AN ASSESSMENT OF A HERPETOFAUNAL COMMUNITY IN HAMILTON COUNTY,

TENNESSEE: BASELINE ECOLOGY, SPECIES RICHNESS, AND RELATIVE

ABUNDANCE

By

Joseph Frellen Simpson, III

Thomas P. Wilson Timothy Gaudin

Associate Professor of Biology Professor of Biology
(Chair) (Committee Member)

Mark S. Schorr Jeffrey Elwell

Professor of Biology Dean, College of Arts & Sciences
(Committee Member)

A. Jerald Ainsworth
Dean of the Graduate School
AN ASSESSMENT OF A HERPETOFAUNAL COMMUNITY IN HAMILTON COUNTY,

TENNESSEE: BASELINE ECOLOGY, SPECIES RICHNESS, AND RELATIVE

ABUNDANCE

By

Joseph Frellen Simpson, III

A Thesis Submitted to the Faculty of the University of Tennessee at Chattanooga

in Partial Fulfillment of the Requirements of the Degree of
Master of Science in Environmental Science

The University of Tennessee

at Chattanooga

May 2013

ii
 ABSTRACT

Long-term investigations are needed to adequately assess herpetofaunal community

structure and dynamics, and habitat alteration remains the most critical threat to these

communities. Herein, I report on a 32-month study investigating species richness, abundance,

sex ratios, and body sizes of amphibians and reptiles in a Hamilton County, Tennessee, wetland.

Utilizing drift fencing in conjunction with pitfall arrays, a total of 14 reptile species and 16

amphibian species was recorded. Evenness was low among all study years due to large sample

sizes of ambystomatid salamanders relative to all other species. Body sizes of Ambystoma were

larger in females, and sex ratios of all Ambystoma were significantly male-biased in all study

years except one female-biased sample of Ambystoma opacum (2009). This study is the first

wetland community assessment for the southeastern Tennessee region and provides baseline data

for future comparisons regarding changes in community structure and dynamics.

iii
 ACKNOWLEDGEMENTS

I am greatly indebted to the numerous undergraduate and fellow graduate students who

have given much time and effort to aid the progress of my project while working with Dr.

Wilson as independent-studies students and with the BIOL 3070 and ESC 3400 courses. Their

man-power has undoubtedly increased the scope and success of this study. Specifically, I would

like to express gratitude to Brett Warren, Jake van Thomison, Kari Baker, Daniel Armstrong,

Dan Huser, Evan Collins, Phil Brown, Jason Hill, Mary Bratton, Andy Shoulders, Ashley Smart,

Charity Hammett, Robert Kennedy, Curtis Cole, “Team Salamander”, and others that I have

likely forgotten but appreciate no less. I especially thank Brett Warren and Jake van Thomison

for help installing the initial sections of drift fence. Daniel Armstrong provided much needed

support during the last year of this study, both in the field and the lab. I am indebted to Adam

Reynolds for his work in delineating the wetland. I am grateful for GIS support from Jon Oakley

and Andy Carroll. I would like to thank committee members Drs. Mark Schorr and Tim Gaudin

for their reviews and guidance. I am indebted to the Department of Biological and

Environmental Sciences at UTC for excellent courses and invaluable experience while serving as

a teaching assistant. Finally, I would like to thank Dr. Thomas Wilson, my advisor and

committee chair, for his daily oversight and tireless efforts both in the field and the lab. I am

also grateful to Dr. Wilson for his personal funding and allowing my generous use of his field

equipment.

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 TABLE OF CONTENTS

ACKNOWLEDGEMENTS ............................................................................................... iv

LIST OF TABLES ............................................................................................................ vii

LIST OF FIGURES ........................................................................................................... ix

LIST OF ABBREVIATIONS ............................................................................................ xi

LIST OF SYMBOLS ........................................................................................................ xii

CHAPTER

I. GENERAL INTRODUCTION ....................................................................1

Introduction ..................................................................................................1
Primary Study Organisms ............................................................................4
Study Area ...................................................................................................6
Research Objectives .....................................................................................6
General Methods ..........................................................................................7

II. RELATIVE ABUNDANCE AND SPECIES DIVERSITY AT LT6 .......14

Abstract ......................................................................................................14
Introduction ................................................................................................15
Materials and Methods ...............................................................................17
Results ........................................................................................................18
Discussion ..................................................................................................20
Implications for Conservation....................................................................24

III. SEX RATIOS OF AMBYSTOMA AND BODY SIZES OF

AMPHIBIANS AND REPTILES PRESENT IN LT6 ........................43

Abstract ......................................................................................................43
Introduction ................................................................................................44
Materials and Methods ...............................................................................45
Results ........................................................................................................47
Discussion ..................................................................................................48
v
 Implications for Conservation....................................................................50

IV. POPULATION ESTIMATES AND PHENOLOGY OF AMBYSTOMA

MACULATUM AND A. OPACUM .....................................................56

Abstract ......................................................................................................56
Introduction ................................................................................................57
Materials and Methods ...............................................................................58
Results ........................................................................................................60
Discussion ..................................................................................................61
Implications for Conservation....................................................................62

V. DIRECTIONS FOR THE FUTURE ..........................................................67

REFERENCES ..................................................................................................................70

VITA ..................................................................................................................................79

vi
 LIST OF TABLES

1.1 Amphibian species likely to occur in Hamilton County, Tennessee, compiled from
Conant and Collins (1998) .......................................................................................9

1.2 Reptile species likely to occur in Hamilton County, Tennessee, compiled from
Conant and Collins (1998) .....................................................................................10

2.1 Amphibian species likely to occur in Hamilton County, TN compiled from Conant
and Collins (1998). Species confirmed in Miller (1996), Redmond and Scott
(1996), Wilson (unpubl. data), and/or this study are noted ...................................27

2.2 Reptile species likely to occur in Hamilton County, TN compiled from Conant
and Collins (1998). Species confirmed in Miller (1996), Redmond and Scott
(2008), Wilson unpubl. data, and/or this study are noted ......................................28

2.3 Total amphibian captures (recaptures excluded) by individual study year and the
entire study period (Sept. 2007-May 2010) ...........................................................29

2.4 Total reptile captures (recaptures excluded) by individual study year and the
entire study period (Sept. 2007-May 2010) ...........................................................30

2.5 Species richness comparisons from previous studies and this study (Sept. 2007-
May 2010) ..............................................................................................................31

2.6 Species richness and heterogeneity measures of LT6 for the entire study (Sept.
2007-May 2010) and by study year (Year 1 = Sept. 2007-Sept. 2008, Year 2
= Oct. 2008-Sept. 2009, and Year 3 = Oct. 2009-May 2010) ...............................32

2.7 Jaccard’s index and percent similarity values between study years (Year 1 =
Sept. 2007-Sept. 2008, Year 2 = Oct. 2008-Sept. 2009, and Year 3 = Oct.
2009-May 2010).....................................................................................................33

3.1 Sex ratios for A. maculatum and A. opacum reported in previous studies
and this study (Sept. 2007-May 2010) ...................................................................52

vii
3.2 Relative biomass (g) of species captured (Sept. 2007-May 2010) and
corresponding percent biomass ..............................................................................53

3.3 Snout-vent length (mm) (mean ± SD) and tail length (mean ± SD), and
body mass (g) (mean ± 1SD) of salamander species (Sept. 2007-May 2010) .......54

3.4 Snout-vent length (mm) (median ± IQR) and body mass (g) (median ± IQR)
of anuran species (Sept. 2007-May 2010) .............................................................55

4.1 Population estimates of Ambystoma opacum and A. maculatum for each

study year (Sept. 2007-May 2010).........................................................................64

4.2 Phenology of Ambystoma maculatum captures according to date and movement

direction (entering or exiting the wetland; recaptures included). Percentages
of the total adult captures (per study period) are also given by month as an
indicator of seasonal activity .................................................................................65

4.3 Phenology of Ambystoma opacum captures according to date and movement

direction (entering or exiting the wetland; recaptures included). Percentages
of the total adult captures (per study period) are also given by month as an
indicator of seasonal activity .................................................................................66

viii
 LIST OF FIGURES

1.1 A gravid female Ambystoma maculatum (top plate) exhibiting increased girth due
to the presence of eggs, and a male A. maculatum (bottom plate) exhibiting
swollen cloacal lips (indicated by arrow) ..............................................................11

1.2 A gravid female Ambystoma opacum (top plate) exhibiting darker crossbands
and increased girth due to the presence of eggs, and a male A. opacum (bottom
plate) exhibiting bright white crossbands and swollen cloacal lips (indicated
by arrow) ................................................................................................................12

1.3 Study area map showing LT6 (delineated by white lines) in Hamilton County,
Tennessee ...............................................................................................................13

2.1 General design of the drift fence (horizontal line) and pitfall traps (circles), with
approximately 6.5 meters spacing from center to center of adjacent pitfalls ........34

2.2 A general view of the installed drift fence. A measuring tape (~60 cm in length)
is attached for scale ................................................................................................35

2.3 Vertical view of an installed 18.9 liter pitfall (with lid attached) adjacent to
flashing. A measuring tape (~20 cm in length) is attached for scale ...................36

2.4 Species area curve compiled using previous studies and this study. Studies
represented on the graph include Greenberg et al. 1994 (A), Gibbons et
al. 1996 (B), Miller et al. 2005 (C), multiple sites from Neimiller et al.
2011 (D), Paton et al. 2000 (E), and this study (F) ...............................................37

2.5 Ranked abundances (log10) for all species recorded during the total study period
(Sept. 2007-May 2010). Species represented on the graph include: Acris
crepitans (A. crep.), Acris gryllus (A.g.), Agkistrodon contortrix (A. cont.),
Ambystoma maculatum (A.m.), Ambystoma opacum (A.o.), Anaxyrus
americanus (A.a.), Anaxyrus woodhousii (A.w.), Aspidoscelis sexlineatus
(A.s.), Chelydra serpentina (C.s.), Chrysemys picta (C.p.), Coluber
constrictor (C.c.), Diadophis punctatus (D.p.), Gastrophryne carolinensis
(G.c.), Hyla cinera (H.c.), Hyla versicolor (H.v.), Kinosternon subrubrum
(K.s.), Lithobates catesbeianus (L. cates.), Lithobates clamitans (L. clam.),
Lithobates sphenocephalus (L.s.), Notophthalmus viridiscens (N.v.),
Opheodrys aestivus (O.a.), Plestiodon fasciatus (P.f.), Plestiodon

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 inexpectatus (P.i.), Plestiodon laticeps (P.l.), Pseudacris crucifer (P.c.),
Pseudacris triseriata (P.t.), Scaphiopus holbrookii (S.h.), Sceloporus
undulatus (S.u.), Terrapene carolina (T.c.), and Trachemys scripta (T.s.) ...........38

2.6 Ranked abundances (log10) for all species recorded during the first study
year. Species represented on the graph include: Agkistrodon contortrix (A.
cont.), Ambystoma maculatum (A.m.), Ambystoma opacum (A.o.), Anaxyrus
americanus (A.a.), Anaxyrus woodhousii (A.w.), Chrysemys picta (C.p.),
Coluber constrictor (C.c.), Diadophis punctatus (D.p.), Gastrophryne
carolinensis (G.c.), Hyla cinerea (H.c.), Lithobates catesbeianus (L. cates.),
Lithobates clamitans (L. clam.), Lithobates sphenocephalus (L.s.),
Notophthalmus viridiscens (N.v.), Pseudacris triseriata (P.t.), Scaphiopus
holbrookii (S.h.), Terrapene carolina (T.c.), and Trachemys scripta (T.s.) ..........39

2.7 Ranked abundances (log10) for all species recorded during the second
study year. Species represented on the graph include: Acris crepitans
(A. crep.), Acris gryllus (A.g.), Ambystoma maculatum (A.m.), Ambystoma
opacum (A.o.), Anaxyrus woodhousii (A.w.), Aspidoscelis sexlineatus (A.s.),
Chelydra serpentina (C.s.), Chrysemys picta (C.p.), Coluber constrictor
(C.c.), Gastrophryne carolinensis (G.c.), Hyla versicolor (H.v.), Kinosternon
subrubrum (K.s.), Lithobates catesbeianus (L. cates.), Lithobates clamitans
(L. clam.), Lithobates sphenocephalus (L.s.), Notophthalmus viridiscens
(N.v.), Opheodrys aestivus (O.a.), Plestiodon fasciatus (P.f.), Plestiodon
inexpectatus (P.i.), Plestiodon laticeps (P.l.), Pseudacris crucifer (P.c.),
Pseudacris triseriata (P.t.), Sceloporus undulatus (S.u.), and Terrapene
carolina (T.c.) ........................................................................................................40

2.8 Ranked abundances (log10) for all species recorded during the third study year.
Species represented on the graph include: Acris crepitans (A. crep.), Acris
gryllus (A.g.), Ambystoma maculatum (A.m.), Ambystoma opacum (A.o.),
Anaxyrus woodhousii (A.w.), Coluber constrictor (C.c.), Kinosternon
subrubrum (K.s.), Lithobates catesbeianus (L. cates.), Lithobates clamitans
(L. clam.), Lithobates sphenocephalus (L.s.), Notophthalmus viridiscens
(N.v.), Pseudacris triseriata (P.t.), and Terrapene carolina (T.c.) ........................41

2.9 A performance curve plotting time versus cumulative species richness as a

measure of effort ....................................................................................................42

x
 LIST OF ABBREVIATIONS

AUP, Animal Use Protocol

cm, Centimeters

g, Grams

kg, Kilograms

ha, Hectares

IQR, Interquartile range

m, Meters

mm, Millimeters

SD, Standard deviation

SVL, Snout-vent length

TL, Tail length

TVA, Tennessee Valley Authority

UTC, University of Tennessee at Chattanooga

VAAP, Volunteer Army Ammunition Plant

xi
 LIST OF SYMBOLS

a, alpha

~, approximately

<, less than

>, greater than

n, total number in sample

±, plus or minus

χ2, Chi-square

xii
 CHAPTER I

GENERAL INTRODUCTION

Introduction

It is well known that herpetofauna worldwide are experiencing population declines

(Gibbons et al. 2000, Laan and Verboom 1990), and several factors have been presented as the

underlying cause behind these declines (Collins and Storfer 2003, Gibbons et al. 2000). The

most commonly cited impact is habitat alteration; indeed, forest-dwelling species are

experiencing an especially rapid loss (Stuart et al. 2004). More than 400 amphibian species are

critically endangered, and many species are poorly studied and thus difficult to properly assess

(Stuart et al. 2004). Similarly, reports of reptile declines are increasing (Santos and Gustavo

2009, Whitfield et al. 2007). Arguably, amphibians are more susceptible to habitat degradation

(e.g., pollution) than reptiles due to their physiology (moist, glandular skin and lack of a

calcareous egg) and life history traits (reliance on aquatic habitats); however, both are

inseparably linked through habitat types and thus equally vulnerable to habitat loss (Gibbons et

al. 2000).

Indeed, approximately 75% of all Central American amphibian species monitored since

1970 have shown drastic reductions in population size, and sympatric reptile species were

subsequently found to exhibit the same pattern of decline (Whitfield et al. 2007). This holds true

in the southeastern United States as well, where both anecdotal and scientific evidence of

herpetofaunal declines has been highlighted (Tuberville et al. 2000, Winne et al. 2007). Other

1
species are less-studied—such as the spotted salamander (Ambystoma maculatum) and the

marbled salamander (A. opacum)—and often lack estimates of basic population and life history

parameters because of their secretive nature and the fossorial habits of adults (Rothermel and

Semlitsch 2006).

Habitat alteration is the best documented cause of the aforementioned declines (Alford

and Richards 1999, Hecnar 1998), and may affect both the wetland itself and the surrounding

uplands. Wetlands in particular can experience rapid transformations after an increase in

development and amount of impervious surface (Kentula et al. 2004). Ephemeral wetlands are

critical habitat because some regions of the United States currently contain less than 20% of the

natural wetlands that were present historically (Leja 1998). Wetland alterations in urban areas

often include large changes in water levels (Kentula et al. 2004), increased sedimentation, and

increased contaminants within the wetland (Schueler 1994). Habitat alteration may also decrease

the wetland hydroperiod, which has been shown to be negatively correlated with the number of

metamorphosed juveniles in amphibian populations (Pechmann et al. 1989). Thus, urbanization

affecting wetland dynamics may greatly impact local herpetofaunal communities.

Habitat fragments that result from landscape alteration have been shown to exhibit lower

species richness than continuous forest for both reptiles and amphibians (Bell and Donnelly

2006). Similarly, land-use studies have shown that patches with low wetland permanency

decrease in species richness (Kolozsvary and Swihart 1999, Vallan 2000). Ambystomatid

salamanders in particular are known to decrease in density with deforestation due to their close

association with forested uplands (Gibbs 1998). These uplands provide habitat for adults when

vernal pools are not being utilized for breeding (Lehtinen et al. 1999). Likewise, upland reptiles

are often confined to any remnant vegetation and may be almost completely removed from the

2
cleared landscape (Driscoll 2004). Often, species found in newly formed pools are highly

dependent upon distance to the nearest woodland; also, connectivity between these woodlands is

significantly important for movement between isolated pools (Laan and Verboom 1990). As

expected, any decreased connectivity is negatively correlated with species richness (Lehtinen et

al. 1999). A synergy of factors can further decrease richness, such as smaller fragment sizes,

increased road density, and increased urban land use (Lehtinen et al. 1999). Although

urbanization negatively affects community structure, different amphibian species exhibit

differing degrees of sensitivity (Gibbs 1998). Red-spotted newts (Salamandridae: Notopthalmus

viridiscens) are extremely sensitive to urbanization, whereas spring peepers (Hylidae: Pseudacris

crucifer) can continue to do relatively well in urbanized areas (Gibbs 1998, Lehtinen et al. 1999).

Spring peepers often exhibit a high local density, which may account for increased resistance to

fragmentation (Gibbs 1998).

However, quantifying the local density of reptiles and amphibians in isolated wetlands is

critical because many herpetofaunal breeding sites may be subdivided into demes (Blaustein et

al. 1994). Metapopulation dynamics are important in these fragmented landscapes because many

organisms can move hundreds of meters if necessary to find suitable habitat. The processes of

extinction and recolonization can be vital to maintaining demes (Buskirk 2005, Gibbons 2003,

Marsh and Trenham 2000, Semlitsch et al. 2008). Further, it should be noted that amphibians

show extremely high site fidelity and many species only move short distances if habitat

requirements are met (Blaustein et al. 1994). Thus, recolonization after local extinction may be

difficult or impossible in some areas (Blaustein et al. 1994). Interestingly, species that are

excellent dispersers (e.g., Notophthalmus viridiscens) may be more susceptible to urbanization

than other species that possess little ability to disperse (Gibbs 1998). Areas with smaller

3
woodland connectivity and fragment sizes increase the risk that dispersing organisms will

migrate into unsuitable areas (Gibbs 1998).

Monitoring changes in community composition that result from land-use alteration

requires long-term investigation (see Blaustein et al. 1994, Husting 1965). While richness and

abundance are two critical measurements for successful species management, monitoring sex

ratios and the degree of sexual dimorphism are also useful tools in determining population trends

that may be resulting from both ecological and/or anthropogenic influences. Further,

conservation strategies are best utilized in conjunction with phenology data that incorporates

temporal movements and breeding patterns among species (Paton and Crouch 2002). In

southeastern Tennessee, data are especially limited with regard to the long-term effects of habitat

degradation on herpetofauna, yet the region is a hotspot for amphibian and reptile biodiversity.

The Southeast is home to 50% of the total U.S. herpetofaunal species, with 20% being endemic

to the region (Tuberville et al. 2005). This lends great urgency to rapid community

investigations that bolster conservation efforts.

Primary Study Organisms

Conant and Collins (1998) highlight 31 amphibian species and 40 reptile species that are

likely occur in Hamilton County, Tennessee (Tables 1.1, 1.2). Although all species listed do not

use wetland habitats, adjacent uplands provide shelter and transient movements may result in

incidental captures. However, two salamander species in the family Ambystomatidae—

Ambystoma maculatum (the spotted salamander) and A. opacum (the marbled salamander)—

utilize ephemeral wetlands for annual breeding migrations (Conant and Collins 1998). Both

4
ambystomatid species are characterized by a large body size (adults >70 mm total length) and

long limbs that overlap when pressed against the body (Powell et al. 1998).

The spotted salamander is a large-bodied salamander with a range extending from eastern

Texas and Oklahoma to the Atlantic coast and north to Canada. Adults average 150-250 mm

total length, and are identifiable by the two irregular rows of yellowish spots that run the entire

length of the body (Conant and Collins 1998, Petranka 1998). Reproduction in A. maculatum

typically begins late winter or early spring when adults migrate from the uplands to temporary

ponds or ephemeral wetlands to lay eggs during periods of rainfall (Husting 1965, Petranka

1998). Males are easily identified before breeding by the presence of swollen cloacal lips, which

are absent in females (Figure 1.1). Mature deciduous forests offer optimal habitats for adults, but

populations are known to occur in coniferous forests with vernal pools as well (Petranka 1998).

Ambystoma opacum is a medium-sized ambystomatid with a range extending from

eastern Texas to the Atlantic coast and northward to Connecticut and New York. Adults average

77-127 mm in total length, and are identifiable by a black background color with conspicuous

white or gray crossbands (Petranka 1998). They prefer deciduous forests, and are one of only

two ambystomatid species that mate and oviposit on land, with females nesting near the dried

beds of temporary ponds (Petranka 1998). Marbled salamanders are relatively unusual because

of their fall breeding cycle, which occurs two to three months (September-October) before the

first A. maculatum breeding movements (January-February). Ambystoma opacum is sexually

dichromatic during the breeding season (Figure 1.2), when males exhibit a distinctly brighter

white than females and also exhibit significantly larger areas of white on the body (Noble and

Brady 1933, Todd and Davis 2007).

5
Study Area

Ecological sampling occurred at the former Volunteer Army Ammunition Plant (VAAP)

property (35.104°N, 85.130°W). The University of Tennessee at Chattanooga (UTC) currently

owns a 36.8 hectare parcel where the study wetland is located (Figure 1.3). Located in the Ridge

and Valley ecoregion, its underlying geology is dominated by limestone and dolomite formations

and rolling hills (Griffith et al. 1997). This wetland, herein referred to as ‘LT6’ (an acronym for

‘long-term study site 6’), has an approximate size of 1.82 ha (ranging from 1.1 to ~4-5 ha during

floods) and is surrounded by mixed deciduous and conifer forests. LT6 holds water from autumn

to early summer each year, and is bordered by parcels owned by Hamilton County, Tennessee

Valley Authority (TVA), Volkswagen, and residential entities. Tennessee State Highway 58 and

several subdivisions create the northern boundary. Volkswagen North America began large-

scale construction in 2009 (significant land-clearing began in 2008), and future studies may thus

be able to correlate the increase in deforestation and impervious surface to changes in

community structure first described here.

Research Objectives

1. Determine and evaluate species richness and relative abundance of amphibians and

reptiles in LT6 for comparison to those reported in the literature for similar habitat type

and across years.

2. Determine mean body sizes, sex ratios, and sexual size dimorphisms of Ambystoma

maculatum and Ambystoma opacum in LT6 for comparison to those reported in the

literature and across years.

6
 3. Use mark-recapture methods to generate population estimates of Ambystoma

maculatum and Ambystoma opacum and determine phenology of these species in LT6.

General Methods

Drift fences have proven to be a useful tool in sampling ambystomatid salamanders and

other small vertebrates, and are considered to be the most effective capture method for species

with a well-defined aquatic breeding season (Dodd and Scott 1994). Drift fences used in

conjunction with pitfalls can capture a very high proportion of the population, and provide

precise estimates of population size when combined with mark-recapture techniques (Crosswhite

et al.1999, Halliday 2006). Additionally, drift fences can be used for several objectives which

include determining species richness and abundance (Karns 1986). Pitfall traps are ‘passive’

sampling gear, and can thus capture higher numbers of pond-breeding amphibians over time as

compared to transects or other opportunistic survey methods (Willson and Gibbons 2009). Thus,

the use of pitfalls provides “ecological perspectives” that could be obtained in no other manner

(Gibbons and Semlitsch 1981).

A general protocol was developed utilizing the S.M.A.R.T. (Specific, Measureable,

Attainable, Relevant, and Timely) approach (Cogalniceanu and Miaud 2010), which increases

the ability to conduct a successful study by first determining resources, duration, target species,

and specific objectives. All captured individuals were classified by gender and age class (e.g.,

juvenile or adult) according to secondary sex characteristics and adult body sizes given in Conant

and Collins (1998). Body size was recorded as snout-vent length (mm), tail length (mm), and

body mass (g). Toe-clipping was utilized for anurans and ambystomatid salamanders to aid in

the identification of recaptures (Martof 1953). More, all Ambystoma were photographed and

7
tissue samples were collected from each individual. Turtles were uniquely marked according to

Cagle (1939), and snakes were marked by clipping ventral scutes (Brown and Parker 1976). All

research activities were conducted pursuant to Tennessee Wildlife Resources Agency Permit #

3082 and UTC AUPs 0907-TPW-03 and 0408-TPW-04.

8
 Table 1.1

Amphibian species likely to occur in Hamilton County, Tennessee, compiled from

Conant and Collins (1998).

Species Name Common Name

Acris crepitans Northern Cricket Frog

Ambystoma maculatum Spotted Salamander
Ambystoma opacum Marbled Salamander
Ambystoma tigrinum Tiger Salamander
Anaxyrus americanus American Toad
Anaxyrus woodhousii Fowler’s Toad
Aneides aeneus Green Salamander
Desmognathus fuscus Dusky Salamander
Desmognathus monticola Seal Salamander
Desmognathus ochrophaeus Mountain Dusky Salamander
Eurycea cirrigera Southern Two-lined Salamander
Eurycea longicauda Longtail Salamander
Eurycea lucifuga Cave Salamander
Gastrophryne carolinensis Eastern Narrowmouth Toad
Gyrinophilus porphyriticus Spring Salamander
Hemidactylium scutatum Four-toed Salamander
Hyla versicolor Gray Treefrog
Lithobates catesbeianus Bullfrog
Lithobates clamitans Green Frog
Lithobates palustris Pickerel Frog
Lithobates sphenocephalus Southern Leopard Frog
Necturus maculosus Mudpuppy
Notophthalmus viridiscens Red-spotted Newt
Plethodon dorsalis Zigzag Salamander
Plethodon glutinosus Slimy Salamander
Plethodon serratus Southern Redback Salamander
Pseudacris crucifer Spring Peeper
Pseudacris triseriata Upland Chorus Frog
Pseudotriton montanus Mud Salamander
Pseudotriton ruber Red Salamander
Scaphiopus holbrookii Eastern Spadefoot Toad

9
 Table 1.2

Reptile species likely to occur in Hamilton County, Tennessee, compiled from

Conant and Collins (1998).

Species Name Common Name

Agkistrodon contortrix Copperhead

Anolis carolinensis Green Anole
Apalone spinifera Softshell Turtle
Aspidoscelis sexlineatus Six-lined Racerunner
Carphophis amoenus Worm Snake
Cemophora coccinea Scarlet Snake
Chelydra serpentina Snapping Turtle
Chrysemys picta Painted Turtle
Coluber constrictor Black Racer
Crotalus horridus Timber Rattlesnake
Diadophis punctatus Ringneck Snake
Graptemys geographica Common Map Turtle
Graptemys pseudogeographica False Map Turtle
Heterodon platirhinos Hognose Snake
Kinosternon subrubrum Eastern Mud Turtle
Lampropeltis calligaster Mole Kingsnake
Lampropeltis getula Kingsnake
Lampropeltis triangulum Milksnake
Nerodia sipedon Midland Water Snake
Opheodrys aestivus Rough Green Snake
Ophisaurus attenuatus Eastern Glass Lizard
Pantherophis obsoletus Rat Snake
Pantherophis guttatus Corn Snake
Pituophis melanoleucus Pine Snake
Plestiodon fasciatus Five-lined Skink
Plestiodon inexpectatus Southeastern Five-lined Skink
Plestiodon laticeps Broadhead Skink
Pseudemys concinna Eastern River Cooter
Regina septemvittata Queen Snake
Sceloporus undulatus Eastern Fence Lizard
Scincella lateralis Ground Skink
Sternotherus minor Stripeneck Musk Turtle
Sternotherus odoratus Common Musk Turtle
Storeria dekayi Brown Snake
Storeria occipitomaculata Redbelly Snake
Tantilla coronata Southeastern Crowned Snake
Terrapene carolina Eastern Box Turtle
Thamnophis sirtalis Garter Snake
Trachemys scripta Slider Turtle
Virginia valeriae Smooth Earth Snake

10
 Figure 1.1

A gravid female Ambystoma maculatum (top plate) exhibiting increased girth due to the presence
of eggs, and a male A. maculatum (bottom plate) exhibiting swollen cloacal lips (indicated by
arrow).

11
 Figure 1.2

A gravid female Ambystoma opacum (top plate) exhibiting darker crossbands and increased girth
due to the presence of eggs, and a male A. opacum (bottom plate) exhibiting the characteristic
bright white crossbands and swollen cloacal lips (indicated by arrow).

12
 Figure 1.3

Study area map showing LT6 (delineated by white lines) in Hamilton County, Tennessee.

13
 CHAPTER II

RELATIVE ABUNDANCE AND SPECIES DIVERSITY AT LT6

Abstract

I utilized a drift fence and pitfall array to sample amphibians and reptiles in LT6 from 15

September 2007 to 1 May 2010. A total of 30 herpetofaunal species was recorded during the

overall study period (H’=2.190; Smith and Wilson’s Evar=0.108), which comprise 45.16% of

amphibian species and 35.0% of reptile species potentially occurring in Hamilton County.

Thirteen anuran species, five lizard species, three salamander species, four snake species, and

five turtle species were recorded. Salamanders, especially Ambystoma maculatum and A.

opacum, were the most numerically abundant species (2,108 and 2, 111 individuals recorded,

respectively). Snakes were the least numerically dominant taxa. Species richness for the first

study year was 18 (H’=1.325; Evar=0.130). Richness for the second and third study year was 24

(H’=2.420; Evar=0.122) and 13 (H’=1.721; Evar=0.133), respectively. Richness was positively

correlated with time, but no new species were recorded after 23 months of sampling. Jaccard’s

index values showed moderate similarity between each study year as values ranged from 0.400

to 0.542. Previous studies utilizing pitfalls to determine abundance and richness at other

locations have had comparable results, and this study recorded more species than a previous

survey at LT6 that did not utilize pitfall or drift fence arrays.

14
Introduction

Few studies have adequately quantified the abundance of amphibians from isolated

wetlands over a well-defined time frame (Gibbons et al. 2006). This is especially true for the

southeastern USA, where descriptive ecological research on herpetofauna is notably almost

absent, with a few exceptions for species presence/absence studies [Miller et al. (1995), Miller

(1996), Neimiller et al. (2011)]. Although Miller (1996) conducted a herpetofaunal survey of the

former VAAP property (Hamilton County, Tennessee), the author highlighted that this study was

not meant to be a thorough investigation (B. Miller pers. comm.). Furthermore, none of these

previous studies has focused primarily on wetland habitats. Miller et al.’s (1995) investigation

took place in south-central Tennessee, approximately 75 miles northwest of Chattanooga.

Similarly, Neimiller et al. (2011) investigated areas in mid-central Tennessee. Hence, a

thorough, local study of species richness and abundance is non-existent in Hamilton County.

Approximately 31 species of amphibians and 40 species of reptiles are likely to occur in

Hamilton County based on distribution maps presented in Conant and Collins (1998); of these,

Redmond and Scott (1996, 2008) have verified 25 amphibian species and 30 reptile species in

Hamilton County. Although many of these species utilize wetlands for breeding, it is reasonable

to assume that few utilize wetlands exclusively because most species lack an entirely aquatic

adult form. This lends great urgency to conservation of entire upland areas, as it is well known

that the diversity of species in areas where these habitats are lost can be seriously impaired

(Semlitsch and Bodie 1998). Salamanders are known to migrate over 100 m into uplands

(Kleeberger and Werner 1983), but some species (e.g., Lithobates catesbeianus and

Notophthalmus viridiscens) may migrate up to 1600 m away from the aquatic habitat (Semlitsch

15
and Bodie 2003). An upland buffer of 275 m beyond the wetland boundary has been

recommended for turtles (Burke and Gibbons 1995).

Rapid assessments, similar to this study, are targeted biodiversity surveys based on an

urgency to determine richness and abundance, and serve as an important prelude to management

plans (Vonesh et al. 2010). Indeed, abundance is a good measure of productivity, and amphibian

species serve a larger ecological role as an important indicator of human impacts on wetland

communities (Gibbons et al. 2006). Also, biodiversity on a regional and national scale is often

measured in terms of species richness and gives an indication of an area’s overall conservation

value (Bock et al. 2007, Gibbons et al. 2006, Hecnar and M’Closkey 1996). Thus, assessments

reporting population estimates, relative abundance data, and presence/absence measurements are

vital because they can often be readily incorporated into adaptive conservation and management

strategies such as road closures during nights of mass amphibian movements (Timm et al. 2007).

Moreover, relatively short investigations can give vital information regarding local biodiversity.

Graham et al. (2010) reported a total of 62 species and 36 new county records after only seven

days of “bioblitz” sampling in southern Georgia. Indeed, site-specific analyses are critical to

producing an overall picture of species status across the range because information on large-scale

changes in population dynamics for most species is almost non-existent (Hecnar and M’Closkey

1996). However, long-term studies are still needed to monitor community fluctuations

(Blaustein et al. 1994). Unfortunately, these studies reporting population statuses are often left

to government agencies that lack sampling intensity, duration, and critical peer-review (Gibbons

et al. 1997).

Thus, my research objective was to determine and evaluate species richness and

16
relative abundance of amphibians and reptiles in LT6 for comparison to those reported in the

literature for similar habitat type and across years.

Materials and Methods

The drift fence utilized in this study was aluminum flashing with a height of 46 cm.

Aluminum flashing was chosen due to its overall durability, as opposed to screenwire used by

Shoop (1965), or mesh hardware cloth used by Sexton et al (1990). Construction of the fence

began late summer of 2007 by burying the flashing 10 to 15 cm deep, and then using exterior-

grade screws to attach wooden posts approximately every 10 meters for stabilization. Pitfall

traps consisted of large, deep 18.9 liter (5 gallon) plastic buckets, which were countersunk in

pairs along the fence every 6.5 m (on average) if the soil was not too compacted or rocky (Figure

2.1-2.3). There were 167 pitfalls and a total fence length of 747 m in use by December 2009.

Sticks, leaf litter, small rocks, moss, and various other natural substrates were placed into pitfalls

to create suitable microhabitat for captured animals and reduce the risk of desiccation.

Moistened sponges were also placed in each pitfall during summer months or when drought

conditions persisted. Bucket lids were suspended several centimeters over the pitfalls using

bamboo skewers to further reduce direct sunlight exposure and protect captured animals from

predation. Each pitfall was checked at least once daily to ensure the survival of captured animals

(Halliday 2006). Species observed outside of pitfalls, but near the fence (approximately 5 m),

were captured if possible and also counted as incidental captures. When not in use, tightly-

fitting pitfall lids were attached at ground level and covered with soil to ensure full closure.

Closure of the fence was very rare, primarily occurring only on weekends during the first few

17
study months, when less than 100 meters of fencing was in place. Maintenance of the fence was

completed as-needed throughout the study.

As detailed in Chapter 1, organisms were uniquely marked and morphological

measurements were obtained. Species richness was calculated for each study year and the study

duration based on first-time captures. However, if a single recaptured individual was the only

individual of that species recorded during a given year, this was included in richness

calculations. Total species richness was compared with species lists compiled from previous

investigations at LT6 (Wilson et al., unpubl. data, obtained from 2004-2007) and the former

VAAP property (Miller 1996), and published field guides (Conant and Collins 1998; Redmond

and Scott 1996, 2008). Jaccard’s index and percent similarity were calculated between each

study year and the cumulative study to indicate the degree of similarity (Krebs 1999). The

Shannon-Weiner index (H’) and Simpson’s index (1-D) were used to determine community

heterogeneity, and Smith and Wilson’s Index of Evenness (Evar) was used to estimate equitability

(Krebs 1999). The Shannon-Weiner index was also expressed in another form (N1) to show the

number of equally common species and was utilized because both richness and abundance are

represented (Krebs 1999). All results were calculated using Ecological Methodology version 5.2

(Krebs 1999).

Results

A total of 30 amphibian and reptile species were captured during the overall study period,

representing over 5,400 individual captures (Tables 2.1-2.4). Species richness for the first,

second, and third seasons was 18, 24, and 13, respectively (Tables 2.5-2.6). Total species

richness exceeded similar studies using a species area curve for comparison (Figure 2.4).

18
Salamanders (especially Ambystoma maculatum and A. opacum) had the highest rate of capture

of all species in each study year, and snakes had the lowest rate of capture in each study year

(Figure 2.5). Species observed in all study years include Ambystoma maculatum, A. opacum,

Anaxyrus woodhousii, Coluber constrictor, Lithobates catesbeianus, L. clamitans, L.

sphenocephalus, Notophthalmus viridiscens, Pseudacris triseriata, and Terrapene carolina.

Species observed only during the first study year include Agkistrodon contortrix, Anaxyrus

americanus, Diadophis punctatus, Hyla cinerea, Scaphiopus holbrookii, and Trachemys scripta

(Figure 2.6; Table 2.3-2.4). Species observed only during the second study year include

Aspidoscelis sexlineatus, Chelydra serpentina, Hyla versicolor, Opheodrys aestivus, Plestiodon

fasciatus, P. inexpectatus, P. laticeps, Pseudacris crucifer, and Sceloporus undulatus (Figure

2.7; Table 2.3-2.4). All species encountered during the third study year had been previously

recorded (Figure 2.8; Table 2.3-2.4). Maximum species richness was recorded after 23 months

of sampling (Figure 2.9).

Overall, 45.16% of amphibian species that occur in Hamilton County based on range

maps in Conant and Collins (1998) were observed, and 35.0% of reptile species (Table 2.1, 2.2).

Importantly, two species that were recorded in this study did not match range maps in Conant

and Collins (1998). These were Acris gryllus (southern cricket frog), a new county record for

Hamilton County (Simpson and Wilson 2009), and Hyla cinerea (green treefrog), a previously

reported Hamilton County record (Miller et al. 2007). Fifty-two percent of amphibian species

and 36.67% of reptile species confirmed in Hamilton County by Redmond and Scott (2008) were

recorded (Table 2.1). A maximum of 76.47% of amphibian species and 55.0% of reptile species

confirmed at the former VAAP property by Miller (1996) was recorded. All amphibian and

19
reptile species observed by Wilson et al. (unpubl. data) were recorded except Ambystoma

tigrinum, Thamnophis sirtalis, and Crotalus horridus (Table 2.1-2.2).

Shannon-Weiner H’ values ranged from 1.325 (Year 1: Sept. 2007 - Sept. 2008) to a

maximum of 2.420 (Year 2: Oct. 2008 - Sept. 2009). The number of equally common species

(N1) ranged from 2.51 during the first year to a maximum of 5.35 in the second study year (Table

2.6). Simpson’s index values ranged from a minimum 0.497 (first year) to a maximum of 0.727

(second year). Evenness in each study year was low, with Evar ranging from 0.122 (second year)

to 0.133 (third year). The overall study (Sept. 2007- May 2010) had an H’ value of 2.190, a

Simpson’s index value of 0.693, and Evar of 0.108 (Table 2.6). Similarity coefficients among

years ranged from a minimum of 51.42% between the first and second years to a maximum of

84.98% between the first and third years (Table 2.7). Jaccard’s index values ranged from a

minimum 0.400 (between years 1 and 2) to a maximum of 0.542 (between years 2 and 3; Table

2.7).

Discussion

This study is comparable to much of previous research using pitfall arrays as the primary

method in determining abundance and richness (see Figure 2.4). Greenberg et al. (1994)

reported capturing 11 species in pitfalls and 15 species in double-ended funnel traps in sand pine

scrub habitat of north-central Florida. Paton et al. (2000) reported capturing 10 species in

pitfalls in southern Rhode Island. A study by Miller et al. (1995) sampled the 15,800 ha property

at Arnold Air Force Base in Tennessee for a total of 64 days and reported the presence of 60

species. Although Miller et al. (1995) documented twice as many species as this study,

numerous survey methods not employed in the present study were utilized including auditory

20
surveys, seine netting, dip-netting, road cruising, and baited minnow traps. Similarly, Neimiller

et al. (2011) investigated multiple sites ranging in size from 61 to 4,130 hectares using multiple

techniques not utilized in the present study and determined richness values between 23 and 41,

respectively. Crosswhite et al. (1999) captured 38 species of reptiles and amphibians during a 91

day study period in upland hardwood habitat using pitfalls and double-ended funnel traps,

whereas Gibbons et al. (2006) captured a total of 24 species in a South Carolina wetland using

pitfalls and funnel traps in a single year. Indeed, my total richness value (30) exceeds both

Greenberg et al. (1994), Gibbons et al. (2006), and some sites in Neimiller et al. (2011),

illustrating that pitfall arrays utilized over many months may be just as effective for

presence/absence inventories as combined sampling methods because the likelihood of capture is

greatly increased.

The survey by Miller (1996) gives insight into previous herpetofaunal species richness

and abundance that occurred on property adjacent to LT6, although much of Miller’s former

study site has likely been developed since 2008. Miller (1996) reported the presence of 17

amphibian species at the former VAAP site; all species were recorded in the present study except

Desmognathus monticola, Eurycea cirrigera, Plethodon glutinosus, and Pseudotriton ruber. A

total of 20 reptile species was recorded by Miller (1996), but several species were not recorded

in the present study (e.g., Carphophis amoenus, Crotalus horridus, Lampropeltis calligaster,

Nerodia sipedon, Pantherophis guttatus, Scincella lateralis, Storeria dekayi, Tantilla coronata,

and Thamnophis sirtalis). However, 3 herpetofaunal species were recorded in the present study

that were not recorded by Miller (1996), including Kinosternon subrubrum, Plestiodon

inexpectatus, and Plestiodon laticeps.

21
 The present study recorded 13 of the 25 amphibian species and 11 of the 30 reptile

species verified in Hamilton County by Redmond and Scott (1996, 2008). As previously

mentioned, two amphibian species represented updated range distributions (Acris gryllus and

Hyla cinerea), and Pseudacris crucifer was a new record for Hamilton County (Simpson and

Wilson 2009). Additionally, several reptile species observed in the present study were not

previously verified by Redmond and Scott (2008), including Chelydra serpentina, Plestiodon

inexpectatus, and Plestiodon laticeps), indicating that more herpetofaunal research in Hamilton

County is needed. Futhermore, additional reptile species not observed by Wilson et al. (unpubl.

data) were recorded in the present study, which included the turtle species Chelydra serpentina,

Chrysemys picta, and Terrapene carolina, and the squamate species Aspidoscelis sexlineatus,

Diadophis punctatus, Opheodrys aestivus, Plestiodon inexpectatus, and Plestiodon laticeps.

Wilson et al. (2005) observed Ambystoma tigrinum (the tiger salamander) at LT6, but A.

tigrinum was not recorded in the present study. Importantly, only a single deceased A. tigrinum

individual was recovered on the perimeter of the wetland by Wilson et al. (unpubl. data). A.

tigrinum also exhibits large breeding movements and is easily sampled using pitfalls (Semlitsch

1983), and it has been reported that A. tigrinum may disperse distances up to 250 meters away

from breeding sites (Semlitsch 1983). At least eight other wetlands are known to occur within

one kilometer of LT6 and were likely the source of the single incidental capture. Tiger

salamanders also prefer deeper waters than spotted salamanders (Conant and Collins 1998), and

LT6 dries completely during the summer months. Further, tiger salamanders are locally rare (D.

Collins, pers. comm.), and thus it is unlikely that any individuals utilized LT6 during the present

study.

22
 It should be noted that most herpetofaunal species reported by Conant and Collins (1998),

Miller (1996), Redmond and Scott (1996, 2008), or Wilson et al. (unpubl. data) that were not

observed in the present study are not likely to be captured in pitfalls surrounding wetlands

because they do not typically utilize wetlands for breeding (Conant and Collins 1998, Gibbons

and Semlitsch 1981). Specific amphibian genera include Aneides, Eurycea, Desmognathus,

Gyrinophilus, Hemidactylium, Necturus, Plethodon, and Pseudotriton species. Aneides prefer

mesic hardwood forests or rocky outcrops, while Desmognathus, Eurycea, Gyrinophilus, and

Pseudotriton primarily utilize streams, springs, and seeps for habitat (Bailey et al. 2006).

Hemidactylium prefer wet meadows and bogs, Necturus prefer rivers or permanent wetlands, and

Plethodon prefer mesic hardwood uplands (Bailey et al. 2006). Eurycea and Plethodon species

were captured in pitfalls by Gibbons and Semlitsch (1981), but at very low abundances in

comparison with other taxa. Thus, only incidental captures should be expected, and none were

observed in the present study.

Similarly, most reptile genera ascribed by the aforementioned sources to Hamilton

County but not captured in this study (e.g., Anolis, Apalone, Carphophis, Cemophora, Crotalus,

Graptemys, Heterodon, Lampropeltis, Ophisaurus, Pantherophis, Pseudemys, Regina, Scincella,

Tantilla, and Virginia species) are not heavily associated with wetland habitats (Bailey et al.

2006). However, six reptile species not observed in the present study are associated with

wetland habitats, including Lampropeltis getula, Nerodia sipedon, Sternotherus minor,

Sternotherus odoratus, Storeria dekayi, and Tantilla coronata (Bailey et al. 2006). The latter six

species are likely to occur at LT6, but may have gone unobserved because snake and turtle

species are not easily sampled using pitfalls (Dodd 1991, Gibbons and Semlitsch 1981). Further,

all snakes except Diadophis punctatus were observed outside of pitfalls, along the drift fence, in

23
this study. Thus, habitat use and ease of escape greatly inhibits adequate sampling of snakes by

the pitfall array at LT6.

Implications for Conservation

LT6 is undoubtedly a speciose area, but relative abundance must be interpreted with

some caution. As expected, species that are most easily sampled using pitfalls always ranked

highest on abundance plots. Notophthalmus viridiscens and Pseudacris crucifer were observed

primarily in the second study year, indicating that these species likely exhibit a biennial mating

cycle, although a longer study duration would be needed for verification. All reptile species had

the largest capture numbers during the summer months of the first and second study years.

However, reptile captures were low (less than 10 total captures for all species), and no clear

temporal patterns were identified. However, although Terrapene carolina was the most

commonly recorded reptile, numerous Coluber constrictor individuals were observed outside of

pitfalls at LT6. Thus, it is likely that this species is much more common than pitfall data

suggests. Anurans, especially Lithobates species, also had the highest capture numbers during

the summer months. All Gastrophryne carolinensis individuals were captured from July through

September, and this is likely the reason that no Gastrophryne individuals were observed during

the third study year, because sampling was halted in May. Interestingly, taxa with peak

movements during the fall and spring months also had relatively low capture numbers in the

third year compared to the previous two study years. Thus, variations in abundance were likely

due to natural fluctuations and were not effort-related, except for Gastrophryne.

It is known that some herpetofaunal species are capable of crossing aluminum fence

arrays, such as Hyla and many snake species (Dodd 1991). Mole and invertebrate tunnels may

24
be a source of escape for some genera, including Acris, Gastrophryne, Hyla, and Notopthalmus

(Dodd 1991). However, no tunnels were visually observed at LT6. Soil at the study site had an

abundance of chert and clay that was heavily compacted during installation, further decreasing

the possibility of tunnel passages. Additionally, the use of large, deep pitfalls (such as those

used in this study) most likely increased the time in which hylids were present in the pitfall by

acting as refugia (Willson and Gibbons 2009). Also, while intermittent gaps present in the drift

fence during construction and maintenance may slightly alter abundance measurements, gaps are

not likely to reduce species richness because wetland size is not a strong correlate of species

richness. Snodgrass et al. (2000) reported that reducing wetland size from 78 hectares to 0.2

hectares would reduce the average species richness from 8.3 to 7.7, a change of less than one

species. However, it is well known that upland habitat is critical to maintaining biodiversity and

that wetland size alone is not capable of predicting species richness (Gibbons 2003, Gibbs 1998).

Thus, the upland buffer surrounding LT6 is undoubtedly a critical source for wetland

biodiversity observed in this study and confirms separate investigations highlighting the

importance of forested wetland buffers (Gibbons 2003).

This study highlights the importance of local conservation and survey efforts that

document the accuracy of published species distributions, which are often constructed with

limited information (Gibbons et al. 1997). First, the expanded ranges of Hyla cinerea (Miller et

al. 2007) and Acris gryllus (Simpson and Wilson 2009), uncovered in association with this study,

are evidence of the usefulness of local surveys and the need for more such surveys. Indeed, the

southern cricket frog (Acris gryllus) is listed in the Guide to the Rare Animals of Tennessee

(Withers 2009) as being found in “far southwest Tennessee”, and given the S2S3 conservation

rank, indicating this species is rare, uncommon, or possibly imperiled within the state. However,

25
this study indicates that A. gryllus may be more common than previously known and more

research regarding its distribution is needed. Similarly, richness data recorded here also

highlights the need for updates to the State Atlas of Amphibians and Reptiles (Redmond and

Scott 1996, 2008) to give more accurate descriptions of distribution patterns.

26
 Table 2.1

Amphibian species likely to occur in Hamilton County, Tennessee, compiled from Conant and
Collins (1998). Species confirmed in Miller (1996), Redmond and Scott (1996), Wilson unpubl.
data, and/or this study are noted.

Species Name Common Name Miller Redmond Wilson This

1996 and Scott unpubl. study
1996 data

Acris crepitans Northern Cricket Frog Yes Yes - Yes

Ambystoma maculatum Spotted Salamander Yes Yes Yes Yes
Ambystoma opacum Marbled Salamander Yes Yes Yes Yes
Ambystoma tigrinum Tiger Salamander - Yes Yes -
Anaxyrus americanus American Toad Yes Yes Yes Yes
Anaxyrus woodhousii Fowler’s Toad Yes Yes - Yes
Aneides aeneus Green Salamander - Yes - -
Desmognathus fuscus Dusky Salamander - Yes - -
Desmognathus monticola Seal Salamander Yes Yes - -
Desmognathus ochrophaeus Mountain Dusky Salamander - - - -
Eurycea cirrigera Two-lined Salamander Yes Yes - -
Eurycea longicauda Longtail Salamander - Yes - -
Eurycea lucifuga Cave Salamander - Yes - -
Gastrophryne carolinensis Eastern Narrowmouth Toad Yes Yes Yes Yes
Gyrinophilus porphyriticus Spring Salamander - Yes - -
Hemidactylium scutatum Four-toed Salamander - Yes - -
Hyla versicolor Gray Treefrog Yes Yes Yes Yes
Lithobates catesbeianus Bullfrog Yes Yes Yes Yes
Lithobates clamitans Green Frog Yes Yes Yes Yes
Lithobates palustris Pickerel Frog - - - -
Lithobates sphenocephalus Southern Leopard Frog Yes Yes Yes Yes
Necturus maculosus Mudpuppy - - - -
Notophthalmus viridiscens Red-spotted Newt Yes Yes Yes Yes
Plethodon dorsalis Zigzag Salamander - Yes - -
Plethodon glutinosus Slimy Salamander Yes Yes Yes -
Plethodon serratus Southern Redback Salamander - - - -
Pseudacris crucifer Spring Peeper Yes Yes* Yes Yes*
Pseudacris triseriata Upland Chorus Frog Yes Yes Yes Yes
Pseudotriton montanus Mud Salamander - - - -
Pseudotriton ruber Red Salamander Yes Yes - -
Scaphiopus holbrookii Eastern Spadefoot Toad - Yes - Yes

* Indicates a new county record for Hamilton Co., Tennessee obtained during this study.
Note: Acris gryllus, also a new county and geographic distribution record, is not included here because the
species does not match distribution maps in Conant and Collins (1998), from which this table was compiled.

27
 Table 2.2

Reptile species likely to occur in Hamilton County, Tennessee, compiled from Conant and
Collins (1998). Species confirmed in Miller (1996), Redmond and Scott (2008), Wilson unpubl.
data, and/or this study are noted.

Species Name Common Name Miller Redmond Wilson This

1996 and Scott unpubl. study
2008 data

Agkistrodon contortrix Copperhead Yes Yes Yes Yes

Anolis carolinensis Green Anole - Yes - -
Apalone spinifera Softshell Turtle - Yes - -
Aspidoscelis sexlineatus Six-lined Racerunner Yes Yes - Yes
Carphophis amoenus Worm Snake Yes Yes - -
Cemophora coccinea Scarlet Snake - - - -
Chelydra serpentina Snapping Turtle Yes - - Yes
Chrysemys picta Painted Turtle Yes Yes - Yes
Coluber constrictor Black Racer Yes Yes Yes Yes
Crotalus horridus Timber Rattlesnake Yes Yes Yes -
Diadophis punctatus Ringneck Snake Yes Yes - Yes
Graptemys geographica Common Map Turtle - - - -
Graptemys pseudogeographica False Map Turtle - - - -
Heterodon platirhinos Eastern Hognose Snake - - - -
Kinosternon subrubrum Eastern Mud Turtle - Yes - Yes
Lampropeltis calligaster Mole Kingsnake Yes Yes - -
Lampropeltis getula Black Kingsnake - Yes - -
Lampropeltis triangulum Milk Snake - Yes - -
Nerodia sipedon Midland Water Snake Yes Yes - -
Opheodrys aestivus Rough Green Snake Yes Yes - Yes
Ophisaurus attenuates Slender Glass Lizard - Yes - -
Pantherophis guttatus Corn Snake Yes Yes - -
Pantherophis obsoleta Rat Snake - Yes - -
Pituophis melanoleucus Pine Snake - - - -
Plestiodon fasciatus Five-lined Skink Yes Yes - Yes
Plestiodon inexpectatus SE Five-lined Skink - - Yes Yes
Plestiodon laticeps Broadhead Skink - - - Yes
Pseudemys concinna River Cooter - Yes - -
Regina septemvittata Queen Snake - - - -
Sceloporus undulatus Fence Lizard Yes Yes Yes Yes
Scincella lateralis Ground Skink Yes Yes - -
Sternotherus minor Stripeneck Musk Turtle - Yes - -
Sternotherus odoratus Common Musk Turtle - Yes - -
Storeria dekayi Brown Snake Yes Yes - -
Storeria occipitomaculata Redbelly Snake - - - -
Tantilla coronata SE Crowned Snake Yes Yes - -
Terrapene carolina Eastern Box Turtle Yes Yes Yes Yes
Thamnophis sirtalis Garter Snake Yes Yes Yes -
Trachemys scripta Slider Turtles Yes Yes - Yes
Virginia valeriae Smooth Earth Snake - Yes - -

28
 Table 2.3

Total amphibian captures (recaptures excluded) by study year and the entire study period (Sept.
2007- May 2010).

Taxon Sept 2007 – Oct 2008 – Oct 2009 – Total

Sept 2008 Sept 2009 May 2010 N

CAUDATA
Ambystoma maculatum 1,172 464 472 2,108
Ambystoma opacum 545 1,251 315 2,111
Notophthalmus viridiscens 14 445 1 460

ANURA
Acris crepitans - 4 3 7
* *
Acris gryllus* - 9 2 11*
Anaxyrus americanus 1 - - 1
Anaxyrus woodhousii 1 23 1 25
Gastrophryne carolinensis 5 5 - 10
Hyla cinerea 1 - - 1
Hyla versicolor - 1 - 1
Lithobates catesbeianus 23 89 43 155
Lithobates clamitans 2 37 55 94
Lithobates sphenocephalus 11 207 9 227
Pseudacris crucifer* - 138 - 138
Pseudacris triseriata 41 40 12 93
Scaphiopus holbrookii 1 - - 1
* Indicates a new county record for Hamilton Co., Tennessee, obtained during this study.

29
 Table 2.4

Total reptile captures (recaptures excluded) by study year and the entire study period (Sept.
2007-May 2010).

Taxon Sept 2007 – Oct 2008 – Oct 2009 – Total N

Sept 2008 Sept 2009 May 2010

TESTUDINES

Chelydra serpentina - 3 - 3

Chrysemys picta 1 1 - 2

Kinosternon subrubrum - 1 1 2

Terrapene carolina 1 4 1 6

Trachemys scripta 1 - - 1

SQUAMATA

Agkistrodon contortrix 1 - - 1

Aspidoscelis sexlineatus - 1 - 1

Coluber constrictor 1 2 1 4

Diadophis punctatus 1 - - 1

Opheodrys aestivus - 1 - 1

Plestiodon fasciatus - 9 - 9

Plestiodon inexpectatus - 1 - 1

Plestiodon laticeps - 2 - 2

Sceloporus undulatus - 1 - 1

30
 Table 2.5

Species richness comparisons from previous studies and this study (Sept. 2007-May 2010).

Reference Habitat type Study area size (ha) Methods Total

Species
Recorded

Gibbons et al. 2006 Open scrub ~10 Pitfall array, funnel traps 24

Greenberg et al. Wetland ~8.5 Pitfall array, funnel traps 20

1994

Miller 1996 Mixed deciduous upland, wetland ~6,587 Auditory surveys, dipnetting, hoop nets, manual searching, 37
minnow traps, seine netting, road cruising, visual surveys

Miller et al. 1995 Mixed deciduous upland, ~15,783 Auditory surveys, coverboards, dipnetting, manual searching, 60
wetland, disturbed area, riparian minnow traps, pitfall array, road cruising, seine netting, visual
zone surveys

Neimiller et al. 2011 Cedar/cedar-hardwood forest, ~288 Auditory surveys, coverboards, dipnetting, funnel box array, hoop 29
(multiple sites) cedar glades and barren, nets, manual searching, minnow traps, road cruising, visual surveys
ephemeral pond, stream

~342 Auditory surveys, dipnetting, minnow traps, visual surveys 23

~250 Auditory surveys, road cruising, visual surveys 24

~4,130 Auditory surveys, dipnetting, minnow traps, road cruising, visual 41

surveys

~61 Auditory/visual surveys 24

Paton et al. 2000 Mixed deciduous upland, wetland ~0.5 Pitfall array 10

This study Mixed deciduous upland, wetland ~2 Pitfall array, incidental captures near array 30

31
 Table 2.6

Species richness and heterogeneity measures of LT6 for the entire study and by study year (Year
1 = Sept. 2007 - Sept. 2008, Year 2 = Oct. 2008 - Sept. 2009, and Year 3 = Oct. 2009 - May
2010).

Study Year Species Shannon-Weiner Shannon-Weiner Simpson’s Smith and

Richness Index of Number of Index of Wilson’s Index
Diversity (H’) Equally Common Diversity of Evenness
Species (N1) (1-D) (Evar)

Total 30 2.190 4.56 0.693 0.108

Year 1 18 1.325 2.51 0.497 0.130

Year 2 23 2.420 5.35 0.727 0.122

Year 3 13 1.721 3.30 0.611 0.133

32
 Table 2.7

Jaccard’s index values and percent similarity values between study years (Year 1 = Sept. 2007 -
Sept. 2008, Year 2 = Oct. 2008 - Sept. 2009, and Year 3 = Oct. 2009 - May 2010).

Study Years Jaccard’s Index Percent Similarity

Total – Year 1 0.600 73.31%

Total – Year 2 0.800 77.75%

Total – Year 3 0.433 80.47%

Year 1 – Year 2 0.400 51.42%

Year 1 – Year 3 0.476 84.98%

Year 2 – Year 3 0.542 59.02%

33
 ~6.5 m

Figure 2.1

General design of the drift fence (horizontal line) and pitfall traps (circles), with approximately
6.5 m spacing from center to center of adjacent pitfalls.

34
 Figure 2.2

A general view of the installed drift fence. A measuring tape (~60 cm in length) is
attached for scale.

35
 Figure 2.3

Vertical view of an installed 18.9 liter pitfall (with lid attached) adjacent to flashing. A
measuring tape (~20 cm in length) is attached for scale.

36
 Species Richness v. log10 Area + 1
70
60 (C)
60
Species Richness
50
41 (D)
40
30 (F) 29 (D)
30 24 (A)
24 (D)
20 20 (B) 24 (D) 23 (D)
10 10 (E)

0
0 1 2 3 4 5 6
log10 Area + 1

Figure 2.4

Species area curve compiled using previous studies and this study. Studies represented on the
graph include Gibbons et al. 2006 (A), Greenberg et al. 1994 (B), Miller et al. 1995 (C), multiple
sites from Neimiller et al. 2011 (D), Paton et al. 2000 (E), and this study (F).

37
 Species Abundance (log10) v. Rank
5

Species Abundance (log10)

4
A.o.
A.m.
3 N.v.
L.s. P.c.
2 L.cates.
L.clam. P.t.
A.crep.
A.w. G.c.
1 A.g. T.c.
C.c. C.p., K.s., P.l.
P.f.
C.s.
0
0 10 A.cont., A.a., A.s.,D.p.,H.c.,H.v., 20
O.a., P.i., S.h., S.u., T.s.
Rank

Figure 2.5

Ranked abundances (log10) for all species recorded during the entire study period (Sept. 2007-
May 2010). Species represented on the graph include: Acris crepitans (A. crep.), Acris gryllus
(A.g.), Agkistrodon contortrix (A. cont.), Ambystoma maculatum (A.m.), Ambystoma opacum
(A.o.), Anaxyrus americanus (A.a.), Anaxyrus woodhousii (A.w.), Aspidoscelis sexlineatus (A.s.),
Chelydra serpentina (C.s.), Chrysemys picta (C.p.), Coluber constrictor (C.c.), Diadophis
punctatus (D.p.), Gastrophryne carolinensis (G.c.), Hyla cinera (H.c.), Hyla versicolor (H.v.),
Kinosternon subrubrum (K.s.), Lithobates catesbeianus (L. cates.), Lithobates clamitans (L.
clam.), Lithobates sphenocephalus (L.s.), Notophthalmus viridiscens (N.v.), Opheodrys aestivus
(O.a.), Plestiodon fasciatus (P.f.), Plestiodon inexpectatus (P.i.), Plestiodon laticeps (P.l.),
Pseudacris crucifer (P.c.), Pseudacris triseriata (P.t.), Scaphiopus holbrookii (S.h.), Sceloporus
undulatus (S.u.), Terrapene carolina (T.c.), and Trachemys scripta (T.s.).

38
 Species Abundance (log10) v. Rank
(Year 1)
Species Abundance (log10) 4
3.5
A.m.
3
A.o.
2.5
2 P.t. A.a., A.w., H.c., S.h.,
1.5 L. cates. C.p., T.c., T.s., A. cont.,
1 L.s. C.c., D.p.
N.v. G.c.
0.5
0 L. clam.
-0.5 0 5 10 15
-1
Rank

Figure 2.6

Ranked abundances (log10) for all species recorded during the first study year. Species
represented on the graph include: Agkistrodon contortrix (A. cont.), Ambystoma maculatum
(A.m.), Ambystoma opacum (A.o.), Anaxyrus americanus (A.a.), Anaxyrus woodhousii (A.w.),
Chrysemys picta (C.p.), Coluber constrictor (C.c.), Diadophis punctatus (D.p.), Gastrophryne
carolinensis (G.c.), Hyla cinerea (H.c.), Lithobates catesbeianus (L. cates.), Lithobates
clamitans (L. clam.), Lithobates sphenocephalus (L.s.), Notophthalmus viridiscens (N.v.),
Pseudacris triseriata (P.t.), Scaphiopus holbrookii (S.h.), Terrapene carolina (T.c.), and
Trachemys scripta (T.s.).

39
 Species Abundance (log10) v. Rank
(Year 2)
Species Abundance (log10) 4
3.5
3 A.o.
A.m. N.v.
2.5 P.c.
L.s.
2 L. cates. L. clam.
1.5 P.t. A.w. A. crep., T.c.
1 H.v., C.p., K.s.,
0.5 A.g., P.f.
G.c. C.c., P.l. A.s., P.i., O.a., S.u.
C.s.
0
0 5 10 15 20 25 30
Rank

Figure 2.7

Ranked abundances (log10) for all species recorded during the second study year. Species
represented on the graph include: Acris crepitans (A. crep.), Acris gryllus (A.g.), Ambystoma
maculatum (A.m.), Ambystoma opacum (A.o.), Anaxyrus woodhousii (A.w.), Aspidoscelis
sexlineatus (A.s.), Chelydra serpentina (C.s.), Chrysemys picta (C.p.), Coluber constrictor (C.c.),
Gastrophryne carolinensis (G.c.), Hyla versicolor (H.v.), Kinosternon subrubrum (K.s.),
Lithobates catesbeianus (L. cates.), Lithobates clamitans (L. clam.), Lithobates sphenocephalus
(L.s.), Notophthalmus viridiscens (N.v.), Opheodrys aestivus (O.a.), Plestiodon fasciatus (P.f.),
Plestiodon inexpectatus (P.i.), Plestiodon laticeps (P.l.), Pseudacris crucifer (P.c.), Pseudacris
triseriata (P.t.), Sceloporus undulatus (S.u.), and Terrapene carolina (T.c.).

40
 Species Abundance (log10) v. Rank
(Year 3)
3.5
Species Abundance (log10) 3
A.m.
2.5 A.o.

2 L. clam.
1.5 L. cates
P.t.
1 L.s.
A. crep.
0.5 A.g.N.v., A.w., K.s., T.c., C.c.
0
-0.5 0 2 4 6 8 10 12 14
Rank

Figure 2.8

Ranked abundances (log10) for all species recorded during the third study year. Species
represented on the graph include: Acris crepitans (A. crep.), Acris gryllus (A.g.),
Ambystoma maculatum (A.m.), Ambystoma opacum (A.o.), Anaxyrus woodhousii (A.w.),
Coluber constrictor (C.c.), Kinosternon subrubrum (K.s.), Lithobates catesbeianus (L.
cates.), Lithobates clamitans (L. clam.), Lithobates sphenocephalus (L.s.), Notophthalmus
viridiscens (N.v.), Pseudacris triseriata (P.t.), and Terrapene carolina (T.c.).

41
 Species Richness v. Time
30
25
Species Richness 20
15
10
5
0
Nov

Nov

Nov
Mar
May

Mar
May

Mar
May
Sept

Jul
Sept

Jul
Sept
Jan

Jan

Jan
2007 2008 2009 2010
Time (months)

Figure 2.9

A performance curve plotting time versus cumulative species richness as a measure of

effort.

42
 CHAPTER III

SEX RATIOS OF AMBYSTOMA AND BODY SIZES OF AMPHIBIANS AND

REPTILES PRESENT IN LT6

Abstract

Pitfall arrays were utilized to capture herpetofaunal species from September 2007 to May

2010 at LT6. Snout-vent length (mm), tail length (mm), body mass (g), gender, and age class

were recorded for all individuals. Species were ranked according to relative biomass and sex

ratios in ambystomatid salamanders were calculated for each study period. Biomass was

dominated by Ambystoma maculatum and A. opacum, together constituting ~87.5% of the total

biomass. The remaining 12.5% was comprised of eleven anuran species, four lizard species, one

salamander species, three snake species, and five turtle species. T-tests were used to test for

dimorphisms in A. maculatum and A. opacum. Females were significantly larger in both species

(p<0.05). Mean SVL for male spotted salamanders was 97.37±7.63 mm, and mean SVL for

female spotted salamanders was 103.56±8.39 mm. Mean SVL for male marbled salamanders

was 68.082±7.755 mm, and mean SVL for female marbled salamanders was 69.280±5.801 mm.

Sex ratios were significantly male biased in two out of three A. maculatum samples

(ratios=2.714*, 1.075, and 2.642*). Sex ratios were significantly female biased in one out of

three A. opacum samples (ratios=0.985, 1.026, and 0.590*). Sex ratio variability can attributed

to a variety of causes which include biennial mating cycles, differential maturation times, and

differential mortality between the sexes.

43
Introduction

Sex ratios are an important parameter in population biology (Viets 1994). Changes in

these ratios are often an indication of ecological or anthropogenic disturbance within the

community, although the extent of previous research varies by taxa. Amphibians exhibit

differing sex ratios among species because of temporal factors (e.g., differing maturation times

for males and females) (Berven 1981, Husting 1965, Sakisaka et al. 2000). Population data on

sex ratios in Ambystoma species have been documented extensively in the literature (see Table

3.1 for a complete list). Sex ratios of ambystomatid salamanders are easily determined because

individuals often move in a predictable manner and external gender differences among adults are

readily apparent. Sex ratios of A. maculatum are often shifted in favor of males (Husting 1965,

Peckham and Dineen 1954, Phillips and Sexton 1989, Sexton et al. 1986, and Whitford and

Vinegar 1966). Ratios in A. opacum are similar, with populations often being significantly male

biased (Krenz and Scott 1994, Stenhouse 1987). Male-biased sex ratios may arise from

differential maturation times, with females often needing an extra year to mature, and thus

suffering an extra year’s mortality (Wacasey 1961). Husting (1965) also suggested that females

may exhibit a biennial mating cycle, and this has been confirmed in the closely related species A.

talpoideum (Raymond and Hardy 1990). Sexton et al. (1986) proposed that misjudgement of

gender may be common after breeding (and the subsequent loss of secondary sex characteristics).

Thus, immediate sex identification when breeding animals first begin migration may be critical

in reporting accurate ratios.

Sexual dimorphism, which often manifests as coloration and size differences among

males and females of the same species, is also an important ecological factor that varies among

taxa. Amphibian species typically exhibit larger female size, likely due to increased fecundity

44
among larger females (Shine 1979). More specifically, size dimorphisms are well known in the

genus Ambystoma (Peckham and Dineen 1954, Raymond and Hardy 1990), although specific

records for A. maculatum and A. opacum do not abound in the literature. Measurements of size

dimorphisms are useful in determining age distribution, stability, and recruitment within

populations (Gibbons et al. 2006). Additionally, relative biomass can be estimated to index

productivity, which illustrates the importance of small wetlands in ecosystem dynamics as they

facilitate the transfer of energy from aquatic to terrestrial habitats (Gibbons et al. 2006). Many

factors can influence dimorphism (e.g., resource competition [Shine 1979]), and sampling bias

may also skew recorded dimorphisms (Gibbons and Lovich 1990).

Thus, my research objective was to determine mean body sizes, sex ratios, and sexual

size dimorphisms of Ambystoma maculatum and A. opacum in LT6 for comparison to those

reported in the literature and across years.

Materials and Methods

Amphibians and reptiles were captured from 15 September 2007 to 1 May 2010 using the

drift fence and pitfall arrays detailed in Chapter 2. Gender was determined if adult body size had

been attained and secondary sex characteristics were apparent (as described in Conant and

Collins 1998). Snout-vent length (hereafter, SVL) and tail length (TL) for amphibians and

reptiles were always determined to the nearest millimeter using a fish measuring board to

decrease handling time (Wildco©, Model #118-40) or dial calipers (Mitutoyo©, Model #505-

672). Body mass was determined to the nearest gram using a digital balance (My-Weigh©,

Model #7001DX).

45
 Male Ambystoma opacum were identified by the presence bright white crossbands and

swollen cloacal lips (Noble and Brady 1933). Individual A. opacum exhibiting gray crossbands,

swollen abdomens (indicating a gravid condition), and a lack of swollen cloacal lips were

regarded as females (Noble and Brady 1933). Similarly, male A. maculatum were identified by

swollen cloacal lips. Sex ratios were calculated for A. maculatum and A. opacum using data

collected from first-time migrations into the wetland during the first half of each breeding season

(based on capture numbers) in order to reduce gender misidentification. I considered the first

half of the A. opacum breeding season to be September through mid-November because the

majority of immigration (>50%) was completed before December. I considered the first half of

the A. maculatum breeding season to be January through the end of February, as the majority of

immigration (>50%) was completed before 1 March. Any early arrivals (before 1 January) of A.

maculatum were, however, considered in sex ratio calculations.

Data were tested for normality using the Shapiro-Wilk statistic and significance was set a

priori at a=0.05 for all significance tests. Analysis was performed using SAS software version

9.2 (2008). Chi-square analysis was used to determine any significant difference in sex ratios

during individual seasons (Raymond and Hardy 1990). Captured species were ranked according

to the relative percent biomass for first time captures. Rankings exclude Acris crepitans, A.

gryllus, and Pseudacris crucifer individuals because most weighed less than 1 gram and were

outside the limits of detection with the MyWeigh scale. Salamanders were divided into age class

(adult or juvenile) and the mean (± 1SD) SVL, tail length, and body mass were recorded. Sexual

size dimorphisms in Ambystoma were tested for significance using an independent samples t-test

between males and females for SVL, tail length, and body mass (Raymond and Hardy 1990).

Mean (± 1SD) reptile SVL, tail length, and body mass were also recorded. Anurans captured

46
were primarily juveniles and sample sizes were relatively small (less than 30 captures for the

majority of species), and thus median values (± IQR) were reported to reduce bias from large

differences in body sizes between adults and sub-adults.

Results

Sex ratios were significantly different during two years of Ambystoma maculatum

breeding movements and one breeding movement of A. opacum. The sex ratio of spotted

salamanders during 2008 was 2.714* (p<0.0001), 1.075 in 2009, and 2.642* (p<0.0001) in 2010

(Table 3.1). The sex ratio of marbled salamanders was 0.985 in 2007, 1.026 in 2008, and 0.590*

(p<0.05) in 2009 (Table 3.1).

Species biomass was dominated by the ambystomatids, with A. maculatum and A.

opacum comprising 61.14% and 26.35% of the total biomass, respectively (Table 3.2). The

remaining 12.5% was comprised of 24 species, and each species was ≤3% of the total biomass.

A. maculatum was the largest salamander recorded by body size, with males having a mean SVL

of 97.155±10.592 mm, a mean tail length of 92.37±9.035 mm, and a mean body mass of

18.428±3.484 grams (Table 3.3). Females were larger overall with a mean SVL of 103.6±8.494

mm, a mean tail length of 94.208±9.188 mm, and a mean body mass of 22.28±5.332 grams.

Measurements were significantly different between male and female SVL (p<0.0001), tail length

(p<0.0001), and body mass (p<0.0001). Male A. opacum had an average SVL of 68.088±7.755

mm, a mean tail length of 46.655±6.699 mm, and a mean body mass of 7.565±2.063 g. Females

averaged 69.280±5.801 mm SVL, 47.303±6.438 mm in tail length, and 8.987±2.344 grams in

body mass. Measurements were significantly different for SVL (p<0.005), tail length (p<0.05),

and body mass (p<0.0001). Juvenile A. opacum emigrating from the wetland after

47
metamorphosis had a mean SVL of 43.33±2.014 mm, mean tail length of 34.003±2.746 mm, and

a mean body mass of 2.177±0.557 grams. All Notophthalmus viridiscens individuals were

juveniles. These juvenile newts (efts) averaged 21.56±6.865 mm SVL, 20.85±7.987 mm in tail

length, and 1.083±0.327 grams in body mass (Table 3.3).

The majority of anuran, snake, and lizard species were encountered less than 10 times,

and thus statistical analysis of sexual dimorphism was not attempted. The largest median body

size among anurans captured was Lithobates catesbeianus (median SVL=44±40-48 mm, n=135;

median body mass=8±6-12 g, n=132) (Table 3.4). The smallest median body size among

captured anurans was Pseudacris crucifer (median SVL=13±0.429 mm, n=138) (Table 3.4).

Discussion

Sample sizes were largest in the two Ambystoma species (A. maculatum and A. opacum)

compared to other taxa during each study year. Drift fence and pitfall arrays are ideal for capture

pond-breeding salamanders, but have limitations in adequately sampling other herpetofaunal

species (Gibbons and Semlitsch 1981), and thus biomass may be somewhat under-represented

for non-salamander taxa. Specifically, turtles and snakes are not readily sampled using pitfalls,

according to Gibbons and Semlitsch (1981). Also, arboreal anuran species possessing adhesive

toe-pads may be able to escape pitfalls by climbing according to Dodd (1991). However, the

present study shows that drift fences may be more suitable to sampling non-traditional taxa than

previously expected. A total of five turtle species were recorded in this study, exceeding both

the total number of all other reptile species and the total number of salamander species. Further,

turtles recorded in the present study represent all turtle species occurring in Tennessee not

usually associated with a riverine habitat, except Sternotherus species. The large size of the drift

48
array coupled with an almost three-year study duration proved to be an effective combination for

increasing incidental captures. This was true even for species known to actively avoid pitfalls,

such as Terrapene carolina (Gibbons and Semlitsch 1981). Furthermore, adding natural

substrates to pitfalls that remain in place for many months may actually attract organisms by

serving as a stable, suitable habitat. Indeed, Lithobates sphenocephalus may seek refuge in

pitfalls and become over-sampled compared to other anuran taxa (Shields 1985). However, all

southern leopard frogs were individually marked in the present study and no indication of over

sampling (e.g., a high recapture rate) was observed.

Temporal variation in breeding among species is also critical when interpreting biomass

measurements (Gibbons and Semlitsch 1981). Sampling ended in May 2010 but was continuous

during the previous two summers. Thus, species most commonly encountered during the

summer months are under-represented during the final study year. These include Agkistrodon

contortrix, Anaxyrus sp., Chrysemys picta, Coluber constrictor, Plestiodon sp., and Trachemys

scripta. However, turtle, lizard, and snake species were primarily incidental captures and it is

difficult to predict how capture numbers were altered because of the low frequency of capture

among these taxa.

Numerous previous studies have reported unequal sex ratios in Ambystoma species.

Samples of A. maculatum from various locations were significantly male-biased in 14 of 18 total

samples (Peckham and Dineen 1954, Husting 1965, Whitford and Vinegar 1966, Sexton et al.

1986, Phillips and Sexton 1989). Similar results have also been reported in A. opacum, with two

of three samples being significantly biased in favor of males (Stenhouse 1987, Krenz and Scott

1994), although Parmelee (1993) reported a small sample of A. opacum (n=18) from Alexander

County, Illinois that was biased in favor of females (ratio=0.64). Husting (1965) was the first to

49
propose that these skewed ratios could be attributed to differential mortality, with females

suffering higher mortality rates. However, there is no proven evidence of increased female

mortality highlighted in the previous literature. A few additional causes for male-biased ratios

have been suggested since Husting (1965). Phillips and Sexton (1989) note that females may

need an extra year to mature or exhibit a biennial mating cycle. Indeed, biennial mating cycles

have contributed to male-biased ratios in Triturus newts (Arntzen 2002). It is also possible that

sampling bias contributes to unequal sex ratios in Ambystoma, with male-biased ratios being

caused by sampling only at the beginning of breeding movements (Whitford and Vinegar 1966).

However, sampling was continuous during the duration of breeding movements and it is thus

unlikely that females were undersampled in this manner in the present study. Long-term studies

are needed to determine if sex ratio fluctuations have an anthropogenic or natural cause, and

future comparisons may be able to correlate changes in average body sizes of Ambystoma to the

availability of resources within LT6.

Peckham and Dineen (1954) and Sexton et al. (1986) also reported significant differences

in SVL and body mass among A. maculatum. Two general reasons have been presented to

explain these dimorphisms. First, males may exhibit an increased mortality and not survive to

attain a larger size, although this hypothesis does not adequately explain male-biased sex ratios

(as previously discussed). However, female fecundity likely increases with size and thus may be

sexually selected (Shine 1979).

Implications for Conservation

Efforts to demonstrate loss of secondary production in wetlands using endemic

amphibians has not been well documented (Gibbons et al. 2006), and this study provides

50
baseline data showing that Ambystoma maculatum and A. opacum production in LT6 does indeed

dominate energy transfer from the aquatic to terrestrial habitat as compared to other local

herpetofaunal species. Future variations in dominance among amphibian species may be useful

in elucidating underlying changes in habitat quality at LT6. Indeed, Ambystoma opacum is given

a high coefficient of conservation (9 on a 1-10 scale) by the Ohio EPA’s Amphibian Index of

Biotic Integrity (Micacchion 2004), because A. opacum is sensitive to disturbance (relative to

other sympatric amphibian species) and requires mature forests in order to thrive. Importantly,

reptiles are also an integral portion of productivity at LT6, and similar to A. opacum, some

species may require undisturbed forests as well. Reptile species (e.g., Terrapene carolina and

Trachemys scripta) with less than 5 total captures also ranked high in relative biomass compared

to other species with over 100 total captures. This further reiterates the importance of forested

buffers to maintain species diversity because reptiles generally have a larger home range and are

more mobile than their amphibian counterparts (Gibbons et al. 2000, Semlitsch and Bodie 2003).

51
 Table 3.1

Sex ratios for Ambystoma maculatum and A. opacum reported in previous studies and this study
(Sept. 2007-May 2010).

Taxon Location Males Females Sex X2 test Reference

ratio (P)*

A. maculatum St. Joseph Co., IN 244 84 2.9 <0.001* Peckham and Dineen 1954
136 51 2.7 <0.001*
Ann Arbor, MI 95 80 1.19 NS Husting 1965
151 109 1.39 0.01*
56 55 1.02 NS
220 119 1.85 <0.001*
Kingston, RI 72 14 5.14 <0.001* Whitford and Vinegar 1966
82 30 2.7 <0.001*
119 34 3.5 <0.001*
St. Louis Co., MO 300 128 2.34 <0.001* Sexton et al. 1986
592 409 1.45 <0.001*
868 340 2.55 <0.001*
381 248 1.54 <0.001*
237 136 1.78 <0.001*
132 93 1.42 .0093*
Orange Co., NC 39 32 1.22 NS Stenhouse 1987
St. Louis Co., MO 510 135 3.8 <0.001* Phillips and Sexton 1989
Alexander Co., IL 8 10 0.8 NS Parmelee 1993
Hamilton Co., TN 635 234 2.714 <0.001* This study
115 107 1.075 NS
428 162 2.642 <0.001*
A. opacum Orange Co., NC 174 52 3.35 <0.001* Stenhouse 1987
Alexander Co., IL 7 11 0.64 NS Parmelee 1993
Aiken Co., IL 769 175 4.4 <0.001* Krenz and Scott 1994
Hamilton Co., TN 131 133 0.985 NS This study
238 232 1.026 NS
48 76 0.590 0.01*
* Ratios significantly different (p≤0.05) from 1 male per 1 female are presented.

52
 Table 3.2

Relative biomass (g) of species captured (Sept. 2007-May 2010) and corresponding percent
biomass.

Species Relative biomass (g) Percent biomass (%)

Ambystoma maculatum 35,783.96 61.142

Ambystoma opacum 15,419.3 26.346
Terrapene carolina 1,699.0 2.903
Lithobates catesbeianus 1,271.44 2.172
Lithobates sphenocephalus 676.0 1.155
Trachemys scripta 573.0 0.979
Agkistrodon contortrix 522.0 0.892
Notopthalmus viridiscens 493.0 0.842
Lithobates clamitans 438.0 0.748
Chelydra serpentina 359.0 0.613
Chrysemys picta 356.0 0.608
Kinosternon subrubrum 259.0 0.443
Coluber constrictor 193.0 0.330
Pseudacris triseriata 142.74 0.244
Pseudacris crucifer 139.0 0.237
Anaxyrus woodhousii 123.44 0.211
Gastrophryne carolinensis 28.0 0.048
Plestiodon fasciatus 15.5 0.026
Scaphiopus holbrookii 14.0 0.024
Plestiodon laticeps 5.0 0.009
Anaxyrus americanus 5.0 0.009
Aspidoscelis sexlineatus 4.0 0.007
Plestiodon inexpectatus 4.0 0.007
Opheodrys aestivus 3.0 0.005
Total 58,526.38 99.998%

53
 Table 3.3

Snout-vent length (mm) (mean ± 1SD), tail length (mm) (mean ± 1SD), and body mass (g)
(mean ± 1SD) of salamander species (Sept. 2007-May 2010).

Taxon Snout-vent length Tail length (mm) Body mass (g)

(mm) (n) (n)
(n)

Ambystoma maculatum

Male 97.155±10.592 92.37±9.035 18.428±3.484

(1121) (1131) (1106)

t=14.20, p<0.0001* t=4.15, p<0.0001* t=16.78, p<0.0001*

Female 103.6±8.494 94.209±9.188 22.28±5.332

(682) (681) (682)

Ambystoma opacum

Male 68.083±7.755 46.655±6.70 7.565±2.063

(777) (772) (771)

t=3.53, P<0.005* t=2.00, p<0.05 t=13.13, p<0.0001

Female 69.280±5.801 47.303±6.438 8.987±2.344

(895) (894) (886)

Juvenile 43.330±2.014 34.003±2.746 2.177±0.557

(694) (694) (694)

Notophthalmus
viridiscens
21.56±6.865 20.85±7.987 1.083±0.327
Juvenile (455) (455) (458)

* Significantly different results between males and females are presented.

54
 Table 3.4

Snout-vent length (mm) (median ± IQR) and body mass (g) (median ± IQR) of anuran species
(Sept. 2007-May 2010).

Taxon Snout-vent length (mm) Body mass (g)

(n) (n)

Acris crepitans 23 ± 20-25 Not reported

(7)

Acris gryllus 20 ± 19-22.5 Not reported

(11)

Anaxyrus woodhousii 33 ± 30-38 4 ± 3-5

(25) (25)

Gastrophryne carolinensis 29.5 ± 27-34 3 ± 2-3

(10) (10)

Pseudacris crucifer 13 ± 12-15 Not reported

(138)

Pseudacris triseriata 27 ± 18-30 1 ± 1-2

(91) (91)

Lithobates catesbeianus 44 ± 40-48 8 ± 6-12

(135) (132)

Lithobates clamitans 37 ± 33-41.5 3 ± 2-6

(92) (90)

Lithobates sphenocephalus 31 ± 29-36 2 ± 2-3

(220) (219)

55
 CHAPTER IV

POPULATION ESTIMATES AND PHENOLOGY OF AMBYSTOMA OPACUM

AND A. MACULATUM

Abstract

Population estimates and phenology are critical tools in understanding the viability and

stability of species. Thus, I utilized drift fencing and pitfall traps from 15 September 2007 to 1

May 2010 to capture and mark as many Ambystoma opacum and A. maculatum as possible.

Total captures of A. maculatum individuals were 1,514, 505, and 513 for 2008, 2009, and 2010,

respectively. Recaptures for each respective year were 342, 47, and 47. As shown in previous

studies, A. maculatum were most active during January and February of each study year.

Population estimates of A. maculatum ranged from a minimum of 1,877 (Schnabel-Schumacher

method) to a maximum of 3,085 (Caughley method) individuals. Total individual captures of A.

opacum were 679, 728, and 405 individuals for 2007, 2008, and 2009, respectively. Recaptures

for each respective year were 73, 180, and 35. Estimates of A. opacum population sizes ranged

from a minimum of 913 (Schnabel-Schumacher method) to a maximum of 2,478 (Caughley

method) individuals. Most individuals were active during October, with juveniles emigrating in

May. Baseline population estimates produced in this study must be compared to future estimates

to determine if fluctuations among study years are typical and caused by natural factors.

56
Introduction

Repeated population estimates of amphibians are useful for predicting future stability and

survival (Blaustein et al. 1994). A high degree of philopatry among amphibians (e.g.,

Ambystoma, Anaxyrus, Hyla, Lithobates, and Pseudacris species [Lannoo et al. 1998]) allows for

numerous individuals to be observed in a relatively short time. Numerical changes that

investigators observe in natural populations may be caused by many natural and anthropogenic

factors that influence the life history of the organism (Blaustein et al. 1994). Indeed, when

golden toads (Incilius periglenes) in Costa Rica were first believed to have drastically reduced in

number, it was noted that these organisms may live 10+ years and perhaps were aestivating until

weather conditions became more favorable (Crump et al. 1992). Hence, quantitative counts may

be difficult to obtain but are needed to adequately assess population size. Population studies are

also challenging because many years are needed to adequately identify patterns and fluctuations

(Blaustein et al. 1994); thus, researchers often report raw census data rather than actual estimates

of population size. However, Amybstoma is an ideal study organism for this type of question

because temporal breeding patterns are often the same each year. Unfortunately, studies

reporting population estimates of Ambystoma maculatum and A. opacum are notably absent,

perhaps because the use of mark-recapture methods must be carefully planned in order to meet

the assumptions of any given model (Phillips et al. 2001).

Phenology is directly useful to maximizing the success of field studies on amphibians

(Paton and Crouch 2002), and is known to vary widely among species. Variation may be present

within a single genus, with Ambystoma annulatum and A. opacum utilizing wetlands heavily

during autumn and A. maculatum breeding in wetlands only during late winter to early spring

(Petranka 1998). Anurans also exhibit wide phenological diversity, with Lithobates catesbeianus

57
beginning breeding as early as January in the southern United States or as late as July in Iowa

(Willis et al. 1956). Likewise, juvenile phenology is an important parameter to consider in

conservation efforts because the length of time until metamorphosis also varies by locale

(Hocking et al. 2008).

Although specific dated chronologies do not abound in the literature, phenology in the

genus Ambystoma is generally well studied (Peckham and Dineen 1954, Husting 1965, Paton et

al. 2000). Previous studies conducted in Michigan, Indiana, and Missouri on A. maculatum have

observed the largest number of immigrating salamanders in March (Husting 1965, Peckham and

Dineen 1954, Sexton et al. 1990, respectively). A population of A. maculatum in Rhode Island

was observed to immigrate in March and emigrate in April (Paton et al. 2000). This pattern is

similar to a Missouri population that underwent the majority of its movements in March and

April (Hocking et al. 2008). Although phenology is further dependent upon various climatic

factors (Semlitsch 1985), it is especially important for developing sampling strategies that can

best utilize resources during predictable, seasonal movements.

Thus, my research objectives were to use mark-recapture methods to generate population

estimates of Ambystoma maculatum and A. opacum and determine phenology of these species in

LT6.

Materials and Methods

Drift fences have proven to be a useful tool in sampling ambystomatid salamanders (see

Chapter 1) and are considered the most effective method for recording individuals (Dodd and

Scott 1994). Blaustein and others (1994) suggest that mark and release of as many individuals as

possible is the best method for monitoring amphibian populations. Thus, mark-recapture

58
methods were used in conjunction with the pitfall arrays so individual identification would be

possible. During the autumn of 2007, photographs were also used as an identification tool for A.

opacum. Photograph identification based upon general coloration patterns has been utilized in

numerous taxa (see references in Gamble et al. 2008 for a thorough list). Manual matching used

in this study was extremely time intensive, with ~800 photographs requiring 50+ hours to

effectively analyze. Although Gamble et al. (2008) utilized photo-recognition technology and

processed over 1,000 A. opacum images in 18 hours, lack of access to this recognition software

(coupled with time-intensive manual matching) was cause for changing methods of individual

salamander identification. Toe-clipping in amphibians has proven to be cost-effective and

lowers the handling time of animals (Ott and Scott 1999), and became the method of choice for

individual identification in the present study. The numbering schemes of D.B. Wake (Donnelly

et al. 1994) and B. Martof (1953) were used. The Wake scheme was used to mark the first 130

marbled salamanders. The Martof scheme was utilized for all remaining marks due to the ease

of memorizing the toe sequence for use while in the field. The first 130 salamanders were also

identified by photographs during the first field season, in order to ensure that the integrity of the

data was maintained. Integrity was further verified by referring to the Wake scheme when

individuals were recaptured that did not logically follow the Martof scheme. All marking was

performed in an appropriate laboratory environment at UTC. Toe-clipping instruments were

disinfected before each use with isopropyl alcohol (70%), and toes were retained in

microcentrifuge tubes in ethanol (95%) for analysis in future studies. Once marked, animals

were returned to the study site within 24 hours and released on the opposite side of the fence

where they were originally captured.

59
 Recapture data were totaled by breeding seasons for both Ambystoma maculatum

(generally January through May) and A. opacum (generally October through December). Results

were analyzed by breeding season because amphibians have the ability to regenerate clipped

digits, making cross-year estimates difficult (Peckham and Dineen 1954). The Schnabel-

Schumacher and Caughley estimates were used to determine population size. Mortality was

included because the Schnabel-Schumacher method accomodates this. The Caughley estimator

is a general, empirical model (Krebs 1999) and assumes that little to no mortality occurs during

sampling. Caughley estimates were thus utilized only during seasons when the total number of

observed deaths was less than 10 individuals. Both recapture models were calculated using

Ecological Methodology version 5.2 (Krebs 1999). Monthly percentages (relative to the yearly

total) for recorded Ambystoma individuals were calculated as an indication of movement activity.

Chi-square analysis was utilized to determine if ingress and egress totals differed significantly

from a 1:1 ratio.

Results

Recaptured individuals of Ambystoma maculatum accounted for 22.59% of the totals

captured in 2008 (Table 4.1). Recaptured individuals accounted for 9.31% and 9.16% of the

totals captured in 2009 and 2010, respectively. Population estimates ranged from 1,877

individuals (95% CI=1,248-3,785) using the Schnabel-Schumacher estimate to 3,085 (95%

CI=2,313-4,188) using the Caughley estimate. Recaptured individuals of A. opacum accounted

for 10.75%, 24.73%, and 8.64% of the totals sampled in 2007, 2008 and 2009, respectively.

Population estimates ranged from 913 individuals (95% CI=740-1,191) using the Schnabel-

60
Schumacher estimate to 3,030 individuals (95% CI= 1,989-6,347) using the Schnabel-

Schumacher estimate (Table 4.1).

Significantly different numbers of immigrating versus emigrating individuals were

observed during each study year (p<0.001), except for A. maculatum in 2009 (Tables 4.2 and

4.3). The greatest numbers of A. opacum movements were observed in October of each study

year. The greatest numbers of A. maculatum movements were observed in January of each study

year, except 2008-2009, when a large movement of emigrating adults was observed in February

and March (Table 4.2).

Discussion

The recapture-based population estimates produced by this study are the first estimates to

be generated for ambystomatid salamanders in the greater metropolitan Chattanooga area, and

offer a unique insight into the ecology of LT6. These baseline data are critical because future

estimates can be compared to this study and a better assessment of anthropogenic effects can be

produced. Although phenology records indicate that not all individuals leaving the wetland were

recaptures, I believe that my estimations of population size are still reliable. Studies do not

require the entire wetland to be encompassed (Willson and Gibbons 2009) because the

comparison of marked individuals versus unmarked individuals is the key component when

estimating population size. It is likely that unmarked salamanders observed exiting the wetland

were already residing inside the drift array when it was installed. Individual Ambystoma opacum

could easily have remained with the drift array after metamorphosing and avoided detection until

a later date; indeed, LT6 has an abundance of habitat and structure occurring within the drift

array (e.g., damp logs, fallen trees) that could readily be utilized by both juvenile and adult

61
salamanders. However, it should also be assumed that some individuals did trespass or enter

the array at locations where the fence was incomplete or difficulties prevented installation.

While this scenario likely occurred to a small extent, both A. opacum and A. maculatum are

known to utilize the same terrestrial routes entering and exiting a wetland (Stenhouse 1985).

Further, they are known to utilize the same paths for consecutive breeding years (Stenhouse

1985). Encounters with unmarked A. maculatum exiting the wetland was most likely due to a

single factor—the wetland held more water during the winter months, and some sections of the

array were completely underwater when A. maculatum exited. The aquatic nature and swimming

ability of A. maculatum may have facilitated movement and escape in areas without fencing—

adults were seen swimming within open wetland waters on numerous occasions.

Implications for Conservation

Genera such as Ambystoma, which have highly predictable, seasonal movements, are

ideal study organisms for monitoring population fluctuations, and phenology is a critical tool

when developing conservation strategies. Yet, for many North American species only qualitative

data is available regarding seasonal movements (Paton and Crouch 2002). Generally, both A.

maculatum and A. opacum utilized rainy nights heavily for movements during this study, as

previously observed by numerous investigators (Blanchard 1930, Husting 1965, Sexton et al.

1990). Thus, phenology is indeed critical in deciding when sampling should begin. The

majority of A. maculatum individuals immigrated in large aggregates during January of all years,

except 2009 (where the majority individuals immigrated in February and March)—two months

in advance of both Missouri and Rhode Island populations (Hocking et al. 2008, Paton et al.

2000). Understanding these temporal movements on a regional scale and across a wide range of

62
species allows land managers to appropriately plan activities such as timber harvesting, road

closures, or controlled burns (Hocking et al. 2008). These movements should also be considered

in conjunction with wetland size and hydroperiod (Paton and Crouch 2002). Although

hydroperiod was not quantitatively measured during the present study, future assessments at LT6

may be able to correlate movement phenology and reproductive success with this parameter.

Estimates of population size can provide an important perspective to help determine

which wetland areas are in the greatest need of conservation. Indeed, estimates of Ambystoma in

the present study offer a unique insight into the stability of LT6. However, short-lived species

(e.g., Acris crepitans and Pseudacris crucifer) may also be an important indicator of stability

because they cannot easily postpone breeding if conditions become unfavorable (Lannoo 1998).

Although spring peepers are resilient and known to occur in urbanized areas (Gibbs 1998), they

are initially sensitive to breeding habitat disturbance (Micacchion 2004). Thus, closely

monitoring small frog species in future years may initially give better indications of negative

ecological impacts—either natural or anthropogenic—in LT6.

63
 Table 4.1

Population estimates of Ambystoma maculatum and A. opacum for each study period (Sept.
2007-May 2010).

Taxon Method Total Percent Population 95%

Captures Recaptures Estimate Confidence
(Recaptures) Interval

Ambystoma maculatum

2008 Schnabel- 1,514 (342) 22.59% 2,432 2,111 – 2,870

Schumacher

2009 Schnabel- 505 (47) 9.31% 1,877 1,248 – 3,785

Schumacher

Caughley 505 (47) 9.31% 3,085 2,313 – 4,188

2010 Schnabel- 513 (47) 9.16% 2,127 1,820 – 2,560

Schumacher

Caughley 513 (47) 9.16% 2,925 2,236 – 3,892

Ambystoma opacum

2007-2008 Schnabel- 679 (73) 10.75% 3,030 1,989 – 6,347

Schumacher

2008-2009 Schnabel- 728 (180) 24.73% 913 740 – 1,194

Schumacher

Caughley 728 (180) 24.73% 1,990 1,700 – 2,364

2009-2010 Schnabel- 405(35) 8.64% 1,666 1,187 – 2,795

Schumacher

Caughley 405 (35) 8.64% 2,478 1,816 – 3,450

64
 Table 4.2

Phenology of Ambystoma maculatum captures according to date and movement direction

(entering or exiting the wetland; recaptures included). Percentages of the total adult captures
(per study period) are also given by month as an indicator of seasonal activity.

Month Ingress Egress X2 Test by Total Percentage of

Year (P)* Recaptures Total Captures

2007 – 2008
January 667 109 76 52.40%
February 227 96 84 21.81%
March 92 215 146 20.73%
April 4 71 37 5.06%
SUBTOTAL 990 491 p<0.001* 343 (100.0%)
2008 – 2009
November 1 3 - 0.83%
December 29 - - 6.04%
January - 9 - 1.88%
February 159 19 2 37.08%
March 19 196 25 44.79%
April 6 38 20 9.17%
May - 1 - 0.21%
SUBTOTAL 214 266 NS 47 (100.0%)
2009 – 2010
November 4 2 - 0.69%
December 5 7 - 1.38%
January 519 2 2 59.89%
February 36 - 1 4.14%
March 70 193 30 30.23%
April 1 31 14 3.68%
SUBTOTAL 635 235 p<0.001* 47 (100.0%)

OVERALL 1839 992 p<0.001* 435

TOTAL

* Significant differences between ingress and egress totals (p≤0.05) are presented.

65
 Table 4.3

Phenology of Ambystoma opacum captures according to date and movement

direction (entering or exiting the wetland; recaptures included). Percentages of the total adult
captures (per study period) are also given by month as an indicator of seasonal activity.

Month Ingress Egress X2 Test by Year Recaptures Percentage of

(juveniles) (P)* Total Captures

2007 – 2008
October 267 226 53 57.73%
November 4 144 12 17.33%
December - 2 - 0.23%
January 3 170 8 20.26%
February - 20 - 2.34%
March - 13 - 1.52%
April - 31 - 0.59%
(26)
SUBTOTAL (adults) 274 580 p<0.001* 73 (100.0%)
2008 – 2009
September 4 2 - 0.83%
October 467 44 39 70.68%
November 1 64 43 8.99%
December 1 137 97 19.09%
February 1 - - 0.14%
March - 1 - 0.14%
May - (666) -
July - 3 - 0.14%
2
SUBTOTAL (adults) 474 249 p<0.001* 179 (100.0%)
2009 – 2010
September 28 11 - 11.85%
October 39 188 34 69.00%
November 1 60 4 18.54%
December - 2 1 0.61%
April - (3) -
SUBTOTAL 68 261 p<0.001* 39 (100.0%)

OVERALL TOTAL 816 1090 p<0.001* 291

(adults)

* Significant differences between ingress and egress totals (p≤0.05) are presented.

66
 CHAPTER V

DIRECTIONS FOR THE FUTURE

Amphibian and reptile populations will continue to decline unless large, concerted

conservation strategies are integrated globally. However, these plans must be locally developed

in order to aid both the targeted species and their habitat (Paton and Crouch 2002). The present

study was designed to elucidate herpetofaunal species richness and abundance, as well as

ecological processes affecting sex ratios and sexual dimorphism. However, this study can

provide baseline data for a host of future ecological studies that are worthy of being conducted

within LT6.

Global amphibian declines have often been detected by researchers initially investigating

the predictable breeding movements of anurans on a local scale (Gibbons et al. 2000).

Therefore, long-term monitoring of frog populations at LT6 may shed light on broader

amphibian declines within the Southeast. Indeed, significant numbers of declining species

belong to three anuran families (Hylidae, Bufonidae, and Ranidae), all of which are found at

LT6. Combining drift array sampling with techniques better suited for hylid monitoring (e.g.,

PVC pipe refugia and auditory surveys [Boughton et al. 2000]), will allow more accurate

monitoring of population fluctuations. Bufonids in particular are known to suffer from

‘enigmatic’ declines (Stuart et al. 2005), but studies of Acris creptians blanchardi (Hylidae)

have also been unsuccessful in determining the leading causes of decline (Lehtinen and Skinner

2006). Future studies within LT6, combined with data first reported here, may be able to better

67
determine persistence and whether local stability exists for these species.

In contrast to amphibians, reptile abundance was not rigorously investigated during this

study, although use of a drift fence did produce captures of fourteen species—only two less than

the observed amphibian richness. As previously stated, reptiles have been less investigated in

the literature with respect to habitat alteration, but are equally affected (Gibbons et al. 2000).

However, discerning reptile species richness and abundance trends may be more difficult than

for amphibians; reptiles are generally more mobile and maintain larger home ranges (Gibbons et

al. 2000). Thus, additional methods suited for increasing reptile observations are needed (e.g.,

coverboards, funnel traps, and hoop nets). Yet, only long-term studies will be able to critically

evaluate patterns (Gibbons et al. 2000). Reptiles, too, suffer from enigmatic declines; the eastern

kingsnake (Lampropeltis getula) is but one example of a species that has declined for reasons

largely unknown (Winne et al. 2007). Tennessee has a paucity of data concerning the

conservation status and population biology of reptiles, and use of additional methods at LT6 to

better understand local abundance can greatly improve what is currently known regarding

southeastern reptile declines in various species.

Finally, investigating the possible long-term effects of urbanization surrounding LT6

should not be overlooked. Industrial activity, although not quantitatively monitored during this

study, increased markedly during the overall study period. Few studies have adequately

investigated the effects of urbanization across all age classes of a given species; further, species

that are considered widespread and common are still being locally extirpated before regional

management plans have been developed (Scheffers and Pazskowski 2012). Indeed,

ambystomatids are locally common within LT6, but the family Ambystomatidae continues to be

one of four amphibian families that have significantly more declining species than the average

68
for all amphibians (Stuart et al. 2005). Also, few studies have documented how amphibians and

reptiles move through an urbanized landscape even though metapopulation dynamics are often

considered critical to these populations (Scheffers and Pazskowski 2012). Overall, any efforts to

keep common species common requires a better understanding of each species’ resistance to

habitat change through continued monitoring. It is only with long-term, species-specific data

that fruitful land management plans can be implemented to promote community stability and

diversity.

69
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 VITA

Joseph F. Simpson, III was born in Hamilton County, Tennessee, and has lived in the

Chattanooga area most of his life. He received a B.S. in Biology from UT Chattanooga in 2006.

Joe has always had a strong interest in wildlife, especially native reptiles and amphibians. Joe is

a former teaching assistant and adjunct faculty at UT Chattanooga, and currently is an adjunct

faculty member at Belhaven University, Chattanooga, and a Watershed Project Coordinator for

the Southeast Tennessee Resource Conservation and Development Council. He is an avid fitness

enthusiast, and can also be found spending free time with a personal flock of backyard chickens

or bagpiping at venues throughout the Southeast.

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