour period, I would record the data as> 175 m moved, which was the total length of

thread in the trailer. Daily measurements were then compared to weather data for Pana

(approximately 14 km from RFSC), Illinois, compiled by the Midwest Climate Center.

Mean values of all five turtles were analyzed using simple regression for mean daily

temperature, daily rainfall, and mean relative humidity.

Results

Daily movement for each turtle varied considerably, from no movement to

movements greater than 175 m, with an average of 64.14 m moved each day. One turtle

was completely inactive for ten days while another turtle moved every day of the study.

Paths were often meandering, circling around trees and crossing each other repeatedly,

with the turtle usually ending her day a short distance from where she had started.

Simple regressions showed no relationship between daily distance moved and daily

temperature (r2 = 0.01, p = 0.61, n = 40) or relative minimum (r2 = 0.02, p = 0.37, n = 39)

and maximum (r2 = 0.01, p = 0.59, n = 39) humidity. A one-way ANOVA did not detect

any relationship between daily distance moved and whether or not it rained that day (F1,Js

= 0.75, p = 0.39).

Discussion

Many people view turtles as slow-moving creatures that lead relatively inactive

30

lives. However, turtles engage in a variety of behaviors and activities in their daily and

seasonal lives. As in other studies (Dodd 2001, Stickel 1950), the turtles in this study

exhibited meandering movements, with a path often circling around objects (one female

continually circled and passed through a camper's tent and around the legs of a cot) and

crossing and recrossing itself. Distance between relocation points could not provide a

true account of the daily distance moved as a turtle was often relocated a short distance

from her last location. One reason for this it that all of the females in this study had a

tendency to have 3 or 4 locations that they used for sleeping, known as forms, and were

repeatedly found at these locations when I checked them in the evening, even though they

had moved a considerable distance away from the form during the day. As an example,

one female moved 67.25 m in one day but was found only 2 m from her last location.

Despite their tendency to meander, occasionally a turtle would lead a very straight

path, often for a considerable distance. Many times a turtle would follow a dry creek bed

or animal path, but other times the turtle would create her own trail through a mix of

vegetation and straight over any obstacles (i.e., fallen trees) in the way. The length of

these straight line paths could be as short as 6 m but were often longer and on 5 occasions

turtles were located 500 m (straight line distance) or more from their previous location.

In these instances, the turtle would always stay in the area for only one or two days and

then the thread trail would show another straight path back to where she had been found

on previous locations. For example, one female that I had located in one of her forms the

previous night was found in the late afternoon of the next day 512 m (the trailer laid a

straight path for 175 m, I then measured a straight line distance from the end of the trailer

to the turtle) away under a multiflora rose (Rosa sp.) in a cow pasture bordering the

31

camp. She was located there the following day but on the third day had made another

straight line path (following almost exactly her previous path) back to the form used on

the first day. My first thought upon encountering her in the pasture was that she had

moved to this location to nest, but no nesting activity was observed and no drop in mass

was recorded (indication eggs had been laid). This behavior has also been observed in a

population of three-toed box turtles (T. c. triunguis) in Texas (J. Koukel, pers. comm.).

While I observed only females, Koukel found that only male turtles would move these

large, straight line distances. The reason for these movements is unknown: previous

studies have shown that female box turtles may move long distances in order to reach a

suitable nesting site (Stickel 1950, Legler 1960). Another hypothesis is that the turtle is

following some olfactory cue (i.e., food source, member of the opposite sex, etc.).

Further study of these movements is important as it may cause a re-evaluation of a

turtle's habitat requirements and home range size.

Unlike previous studies (Stickel 1950, Claussen et al. 1997, Dodd 2001), my

turtles showed no correlation between daily distance moved and daily mean temperature,

occurrence of rainfall, or daily mean relative humidity. These results should be

interpreted with caution, however, as analyses are based on a pseudoreplicated data set

(Hurlbert 1984). Also, my sample size was small and I was recording data from only

gravid females. It is possible that their behavior may have been modified by their

condition and therefore affected the results, (i.e., the demands of egg production and the

motivation to find a nest site for oviposition may have been a larger determinant of

distance moved than meteorological factors). In a study of ornate box turtles (T. ornata),

Legler (1960) found that gravid females had the greatest daily mean distance moved

32

(110.6) as compared to males (88.1) and non-gravid females (68.9). A larger sample size

including males and both gravid and non-gravid females would be required to gain a

better understanding of what factors influence daily movements and activities at RFSC.

Conclusions

Female box turtles at RFSC showed no correlation between daily distance moved

and either daily mean temperature, occurrence of rainfall, or daily mean relative

humidity. These results could be explained either by the small sample size or the fact

that gravid turtles may behave differently than non-gravid or male turtles. Their daily

movements showed meandering paths that covered little straight line distance and ended

in often-utilized resting locations, or forms.

An interesting aspect of this study was the occurrence of lengthy straight line

movements by three of the females to a particular spot and then the direct movement

back to their previous locations a few days later. I believe that investigating the reasons

(i.e., nesting, hibernation, mating, finding a food source, etc.) behind these movements in

the turtle warrants further study for many reasons. If one discounts, or is unaware of,

these long distance movements, the amount of habitat needed by a turtle could be

drastically underestimated. If it is necessary for some turtles to move long distances to

nest, it may shed light on the inappropriateness of captive nest site selection studies.

And, with the ever increasing amount of habitat loss and fragmentation occurring in our

environment, understanding these long distance movements may have important

conservation implications as well.

33