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Spatial Ecology and Demographics of a Population of Sternotherus odoratus (Testudines

Permanent Link: http://ufdc.ufl.edu/UFE0022630/00001

Material Information

Title: Spatial Ecology and Demographics of a Population of Sternotherus odoratus (Testudines Kinosternidae) in an Ozark Stream
Physical Description: 1 online resource (52 p.)
Language: english
Creator: Tavano, Joseph
Publisher: University of Florida
Place of Publication: Gainesville, Fla.
Publication Date: 2008

Subjects

Subjects / Keywords: degradation, ecology, freshwater, habitat, missouri, ozark, river, sternotherus, turtle
Interdisciplinary Ecology -- Dissertations, Academic -- UF
Genre: Interdisciplinary Ecology thesis, M.S.
bibliography   ( marcgt )
theses   ( marcgt )
government publication (state, provincial, terriorial, dependent)   ( marcgt )
born-digital   ( sobekcm )
Electronic Thesis or Dissertation

Notes

Abstract: From 1969 through 2004, the turtle community of a 4.7 km section of the North Fork of the White River, Ozark County, Missouri was studied periodically. The Stinkpot, Sternotherus odoratus, was found in all years in low abundance. From 2005 through 2007, I continued this survey to determine the distribution and related parameters, movement patterns, population structure, and demographic parameters of S. odoratus within the research section. Distribution was similar from 1969 through 2006, with expansion to the terminal station of the research section in 2007. Turtle distribution was positively correlated with dense submerged patches of mud and rooted vegetation. Movement between captures from 2004 to 2007 averaged 198 m for males and 102 m for females, with a range of 0 m to 740 m. There was no difference between male and female carapace lengths or weight. There was a significant difference in male and female plastron lengths with females being larger. Sex ratios were slightly female biased but not significantly so. My study provides insight into the effects of land use change and habitat alteration on aquatic ecosystems. Continued habitat alteration, by affecting the behavior and abundance of species, may have consequences not only for S. odoratus but for the entire community.
General Note: In the series University of Florida Digital Collections.
General Note: Includes vita.
Bibliography: Includes bibliographical references.
Source of Description: Description based on online resource; title from PDF title page.
Source of Description: This bibliographic record is available under the Creative Commons CC0 public domain dedication. The University of Florida Libraries, as creator of this bibliographic record, has waived all rights to it worldwide under copyright law, including all related and neighboring rights, to the extent allowed by law.
Statement of Responsibility: by Joseph Tavano.
Thesis: Thesis (M.S.)--University of Florida, 2008.
Local: Adviser: Nickerson, Max A.
Electronic Access: RESTRICTED TO UF STUDENTS, STAFF, FACULTY, AND ON-CAMPUS USE UNTIL 2010-08-31

Record Information

Source Institution: UFRGP
Rights Management: Applicable rights reserved.
Classification: lcc - LD1780 2008
System ID: UFE0022630:00001

Permanent Link: http://ufdc.ufl.edu/UFE0022630/00001

Material Information

Title: Spatial Ecology and Demographics of a Population of Sternotherus odoratus (Testudines Kinosternidae) in an Ozark Stream
Physical Description: 1 online resource (52 p.)
Language: english
Creator: Tavano, Joseph
Publisher: University of Florida
Place of Publication: Gainesville, Fla.
Publication Date: 2008

Subjects

Subjects / Keywords: degradation, ecology, freshwater, habitat, missouri, ozark, river, sternotherus, turtle
Interdisciplinary Ecology -- Dissertations, Academic -- UF
Genre: Interdisciplinary Ecology thesis, M.S.
bibliography   ( marcgt )
theses   ( marcgt )
government publication (state, provincial, terriorial, dependent)   ( marcgt )
born-digital   ( sobekcm )
Electronic Thesis or Dissertation

Notes

Abstract: From 1969 through 2004, the turtle community of a 4.7 km section of the North Fork of the White River, Ozark County, Missouri was studied periodically. The Stinkpot, Sternotherus odoratus, was found in all years in low abundance. From 2005 through 2007, I continued this survey to determine the distribution and related parameters, movement patterns, population structure, and demographic parameters of S. odoratus within the research section. Distribution was similar from 1969 through 2006, with expansion to the terminal station of the research section in 2007. Turtle distribution was positively correlated with dense submerged patches of mud and rooted vegetation. Movement between captures from 2004 to 2007 averaged 198 m for males and 102 m for females, with a range of 0 m to 740 m. There was no difference between male and female carapace lengths or weight. There was a significant difference in male and female plastron lengths with females being larger. Sex ratios were slightly female biased but not significantly so. My study provides insight into the effects of land use change and habitat alteration on aquatic ecosystems. Continued habitat alteration, by affecting the behavior and abundance of species, may have consequences not only for S. odoratus but for the entire community.
General Note: In the series University of Florida Digital Collections.
General Note: Includes vita.
Bibliography: Includes bibliographical references.
Source of Description: Description based on online resource; title from PDF title page.
Source of Description: This bibliographic record is available under the Creative Commons CC0 public domain dedication. The University of Florida Libraries, as creator of this bibliographic record, has waived all rights to it worldwide under copyright law, including all related and neighboring rights, to the extent allowed by law.
Statement of Responsibility: by Joseph Tavano.
Thesis: Thesis (M.S.)--University of Florida, 2008.
Local: Adviser: Nickerson, Max A.
Electronic Access: RESTRICTED TO UF STUDENTS, STAFF, FACULTY, AND ON-CAMPUS USE UNTIL 2010-08-31

Record Information

Source Institution: UFRGP
Rights Management: Applicable rights reserved.
Classification: lcc - LD1780 2008
System ID: UFE0022630:00001


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SPATIAL ECOLOGY AND DEMOGRAPHICS OF A POPULATION OF .Sicinotheins //
odoratus (TESTUDINES: KINOSTERNIDAE) INT AN OZARK STREAM




















By

JOSEPH J. TAVANO


A THESIS PRESENTED TO THE GRADUATE SCHOOL
OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT
OF THE REQUIREMENTS FOR THE DEGREE OF
MASTER OF SCIENCE

UNIVERSITY OF FLORIDA

2008


































O 2008 Joseph J. Tavano




































To Amber, who always knew I could do it.









ACKNOWLEDGMENTS

I would like to thank Drs. Max Nickerson, Mike Moulton, and Kent Vliet for their

guidance, advice and recommendations in the writing of this thesis. I would like to thank

Gustavo Vasques for assistance with the GIS, and Dr. Kenney Krysko for help with the statistical

analyses. I thank all of the wonderful people I met while in the field in Missouri: Connie

Morgan and Randy Padgett for their friendship and hospitality; Amy and Justin Spencer at

Sunburst Canoe Ranch for providing camping accommodations; Jeff Ettling, Eric Miller, Mark

Wanner, and everyone else at the Saint Louis Zoo for all of their help and support through the

years; and Karen and Ron Goellner, two wonderful people and great friends, without whom this

research would not have been possible. I would like to thank the Saint Louis Zoological Park

and the Reptile and Amphibian Conservation Corps for providing the funding for this research.

I thank my parents, my grandmother, my brothers and their families for all of their

patience, encouragement and support over the years. Finally, I would like to thank Amber Pitt

for her invaluable help both in the field and in reviewing this thesis.












TABLE OF CONTENTS


page

ACKNOWLEDGMENTS .............. ...............4.....


LIST OF TABLES .........__... ......._. ...............6....


LIST OF FIGURES .............. ...............7.....


AB S TRAC T ......_ ................. ............_........8


CHAPTER


1 INTRODUCTION ................. ...............9.......... ......


The Sti nkp ot, .9t-neshe- u slr~ odoratus (Latrei lle) ................. ...............10..........
.Yievl mothe-i usl odoratus in M issouri ................. ...............14........... .

Justification and Obj ectives ................. ...............16................


2 SPATIAL ECOLOGY AND DEMOGRAPHICS OF A POPULATION OF
STERNOTHERUS ODORATUS (TESTUDINES: KINOSTERNIDAE) IN AN
OZARK STREAM .............. ...............17....


Introducti on ........._._... ....__.. ...............17....
M materials and M ethods .............. ...............17....

Study Site............... ...............17..
Capture M ethods .............. ...............20....
M parking M ethods .............. ...............21....
Data Collection ........._..... ...._... ...............21.....
Data Analysis............... ...............22
Re sults........._..... ...._... ...............22.....
Distribution ........._..... ...._... ...............22....
H habitat .............. ...............30....
M ovem ent ........._...... .. ......._. ........ .............3

Population Structure and Demographics ......... ......._.._.._ ......... ............3
D discussion ................... ........._._ ...............37......
Distribution and Habitat .............. ...............37....
M ovem ent ........._...... .. .......... ........ .............4

Population Structure and Demographics ......... ......._.._.._ ......... ............4
Conclusions............... ..............4


3 CONCLUSIONS .............. ...............45....


LIST OF REFERENCES ....._.._................. ........_.._.........4


BIOGRAPHICAL SKETCH .............. ...............52....










LIST OF TABLES


Table page

2-1 Behavior of .Yievl ul~rr,~ sth-i odoratus at site of capture, North Fork of the White River,
Ozark County, Missouri............... ...............30

2-2 Distance traveled by recaptured .%eiesl; White River, Ozark County, Missouri, 2004-2007 ........__............_. ........._.._. ...31

2-3 Mean carapace length (CL), plastron length (PL) and weight (WT) of Sternotherus
odoratus in the North Fork of the White River, Ozark County, Missouri, 2005 2007....32

2-4 Test for difference in male versus female carapace length of.rl .%inesdiciit s odoratus
in the North Fork of the White River, Ozark County, Missouri, 2005-2007. ........._.........32

2-5 Test for difference in male versus female weight of.%e~linesdiciit s/ odoratus in the
North Fork of the White River, Ozark County, Missouri, 2005-2007. ........._..... .............32

2-6 Test for difference in male versus female plastron length of.rl .tinesdlreiits odoratus in
the North Fork of the White River, Ozark County, Missouri, 2005-2007. ................... .....32

2-7 Sex ratios and chi-square statistics for Sternotherus odoratus, North Fork of the
White River, Ozark County, Missouri, 2005-2007 .....__.___ .........__ ........_.... ....33










LIST OF FIGURES


1-1 Distribution of the Stinkpot, .%ei~ mothe-l~ us odoratus ................. ........__........._... 10

1-2 Distribution of .%ei~ mothei,~ us odoratus in Missouri ........._.._ ...... .___ ......_........1

2-1 Stations 0-50 of the 4.7 km research section of the North Fork of the White River,
Ozark County, Missouri............... ...............19

2-2 Distribution of Sternotherus odoratus within the 4.7 km study section of the North
Fork of the White River, Ozark County, Missouri .......... ................ ................ ...23

2-3 Size class distribution by carapace length for Sternotherus odoratus captured in the
4.7 km research section of the North Fork of the White River, Ozark county
M i ssouri. ........._ ......._. ...............33...









Abstract of Thesis Presented to the Graduate School
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Master of Science

SPATIAL ECOLOGY AND DEMOGRAPHICS OF A POPULATION OF .Sicinestheins //
odoratus (TESTUDINES: KINOSTERNIDAE) INT AN OZARK STREAM


By

Joseph J. Tavano

August 2008

Chair: Max A. Nickerson
Major: Interdisciplinary Ecology

From 1969 through 2004, the turtle community of a 4.7 km section of the North Fork of

the White River, Ozark County, Missouri was studied periodically. The Stinkpot, .Sicinestheins //

odoratus, was found in all years in low abundance. From 2005 through 2007, I continued this

survey to determine the distribution and related parameters, movement patterns, population

structure, and demographic parameters of S. odoratus within the research section.

Distribution was similar from 1969 through 2006, with expansion to the terminal station of

the research section in 2007. Turtle distribution was positively correlated with dense submerged

patches of mud and rooted vegetation. Movement between captures from 2004 to 2007 averaged

198 m for males and 102 m for females, with a range of 0 m to 740 m. There was no difference

between male and female carapace lengths or weight. There was a significant difference in male

and female plastron lengths with females being larger. Sex ratios were slightly female biased but

not significantly so.

My study provides insight into the effects of land use change and habitat alteration on

aquatic ecosystems. Continued habitat alteration, by affecting the behavior and abundance of

species, may have consequences not only for S. odoratus but for the entire community.









CHAPTER 1
INTTRODUCTION

Turtles date back in the fossil record 210 million years to the late Triassic, pre-dating the

dinosaurs (Gaffney, 1990). Turtles have evolved several unique adaptations, the most

characteristic being the shell that encases the body. Extant turtles, or chelonians, are long-lived

organisms, and may have the longest life spans of any extant vertebrate (Ernst et al., 1994).

Giant tortoises have been known to live 120 to 150 years, and the oldest known box turtle lived

123 years (Pritchard, 1967).

There are approximately 300 species of chelonians. Turtles can be found worldwide in

terrestrial, aquatic, and marine habitats except at high latitudes or altitudes (Pough, 2004).

Turtles, and river turtles in particular, can comprise a significant portion of the biomass of

ecosystems they inhabit (Congdon et al., 1986; Moll and Moll, 2004). They serve as predators,

frugivores, seed dispersers, and scavengers, as well as prey for crocodilians and some birds and

mammals (Moll, 1990; Ernst et al., 1994; De Lima et al., 1997; Ford & Moll, 2004). These roles

act as important linkages both within and between terrestrial and aquatic food webs (Moll and

Moll, 2004). Turtles also play a vital role in nutrient cycling, maintenance of water quality, and

energy flow (Moll and Moll, 2004). Moreover, turtle species with specialized diet or habitat

requirements can serve as indicators of habitat quality (Ernst et al., 1994).

Despite their unique characteristics and the important impacts they have on their

ecosystems, turtles remain largely understudied. Many turtle populations are believed to be in

decline (Moll, 1986; Ernst et al., 1994; Buhlman and Gibbons, 1997; van Dijk et al., 2000), yet

not enough is known about the natural and life histories of many species, nor have there been

sufficient long-term population studies to support that claim (Congdon et al., 1993; Dodd and

Franz, 1993; Moll and Moll, 2004). Chelonians face many threats to survival including









collection for the pet trade, utilization as a food resource, and habitat loss and alteration (Ernst et

al., 1994; Buhlman and Gibbons, 1997; van Dijk et al., 2000; Moll and Moll, 2004).

The Stinkpot, Sternotherus odoratus (Latreille)

The Stinkpot, .Sicinestheins // odoratus, is a relatively small aquatic turtle found throughout

North America from Maine to Florida, and west to Wisconsin and Texas (Ernst et al., 1994;

Figure 1-1). Although it is considered to be a chiefly nocturnal species (Ernst, 1986), some

studies have found peak activity periods to be bimodal. April to September activity periods in

both Oklahoma and Pennsylvania have been reported between 0400 h to 1100 h and 1700 h to

2100 h (Mahmoud, 1969; Ernst, 1986). Peak activity for the rest of the year was between 1000 h

and 1600 h (Mahmoud, 1969; Ernst, 1986).























Figure 1-1. Distribution of the Stinkpot,.rl Siimotheius/ odoratus. Reprinted from Ernst, C.H., J.E.
Lovich, and R.W. Barbour. 1994. Turtles of the United States and Canada.
Smithsonian Institution Press, Washington D.C. and London. P. 142. Used with
permission of the author.









.Srlilcim 'Ithins odoratus are omnivorous. Diet analyses reveal consumption of earthworms,

leeches, clams, snails, crayfish, insects, minnows, tadpoles, algae, plants, seed, seed pods, fish

eggs, and carrion (Lagler, 1943; Mahmoud, 1969; Ernst et al., 1994; Palmer and Braswell, 1995;

Davis, 1996; Ford, 1999). Diets of Oklahoma Stinkpots consisted primarily of insects, snails,

and aquatic vegetation (Mahmoud, 1969). Snails also represented a significant part of the diet of

S. odoratus in Wisconsin (Vogt, 1981), Michigan (Lagler, 1943), and Florida (Bancroft et al.,

1983).

.Srlilcim 'Ithins odoratus serve as prey for numerous species. Eggs are eaten by snakes

(Lampropeltis, Cemophora), raccoons (Procyon lotor), skunks (M~ephitis mephitis), and herons

(Ernst et al., 1994), and adults and juveniles are preyed upon by raccoons, foxes (Vulpes vulpes),

American Alligators (Alligator mississippiensis), Common Snapping Turtles (Chelydra

serpentina), Bald Eagles (Haliaeetus leucocephalus), Cottonmouths (Agkistrodon piscivorous),

water snakes (Nerodia sp.), Largemouth Bass (M~icropterus salmoides), and wading birds (Clark,

1982; Delaney and Abercrombie, 1986; Pritchard, 1989; Ernst et al., 1994; Mitchell, 1994;

Palmer and Braswell, 1995).

Although many species of freshwater turtles exhibit pronounced sexual size dimorphism,

sexual dimorphism in S. odoratus is not as pronounced. Tinkle (1961) reported that sexual

dimorphism in S. odoratus body size is only seen in southern populations. This observation is

supported by Bancroft et al. (1983) and Meshaka (1988) who found that females in Florida have

longer carapace lengths (CL) than males. In contrast, Edmonds and Brooks (1996) found males

in a population in Ontario to have significantly longer carapace lengths than females. They

hypothesized that this dimorphism was a result of selection for larger male body size in response

to combat related competition for mates (Edmonds and Brooks, 1996). Aside from body size,









tails of mature males are thicker and longer than that of females, and males develop elevated

scale patches on the inner hind legs (Risley, 1930).

Size at maturity varies with location and sex. In Michigan, Risley (1933) and Tinkle

(1961) found that males reached maturity in three or four years at 60-70 mm carapace length

(CL). Risley (1933) found that females matured between ages nine and eleven at 80 mm CL,

while Tinkle (1961) found mature females between two and seven years old, also at 80 mm CL.

Mitchell (1988) in Virginia reported seeing male secondary sex characteristics at two years and

51 mm CL, and females at 4 years and 66 mm CL. Mahmoud (1969) saw mature males at 65

mm CL between four and seven years of age and females matured between 65 mm and 85 mm

CL at five to eight years of age in Oklahoma.

Sex ratios for populations vary widely, but generally follow the pattern of being male-

biased in northern populations and female-biased in southern populations (Iverson and Meshaka,

2006). Mahmoud (1969) and Mitchell (1988) both found sex ratios not significantly different

from 1:1 in Oklahoma and Virginia, respectively. An Ontario population consisted of 72%

males (Edmonds and Brooks, 1996), and Dodd (1989) reported a sex ratio of 1:2.8 males to

females in Alabama. A range of explanations have been proposed to account for the disparity in

sex ratios, including differential mortality of nesting females (Edmonds and Brooks, 1996;

Iverson and Meshaka, 2006), sampling bias as a result of trapping technique or behavioral

differences (Dodd, 1989; Edmonds and Brooks, 1996; Iverson and Meshaka, 2006), and

temperature-dependent sex determination of hatchlings (Edmonds and Brooks, 1996; Iverson and

Meshaka, 2006).

Sicinothelr'lins odoratus can be long-lived. Ernst (1986) observed two individuals in his

field study, a male and a female, who were known to be 27 and 28 years old at the date of last










capture, and a specimen at the Philadelphia Zoo lived for 54 years, 9 months (Snider and Bowler,

1992).

.Siclinother'lins odoratus can be found in most any body of freshwater, including ponds and

lakes, rivers and streams, urban lakes, reservoirs, and other man-made impoundments, although

Iverson and Meshaka (2006) report a preference for permanent bodies of water with little or no

current. While S. odoratus has been found on rocky and gravelly substrates (Mahmoud, 1969;

Emnst et al., 1994; Palmer and Braswell, 1995), this species is most commonly found in the soft-

bottomed littoral zones of lentic systems with abundant submerged vegetation (Kingsbury,

1993).

Thermoregulatory basking in S. odoratus typically occurs while submerged in shallow

water or floating at the surface, or while partially submerged atop lily pads or mats of algae or

emergent vegetation (Emnst, 1986; Ernst et al., 1994; Iverson and Meshaka, 2006). Basking also

occurs on logs, stumps, or trees overhanging the water, with some individuals climbing as high

as three meters above the water's surface (Vogt, 1981; Ernst, 1986; Nickerson, 2000).

Several studies have examined movement of S odoratus. Most studies reveal a pattern of

a larger home range in males than females (Mahmoud, 1969; Ernst, 1986; Edmonds, 1998).

Average distance traveled between captures by males and females ranges from 69 m and 51 m,

respectively, in an Oklahoma stream (Mahmoud, 1969) to 2052 m and 828 m, respectively, in an

Ontario lake (Edmonds, 1998).

Most research on S. odoratus has been carried out in closed bodies of water such as lakes

or reservoirs, with several exceptions. Emst (1986) sampled a small creek in Pennsylvania

together with marsh, floodplain pool, and mill pond habitats. Mahmoud (1969) studied S.

odoratus in a small stream (average depth and width, approximately 1 m and 3 m, respectively)










in Oklahoma, and Meylan et al. (1992), studied the turtle community of Rainbow Run in Marion

County, Florida, with an emphasis on two kinosternid turtles, the Loggerhead Musk Turtle,

.Sicinesther'lins minor, and S. odoratus.

Sternotherus odoratus in Missouri

.Siclinesthe'lins odoratus is found statewide in Missouri except in the northern and

northwestern third of the state (Johnson, 2000; Figure 1-2). The species is most commonly

encountered in slow-moving sections of larger rivers and streams in the Ozarks, the sloughs and

backwaters of the Bootheel, and rivers in the northeast part of the state (Johnson, 2000).

Missouri's smallest turtle species, S. odoratus typically range in size from 51 mm to 115 mm

carapace length (Johnson, 2000), with the Missouri record specimen measuring 117 mm

carapace length (Powell et al., 1982).

Research on S. odoratus in Missouri has been restricted mainly to two studies (Davis,

1996; Ford, 1999) that investigated reproduction, diet, and demographics. Davis (1996) studied

the effects of cold water effluent on S. odoratus diet and reproduction in a reservoir in southwest

Missouri. Ford (2000) gathered data on foraging ecology and demographics of a population of S.

odoratus in a lake created by the damming of a river, also in southwest Missouri. Ford (1999)

found that there was seasonal and sexual variation in the diet of S odoratus in Missouri, with

females eating more grass seeds, Heliosoma snails, crayfish, mayflies, beetles, isopods, Ludwuigia

seeds, clams, and overall animal matter, and males consuming more nuts, water striders, and

lepidopterans. Seed consumption in September was higher than for June-August, larger volume

of vegetative material was consumed in July and August compared to June, and more crayfish,

snails, clams, and insects were consumed in June.


































Figure 1-2. Distribution of Sicinotheinsl // odoratus in Missouri (Johnson, 2000). Range map for
Sternotherus odoratus reproduced from "The Amphibians and Reptiles of Missouri."
p. 182. Copyright 2000 by the Conservation Commission of the State of Missouri.
Used with permission.



From 1969 through 1980, M. A. Nickerson conducted a mark-recapture study of the turtle

community in a 4.7 km section of the North Fork of the White River (NFWR), Ozark County,

Missouri (Pitt, 2005). Pitt' s (2005) analyses revealed that S. odoratus was present in the research

section, but in very low numbers. Pitt (2005) reassessed the turtle community of NFWR in 2004

with a focus on the Northern Map Turtle, Guraptemys geoguraphica, historically the most

abundant turtle in this community (Nickerson, unpubl. data). She found that whereas the river

and surrounding terrestrial habitat had undergone significant changes since Nickerson's surveys

from 1969 through 1980, the number of S. odoratus captured was similar (Nickerson, unpub.

data; Pitt, 2005).










Justification and Objectives

Although several ecological studies of S. odoratus have been conducted across its

geographic range, little data exist on the life history of S. odoratus in one of its common habitats-

the river ecosystem. Likewise, there has been only one published long-term study of a S

odoratus population (Ernst 1986). This study will add to the body of knowledge concerning S.

odoratus by addressing these two crucial issues, as well provide additional data for future

comparisons.

Obj ectives of my study were to map the historical and current distribution of .%irl~liri,~ /i t

odoratus in the 4.7 km study section of the North Fork of the White River; identify habitat

variables positively and negatively associated with current.%e~linesdiciit s/ odoratus distribution in

NFWR; determine movement patterns of the NFWR .9t-neshe- tislr~ / odoratus population; assess

population structure and demographics of.rl .tinesdient s/ odoratus in NFWR.









CHAPTER 2
SPATIAL ECOLOGY AND DEMOGRAPHICS OF A POPULATION OF .Sicinestheins //
odoratus (TESTUDINES: KINOSTERNIDAE) INT AN OZARK STREAM

Introduction

The Stinkpot, .Sicinestheins // odoratus (Latreille), is a small aquatic turtle found statewide in

Missouri except in the northern and northwestern third of the state (Johnson, 2000). Periodic

surveys of the turtle assemblage of the North Fork of the White River, Ozark County, Missouri

from 1969 through 1980 (Nickerson, unpubl. data) and in 2004 (Pitt, 2005) revealed the presence

of a small but consistent number of S. odoratus. Pitt (2005) also found an increase in the likely

habitat of S odoratus within the research section. With this in mind, I conducted a mark-

recapture study of S odoratus in this same research section over the summers from 2005 through

2007, to compare current and historical distribution, determine microhabitat factors associated

with current S. odoratus distribution, analyze movement patterns, and to assess population

structure and demographics.

Materials and Methods

Study Site

The study area consists of a 4.7 km section of the North Fork of the White River (NFWR),

Ozark County, Missouri. This stretch of river includes fast-flowing riffles, deep, calm pools, and

long, straight, relatively shallow runs. Depths range from approximately 2.5 m (non-flood stage)

to less than 50 cm (Nickerson and Mays, 1973).

The substrate is comprised of extensive chert-gravel beds, exposed limestone, sandstone,

and dolomite bedrock, and numerous rocks and boulders, both submerged and emergent, ranging

in size from approximately dinner plate size to the size of small automobiles (Nickerson and

Mays, 1973). Patches of mud or sand can be found along the river banks where the current is

slow or the banks are eroding. Water clarity is generally high, although siltation has increased










significantly since construction of an upstream high-water bridge in 1995 (Missouri Department

of Conservation (MDC), 2001). Still pools and slow moving stretches collect sediments and silt

which can cloud the water for more than 20 minutes when disturbed (personal observation).

NFWR is primarily surrounded by oak-hickory forest with numerous shrubs and grasses

(see Nickerson & Mays, 1973 for species list). The upstream end of the research section

includes an extensively cleared private campground on the west bank and an MDC campsite and

boat ramp access point on the east bank (Pitt, 2005). Clearing within the study section also

occurs downstream at two canoe ranches/campgrounds, two private residences, and an MDC

campground at the end of the study section (Pitt, 2005). More than 100 aquatic plant species

occur in the NFWR drainage (Nickerson & Mays, 1973 for list) with the most common species

in the study section being Justicia amnericana, Water Willow; Ceratophyllum demersum,

Coontail; and M~yriophyllum heterophyllum and M. pinnatum, Water Milfoil. Additionally, there

are mosses and a species of long, fibrous algae from the genus Cladophora. The long, fibrous

algae become very abundant mid- to late-summer and cling to submerged and emergent

vegetation, rocks, and logs, forming extensive floating mats (Pitt, 2005). Other chelonian

species found within the research section are the Northern Map Turtle, Guraptemys geoguraphica,

the Red-Eared Slider, Trachemys scripta, the Spiny Softshell, Apalone spinifera, Common

Snapping Turtle, Chelydra serpentine, and the River Cooter, Pseudemys concinna.

I followed the same methodology set forth in initial turtle surveys by Nickerson and Mays

in 1969 (unpubl. data), Nickerson in 1980 (unpubl. data), and Pitt in 2004 (2005) who reassessed

the turtle community with a focus on G. geoguraphica. The 4.7 km section of river was divided

into fifty-one 92 m-long stations (stations 0-50; Figure 2-1). The research section was surveyed

every other day from mid-June to mid-August 2005 through 2007, between 0900 h and 1800 h,











































Figure 2-1. Stations 0-50 of the 4.7 km research section of the North Fork of the White River, Ozark County, Missouri. Whitewater
riffles are located in or between stations 2-3, 9-10, 17, 29-30, and 38-39.









weather permitting. Summer rainfall in this region occurs most often in the form of brief but

often intense thunderstorms (Nickerson and Mays, 1973). Nickerson and Mays (1973) reported

that in 1969 water clarity was difficult to disrupt, and they were able to work the day after a

heavy rainfall (e.g., 5 cm). On 1 July 2007, between 6 cm and 9 cm of rain fell at the study site

(National Climatic Data Center, 2008), reducing underwater visibility to less than 50 cm for 5

days (personal observation).

Capture Methods

Surveys were conducted via snorkeling and hand capturing, paired with a novel capture

technique (see below). Trapping with hoop or fyke nets is the common method of capturing S.

odoratus (Plummer, 1979), and use of hoop traps was attempted in this study. Traps were set

between 1800 h 0900 h several times and yielded only captures of non-target species (e.g.,

crayfish). Trapping proved to be unsuitable for use in this study. NFWR is a popular river for

recreation, receiving anywhere from 10 to > 1000 recreational users/day in the summer. Theft or

other disruption of traps, as well as removal of trapped turtles by recreational river users were

concerns. Additionally, trap-related mortality was a concern, since traps would need to be

placed where they would be fully submerged to avoid being seen by the public. Trap mortality

of turtles is not uncommon (Barko et al., 2004), with one researcher having 41 S. odoratus drown

in a trap within 24 hours (see Dodd, 1989).

In a pilot study, S. odoratus was primarily encountered basking atop mats of floating

emergent vegetation consisting ofhyriophyllum sp. and Ceratophyllum demersum which were

too dense to move through, and which would release a cloud of silt if touched, reducing visibility

to < 10 cm. Neither the silt substrate nor the vegetation mats were present in the research section

during historical surveys.









The novel capture method involved one researcher sitting in the rear of an eight foot, flat-

bottomed j ohn boat with the other researcher standing in the water holding the front of the boat.

The latter researcher would push the boat along the edge of the vegetation mats until stinkpots

were sighted. The researcher holding the boat would then quickly push the boat forward onto

and over the mat and the researcher in the boat would pick the turtles off the surface. The

researcher pushing the boat through the water walked with his head submerged while wearing a

facemask. The boat was pulled back upstream following the use of the novel technique and the

area was surveyed while snorkeling in order to stay consistent with the other survey method.

This method proved highly effective for both locating and capturing stinkpots in this study.

Marking Methods

Each captured turtle was numbered with nail polish (AM Cosmetics, Inc., North Arlington,

NJ). The polish marks allowed for visual identification from a distance if the turtle was able to

escape before recapture and minimized frequent, unnecessary recaptures and possibility of stress

that may come with handling. A clear topcoat was also applied to assure the polish would not

come off over the course of the season. Turtles that were large enough to be sexed in 2006 and

2007 were inj ected with a passive integrated transponder (PIT) tag (Destron-Fearing

Corporation, So. St. Paul, MN) to identify recaptures in subsequent years. Tags were injected

into the anterior inguinal region next to the bridge of the carapace following the protocol of

Buhlmann and Tuberville (1998). The injection site was prepped by cleaning with 70%

isopropyl alcohol. The hole was covered with New-Skin liquid bandage (Medtech Laboratories

Inc., Jackson, WY).

Data Collection

Each turtle was weighed to the nearest 0. 1 g using an Ohaus Scout Pro digital scale. I

measured straight-line carapace length (SCL) to the nearest millimeter using calipers. I measured










plastron length (PL) to the nearest millimeter using a tape measure. Turtles were sexed visually

when possible based on secondary sexual characteristics. Mature male Stinkpots have longer

and thicker tails than females, as well as scale patches on the inner hind legs (Ernst et al., 1994).

Water depth, substrate type and depth, behavior, and distance from shore were recorded at the

site of each capture, as well as GPS coordinates or station number.

Data Analysis

GPS coordinates taken at capture sites were entered into ArcGIS 9.2 (ESRI). Distance

between captures was determined using the "measure" tool. A geo-referenced map of the fifty-

one 92 m study sections was created by tracing over a USGS 7.5 minute quadrangle

topographical map in ArcGIS using the "edit" tool. Turtles from the historical and current data

were placed in the stations in which they were captured. Chi-square (X2) gOodness-of-fit tests

were performed to determine if sex ratios differed from the expected value of 1:1. I performed

Mann-Whitney Rank Sum tests to determine if there were any significant differences in CL, PL,

or weight between sexes. Statistical analyses were performed at the a = 0.05 level using

SigmaStat 2.0 (Jandel Corporation 1995). Small sample sizes in 1969, 1970, 1980, and 2004

prevented more rigorous statistical comparisons.

Results

Distribution

Distribution of S. odoratus in the research section from 1969 through 2007, displayed as

number of S odoratus captured per station, is shown in Figures 2-2A-G. Distribution was

similar for all years, with the exception of S odoratus present at the downstream end (station 50)

of the research section in 2007. No S~illinotheins/ odoratus were encountered in any year in

stations 3, 5-15, 17, or 30-49.
























S. odoraltus By Staltion



















Source Data: Scanned USGS
1:24,000 scale 7.5 minute
qua drang le topo grap hic ma p


0137.5 275 550 825 1 ,100




Figure 2-2. Distribution of.rl Sillinotheiits odoratus within the 4.7 km study section of the North Fork of the White River, Ozark
County, Missouri. A) 1969. B) 1970. C) 1980. D) 2004. E) 2005. F) 2006. G) 2007.

















































0 137.6 275 550 825 1,100


S. odoratus By Station )'L




m 2 --~rr ne ,

IIB


Figure 2-2. Continued



























































550 826 1,100


0 137.5 276


S. odoratus By Station









5









Source Data: Scanned USGS
1:24,000 scale 7.5 rninute
quadrangle topographic map


Figure 2-2. Continued















































Source Data: Scanned USGS
1:24,000 scale 7.5 minute
quadrangle topographic map


I I I Meters


S. odoratus ELy Staltion


137 5 275


825 1,100


Figure 2-2. Continued
























S. odoratus By Station




1 1-2




11 -16








Source Data: Scanned USGS
1:24,000 scale7T.5 rninute
quadrangle topographic rnap


I I I Meters


0 137 5 275


550 825 1,100


Figure 2-2. Continued























S. odoratus By Station




1 1-2


7-10

11 -16










Source Data: Scanned USGS
1:24,000 scale 7.5 minute
quadrangle topographic map


I I I Meters


O 137.5 275


550 825 1,100


Figure 2-2. Continued










N



S. s atuBy Staion II~







quadrangl to gapi map --

O 3 256082 ,0


Figure 2-2. Continued































Number of Observations


Behavior 2004-2007 1969-1970
Basking on vegetation or algal mat 133 2
Basking on log 6 13
Swimming 2 1
On bottom 7 12
Total 148 28

Movement

Most recaptured turtles were found either within their original capture station or the

adjacent station (Table 2-2). The mean distance traveled by recaptured females was 102 m

(n=17) and by males 198 m (n=10). One female had no measurable change in movement after

348 days. The longest movement recorded was by a male that traveled downstream

approximately 780 m in 401 days. This included a movement of 440 m in 49 days. The longest

movement by a female was 525 m downstream in 380 days. No turtle was captured on the

opposite side of the river from the side where it was originally captured. Most movement was in

the direction of upstream to downstream.


_Ij


Habitat

Mean water depths at capture sites from 2005 through 2007 were 74.8 cm, 77.9 cm, 62.4

cm, and 76.0 cm for males, females, juveniles, and all pooled, respectively. From 1969 through

1980 .Sicinesthesit s/ odoratus were most often observed basking on logs (13/28 observations) or

walking on the river bottom (12/28 observations). From 2004 through 2007, S. odoratus were

most often observed basking on emergent vegetation or algal mats (Table 2-1). .Sicinesthes,~ its

odoratus was found on all substrate types, but was only found over sand, rock, or gravel if there

was also dense vegetation present.

Table 2-1. Behavior of.rl Silinesthes its odoratus at site of capture, North Fork of the White River,
Ozark County, Missouri.


















































Population Structure and Demographics

A total of 112 different turtles was captured from 2004 through 2007 with 28 turtles

recaptured a total of 35 times. Mean CL, PL and weight from 2005-2007 are reported in Table

2-3. There were no statistically significant differences between male and female carapace

lengths or weight in any year (Table 2-4, 2-5). There were statistically significant differences in

male versus female plastron length in 2007 and for the years pooled (Table 2-6). Sex ratios did


Table 2-2. Distance traveled by recaptured .icilthe tisrrll / odoratus from the North Fork of the
White River, Ozark County, Missouri, 2004-2007.


Days Between Capture
34
25
25
7
15
1
6
18
356
328
19
1
352
49
720
1
6
26
772
15
333
348
731
374
380
7
12
31
368
408


Turtle ID
1
2
3
4
5
6
7
8
9
10
11
11
12
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28


Sex
Female
Female
Female
Male
Female
Female
Male
Male
Male
Female
Female
Female
Male
Male
Female
Unknown
Female
Male
Female
Female
Male
Female
Female
Female
Female
Female
Female
Male
Male
Male


Station
50
29
29
24
24
23
23
2328
2223
2223
22
22
1924
2429
1824
0.5
24
23
2321
27 24
20.5 23.5
23
23 -21 .75
2322
2428
24
2423
2422
24
0.5


Distance Traveled (m)
10
<5
40
6
14
16
10
530
72
80
10

340
440
470


70
100
300
270

100
50
525

100
180
45
20
















Table 2-3. Mean carapace length (CL), plastron length (PL) and weight (WT) of .9t-inesthe-iti s/
odoratus in the North Fork of the White River, Ozark County, Missouri, 2005 2007.
Year Sex Mean CL (mm) Mean PL (mm) Mean WT (g)
2005 M 97 71 162.4
F 89 69 139.3
2006 M 103 70 180.5
F 97 75 164.2
2007 M 91 62 131.3
F 95 74 172.2
All Years M 96 67 155.2
F 94 73 159.5



Table 2-4. Test for difference in male versus female carapace length of.rl .tinesdlreiits odoratus in
the North Fork of the White River, Ozark County, Missouri, 2005-2007.
Year Statistic P
2005 T =201.0 0.312
2006 T = 262.0 0.597
2007 T =181.5 0.569
All Years T =1454.0 0.860



Table 2-5. Test for difference in male versus female weight of .9t-inesdlr, eiits odoratus in the
North Fork of the White River, Ozark County, Missouri, 2005-2007.
Year Statistic P
2005 T = 200.0 0.332
2006 T = 258.0 0.928
2007 T =153.0 0.107
All Years T =1392.0 0.685



Table 2-6. Test for difference in male versus female plastron length of.rl .%inesdiciit s odoratus in
the North Fork of the White River, Ozark County, Missouri, 2005-2007.


0.804
0.131
0.034*
0.035*


not differ from the expected ratio of 1:1 (Table 2-7). Size class distributions by carapace length


for each year are shown in Figures 2-3A-G.


Year Statistic
2005 T =182.5
2006 T = 207.0
2007 T =142.0
All Years T =1253.5
* Denotes significance at the a=0.05 level.










Table 2-7. Sex ratios and chi-square statistics for .9e~lineithein s/ odoratus, North Fork of the
White River, Ozark County, Missouri, 2005-2007.
Year Ratio males:females Chi-square P-value (2-tailed)
2005 1:1.82 2.065 0.151
2006 1:1.27 0.265 0.607
2007 1:1.38 0.516 0.472
All Years 1:1.43 2.306 0.129
Yates' correction factor for continuity was applied to all cases to compensate for small sample
sizes.


6
5
u- 4


O Males
inFemales
ci Juveniles


0-39 40-49 50-59 60-69 70-79 80-89
Size Class (mm)


90-99 100-109 110-119


8120


Figure 2-3. Size class distribution by carapace length for.rl .%imothe-iu s/ odoratus captured in the
4.7 km research section of the North Fork of the White River, Ozark county Missouri.
A) 1969. B) 1970. C) 1980. D) 2004. E) 2005. F) 2006. G) 2007.


nnn
























6

5

L4


O Males
mFemales
5 JuvenilesB


0-39 40-49 50-59 60-69 70-79 80-89 90-99 100-109 110-119 81 20

Size Class (mm)





10-











5 -

u- 4-










0-39 40-49 50-59 60-69 70-79 80-89 90-99 100-109 110-119 81 20

Size Class (mm)



Figure 2-3. Continued


O Males
a Females
a Juveniles
























O Males
m Females D
5 Juveniles


5

u- 4









0-39 40-49 50-59 60-69 70-79 80-89 90-99 100-109 110-119 81 20
Size Class (mm)



Figure 2-3. Continued


O Males
m Females E
5 Juveniles


6

5

u- 4

3

2





0-39 40-49 50-59 60-69 70-79 80-89 90-99 100-109 110-119 81 20
Size Class (mm)
















9

8

7

6
c*
5

u- 4


O Males
m Females F
5 Juveniles
































O Males
females

m Juveniles G


0-39 40-49 50-59 60-69 70-79 80-89 90-99 100-109 110-119 81 20

Size Class (mm)


Figure 2-3. Continued


5

L4









0-39 40-49 50-59 60-69 70-79 80-89 90-99 100-109 110-119 81 20

Size Class (mm)









Discussion


Distribution and Habitat

The distribution of S. odoratus in the research section has remained virtually unchanged

since 1969, with the primary difference being the presence of the species in the farthest

downstream station (station 50) of the section in 2007. .Sillinotheiits odoratus is found in three

distinct areas: the west stream margins of stations 0-3 and 50, and the east stream margin from

station 18-29. These three areas are separated by long stretches of river with gravel and bedrock

substrates as well as by whitewater riffles, which both probably serve as barriers to movement of

S. odoratus. No individual was observed to move between these areas in either a single season

or between years. This observation, together with the presence of potential barriers to

movement, indicates that there are probably three subpopulations of S. odoratus within the

research section. In addition to being the only areas of the research section that contain thick

muddy substrates, they are the only areas that also contain dense stands of rooted underwater

vegetation. These stands were considerably smaller or absent in 1969-1970, 1980 (Nickerson,

personal comm.), and 2004 (personal observation) than from 2005-2007. There are several

likely explanations for the observed increase in vegetation. A flood event occurred just prior to

the field season in 2004 that registered a daily mean discharge of over 15,000 cubic feet per

second (cfs) (United States Geological Survey (USGS), 2008). This flood may have scoured

submerged vegetation from the stream bed as well as displaced turtles downstream, both of

which returned to the research section in 2005 in the absence of any subsequent flood events.

Nutrient loading is another likely cause of the increase in both aquatic vegetation and earlier and

larger seasonal blooms of long fibrous algae. The research section of NFWR flows through a

narrow valley, and atop bluffs on both sides of the river are cow pastures. In addition, much

residential development has occurred on the river upstream of the research section, and NFWR is










heavily used in the summer for canoeing, kayaking, and tubing. All of these activities could be

adding significant levels of nitrogen and phosphorus to the river, resulting in increased growth of

aquatic vegetation. Water quality testing upstream of, in, and downstream of the research

section in summer 2007 revealed levels of total and fecal coliform bacteria that exceed those

recommended for full body contact (Pitt, personal comm.). These high coliform counts indicate

that human or animal waste is entering the river, and most likely causing an increase in nutrient

levels. This could explain the increase in aquatic vegetation. More vegetation, and thus

potential habitat, is probably why more S. odoratus were found in 2005-2007 than in historical

surveys.

While Ernst et al. (1994) report finding an individual in a gravel-bottomed stream in

Arkansas, no S. odoratus were observed in open water on rock or gravel substrates in NFWR.

Several individuals were captured on emergent mats ofhyriophyllum sp. and Ceratophyllum

demersum that were rooted in and growing over rock and gravel. These areas were in close

proximity to (~10 m) areas with mud substrate. These data indicate that presence of rooted

vegetation, as opposed to soft-bottom substrates, are more important to the distribution of S

odoratus in NFWR. Taking into account the preferred microhabitat of S odoratus from this

study as well as that reported in the literature (i.e., soft bottom, emergent vegetation) and the area

of that habitat in the research section, I estimate that S. odoratus could inhabit approximately 1.5

hectares, or 6% of the total area of the research section. This figure is arrived at by multiplying

the approximate distance vegetation mats extend out from the banks (11 m) times the length of

the number of stations occupied by vegetation mats in 2007 (1380 m) It is possible that S.

odoratus utilizes the rock and gravel-bottomed sections of river not included in this estimate,

particularly during nighttime hours when surveys were not conducted, but they were not









observed doing so in this or previous studies of NFWR. Future studies of the NFWR should

include nocturnal sampling to determine if S odoratus uses these widely available habitats.

In contrast to Carr (1952) and Ernst et al. (1994) who reported that aerial basking (basking

on objects out of the water) was uncommon in S. odoratus, Nickerson (2000) found that North

Fork individuals in 1969 and 1970 were most often observed basking aerially on logs or trees (15

observations), while aquatic basking was observed only three times. From 2004 to 2007, there

were only six observations of aerial basking, while observations of partially submerged basking

on mats of aquatic vegetation totaled 133. Over the period of Nickerson's sampling, sycamores,

willows, and other large trees were observed at the stream banks extending out over the river

providing ample basking sites for turtles, as well as refugia in the exposed underwater root

masses. These conditions no longer exist in the research section. Between surveys in 1980 and

2004, several massive floods occurred in the NFWR. Flooding in February and November of

1985 registered daily mean discharge of 32,000 ofs and 45,000 ofs, respectively (peak flow:

108,000 ofs and 133,000 ofs, respectively) at the downstream USGS monitoring station (USGS,

2008). Floods in September and November of 1993 registered daily mean discharge of 25,100

cubic feet per second (cfs) and 32,600 ofs, respectively (peak flow: 52,500 ofs and 71,200 ofs,

respectively) (USGS, 2008). The non-flood discharge of NFWR is less than 2500 ofs (Nickerson

et al., 2007). Flooding in NFWR between 3700 ofs and 5100 ofs in 1970 is known to have

removed a massive sycamore tree from the research section (Nickerson et al., 2007). The floods

of 1985 and 1993 likely washed out many more trees and may have led to the observed change in

S. odoratus basking behavior. In addition, Pitt (2005) noted the general lack of basking sites in

2004 compared to historical studies. G. geoguraphica, by far the most abundant chelonian species

in the research section, may be out-competing S. odoratus for a limited number of woody or










rocky basking sites. Basking plays an important thermoregulatory role in digestion and egg

development, as well as aids in parasite removal. Boyer (1965) found that turtles basking on

partially submerged substrates (such as vegetation or algal mats) were able to raise body

temperature above that of the air or the water. .Slillinotheins/ odoratus in the NFWR may be able

to thermoregulate effectively in this way, but apparently are unable to receive the benefit of

parasite removal, demonstrated by the prevalence of leech infestation on captured turtles.

Movement

Movement of S odoratus observed in this study is similar to patterns observed in other

systems. Mahmoud (1969) found that the average distance traveled between captures was 66 m

and 38 m for males and females, respectively, in less than 100 days, and 69 m and 51 m for

males and females, respectively, in greater than 100 days. Ernst(1986) reported average

movements of 117 m and 90 m for males and females, respectively. The average distance

traveled between captures for males and females in this study, 198 m and 102 m, respectively,

follow the same pattern of longer distances traveled by males. Iverson and Meshaka (2006),

analyzing data from several studies, speculated that local conditions, rather than geography,

dictate movement of S odoratus. This seems to be the case in NFWR. Both males and females

were recorded traveling distances greater than 500 m, as well as not traveling any measurable

distance. All recorded movements occurred within and may have been confined to the preferred

microhabitat: submerged patches of rooted vegetation. A small-scale experiment intended as a

pilot study was conducted during this survey to examine barriers to movement in S. odoratus.

Four adults were displaced to the opposite bank of the river from where they were initially

captured, a distance of approximately 50 m. Between the initial capture site and release site was

a 1.5 m 2.5 m deep channel with a gravel/bedrock substrate. To find the preferred habitat,

turtles would need to cross the river or travel approximately 300 m downstream on the same side









of the river. Displacement occurred in August 2006, and the turtles were not recaptured in 2007.

Both the initial capture sites and the release sites, including preferred and non-preferred

microhabitat, were surveyed. While definitive results were not obtained from this side

experiment, similar experiments could be done in rivers like the North Fork using radio

telemetry to track the released turtles to answer this question.

Population Structure and Demographics

Small sample sizes combined with few recaptures during each sampling period prevented

the determination of any reliable population estimates for S. odoratus in the research section.

The sex ratios for 2005-2007 were all female-biased, but none were significantly different from

the expected ratio of 1:1 (Table 2-7). Ford (1999) also reported a slightly female-biased

population in Missouri, and female-biased populations have also been reported for Michigan

(Risley, 1933), Illinois, (Tinkle, 1961), Alabama (Dodd, 1989, Prassak et al., 1992), Louisiana

(Rayburn et al., 1989), and Florida (Meshaka, 1988, Aresco, 2005). Risely's (1933) high

proportion of females in the population was attributed to sampling bias (Gibbons, 1970 in Bury,

1979). Likewise Dodd (1989) stated that sampling over a wider area or over different portions of

the reproductive season may have yielded a different sex ratio in Alabama. Females in the

NFWR may have been captured more often than males because of increased activity related to

nesting in the early-to-mid summer when sampling primarily occurred. Males would likely be

encountered more often in the fall due to their search for mates.

An examination of size class based on CL reveals that all size classes are fairly well

represented and evenly distributed. The exceptions to this are the hatchling size class (under 40

mm CL) in all years and the distribution for the year 1980. Hatchlings were likely missed in

sampling due to their small size and cryptic nature. Hatchlings are most likely inhabiting the

shallowest areas of the river that are not amenable to sampling via snorkeling. In addition, eggs









in Missouri are laid from June through August, and take two to three months to hatch (Johnson,

2000). Eggs may not have hatched during the field season that may explain the absence of

hatchlings in samples. The 1980 data stand out in that only the largest size classes are

represented (over 90 mm CL). While sampling for all other years occurred in the early to mid-

summer, 1980 sampling occurred late-summer to fall. Larger size classes may have been more

conspicuous during this time of year due to movements related to the fall mating season.

Juveniles were captured in every year except 1980, suggesting that there is recruitment. The

most striking detail that comes from the size class distribution graphs is the large increase in the

number of turtles captured between 2004 and 2005-2007. As discussed above, 2005 saw an

increase in the area of mats of emergent vegetation, as well as an increase in the amount of time

the mats were present. In 2004, mats of vegetation or algae did not appear until early August,

whereas from 2005-2007, these mats were present from the start of the field season in early June.

The newly available habitat may have drawn S. odoratus from other parts of the river. In

addition, drought conditions in the area may have caused S. odoratus to migrate into the study

section from nearby cow ponds, creeks or backwaters. It is also possible that in 2004, more S.

odoratus were present in the research section but highly cryptic, spending daytime hours buried

in mud or hiding in muskrat or otter dens when the protection of dense submerged vegetation

was not available. When the vegetation was present from 2005-2007, these turtles may have

become more apparent by basking and feeding in the relative safety of the vegetation mats.

It was not surprising that dimorphism was not found between male and female carapace

length, as dimorphism has been rarely reported in the species. The significant difference in

plastron length may confer fitness benefits to males with small plastrons. Ford (1999) also found

female S. odoratus in Missouri with larger plastron lengths than males, while the carapace










lengths were similar. The biological significance of this, cited by Ford in Gibbons and Lovich

(1990), is that males with smaller plastrons may have a copulatory advantage through greater

flexibility. Since greater plastron lengths in females were not observed in each year, further

investigation of this phenomenon in NFWR is warranted.

Male S. odoratus in the NFWR reach sexual maturity when CL ranges between 61 mm and

70 mm. The smallest male that could be identified based on secondary sexual characteristics was

70 mm. This individual was initially captured in 2006 with a CL of 61 mm and sex could not be

determined. It was recaptured 408 days later and was clearly a male based on secondary sex

characteristics. Its age could not be determined accurately because of obscured plastral annuli,

but the size at maturity is similar to that found by Mahmoud (1969) for males in Oklahoma (65

mm CL). Females in the NFWR were sexable between 57 mm CL and 65 mm CL. Most studies

have found that female S. odoratus mature at larger sizes and greater ages than males (Risley,

1933; Tinkle, 1961; Mitchell, 1988; Edmonds, 1998). The Eindings on size at maturity in this

study could be an artifact of small sample size, so future studies should involve some histological

examination of specimens to more accurately determine the age and size at maturity of S

odoratus in the NFWR.

Conclusions

It is clear that while there currently are more S. odoratus within the 4.6 km research section

than were found in historical studies, the number is still small, even without accurate population

estimates. The increase in number of turtles captured in recent years corresponds to the increase

in available habitat. While significant habitat alteration has occurred in and around the research

section in the form of development, land clearing, siltation, and basking site removal,

distribution of S odoratus within the research section has changed little in the last 38 years, with

the species confined to three distinct areas.









This study provides insight into the effects of land use change and habitat alteration on

aquatic ecosystems. The NFWR has changed from a relatively pristine, remote river where few

people were encountered during sampling periods in 1969 and 1970, to a heavily used recreation

destination in 2007. Continued habitat alteration, by affecting the behavior and abundance of

species, may have consequences not only for S. odoratus but for the entire community. These

data can inform future studies by providing a long-term perspective on a turtle species, as well as

baseline data against which future surveys can be compared.









CHAPTER 3
CONCLUSIONS

Pitt (2005) and Nickerson et al. (1984) documented dramatic changes in the NFWR

watershed between 1969 and 2004, including large-scale land and riparian zone clearing,

development, the removal of basking sites, nutrient loading, and increased recreational use.

These factors appear to be benefiting the .Sillinotheins/ odoratus populations of the North Fork

through an increase in mud, sedimentation, and growth of submerged aquatic vegetation.

Many other ecologically generalized chelonians have been shown not only to tolerate but

flourish in polluted or degraded habitats. Moll's (1977) study on the polluted Illinois River

revealed that Common Snapping Turtles, Spiny Softshells, False Map Turtles, and sliders were

thriving while Yellow Mud Turtles, Blanding's Turtles, and Smooth Softshells were greatly

reduced or absent. Likewise, Moll and Moll (2004) note that several species, such as the

Malayan Box Turtle, and the Black Pond Turtle have been more successful in impacted habitats

than in more natural habitats (Moll and Khan bin Momin Khan, 1990 in Moll and Moll, 2004).

Presumably, if generalists tend to benefit from degraded conditions, specialists likely

would not. Pitt (2005) documented a decline in G. geoguraphica, a mollusk specialist, within the

NFWR research section between 1969 and 1980, as well as a failure to rebound from the 1980

level by 2004. She attributed as least part of this failure to degraded habitat conditions. In

addition, the generalist Trachemys script elegan2s became established in the research section

over this time period, and Pitt (2005) determined that its distribution had expanded between 1980

and 2004.

My study should not prove to be the end of research in NFWR, as additional questions

arose over the course of my research. While I did not encounter S. odoratus on rock or gravel

substrates in open water, I suggest that future studies in NFWR vary sampling both seasonally










and temporally (nocturnal and diurnal) to determine variation in habitat use. Radio telemetry

could also be employed to determine nocturnal habitat use. I suggest that the phenomenon of

NFWR S. odoratus males having smaller PL's than females should be confirmed, as this

phenomenon was not found to occur in 2005 or 2006. Likewise, I suggest that the age and size

at maturity of NFWR S. odoratus should be confirmed, as findings in my study were based on

just a few individuals. This could be accomplished by x-raying individuals in the lab for

presence of ova or follicles, or by harvesting some individuals for histological examination.

Finally, while my study has shown that the increase in S. odoratus abundance is correlated with

an increase in rooted underwater vegetation, I suggest that the causative factors contributing to

the increase in S. odoratus abundance be examined. Manipulative experiments involving

vegetation and substrate, as well as examination of reproductive rate, recruitment, and migration

from outside the study section could point to the factors causing the increase in S. odoratus

abundance.

My study provides further evidence of how generalist species can increase in abundance

when habitats become degraded. Studies such as this, while important in a broad ecological

context, will also add to the knowledge of Urban Ecology, a growing Hield with increasing

importance as growing human populations expand into less populated, relatively undisturbed

areas.










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BIOGRAPHICAL SKETCH

Joseph J. Tavano was born in 1978 in Hyannis, Massachusetts. He grew up in

the town of Harwich, Massachusettes, on Cape Cod, where he graduated from Harwich High

School in 1996. He earned a B.A. in biology from the University of Vermont in 2005. He

began graduate school at the University of Florida in 2006, working toward an M. S. in

interdisciplinary ecology. Upon graduating, Mr. Tavano plans to pursue a career in teaching or

as a Wildlife Biologist.





PAGE 1

1 SPATIAL ECOLOGY AND DEMOGRAP HICS OF A POPULATION OF Sternotherus odoratus (TESTUDINES: KINOSTERNIDAE) IN AN OZARK STREAM By JOSEPH J. TAVANO A THESIS PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLOR IDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE UNIVERSITY OF FLORIDA 2008

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2 2008 Joseph J. Tavano

PAGE 3

3 To Amber, who always knew I could do it.

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4 ACKNOWLEDGMENTS I would like to thank Drs. Max Nickerson, Mike Moulton, and Kent Vliet for their guidance, advice and recommendations in the wri ting of this thesis. I would like to thank Gustavo Vasques for assistance with the GIS, and Dr. Kenney Krysko for help with the statistical analyses. I thank all of the wonderful peopl e I met while in the field in Missouri: Connie Morgan and Randy Padgett for their friendship an d hospitality; Amy and Justin Spencer at Sunburst Canoe Ranch for providing camping acco mmodations; Jeff Ettling, Eric Miller, Mark Wanner, and everyone else at the Saint Louis Zo o for all of their help and support through the years; and Karen and Ron Goellner, two wonderfu l people and great friends without whom this research would not have been possible. I woul d like to thank the Sain t Louis Zoological Park and the Reptile and Amphibian Conservation Corp s for providing the funding for this research. I thank my parents, my grandmother, my brothers and their families for all of their patience, encouragement and support over the year s. Finally, I would like to thank Amber Pitt for her invaluable help both in the field and in reviewing this thesis.

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5 TABLE OF CONTENTS page ACKNOWLEDGMENTS...............................................................................................................4LIST OF TABLES................................................................................................................. ..........6LIST OF FIGURES.........................................................................................................................7ABSTRACT.....................................................................................................................................8 CHAPTER 1 INTRODUCTION....................................................................................................................9The Stinkpot, Sternotherus odoratus (Latreille)..................................................................... 10Sternotherus odoratus in Missouri......................................................................................... 14Justification and Objectives....................................................................................................162 SPATIAL ECOLOGY AND DEMOGRAP HICS OF A POPULATION OF STERNOTHERUS ODORATUS (TESTUDINES: KINOSTERNIDAE) IN AN OZARK STREAM................................................................................................................. 17Introduction................................................................................................................... ..........17Materials and Methods...........................................................................................................17Study Site..................................................................................................................... ....17Capture Methods.............................................................................................................20Marking Methods............................................................................................................21Data Collection................................................................................................................21Data Analysis...................................................................................................................22Results.....................................................................................................................................22Distribution................................................................................................................... ...22Habitat.............................................................................................................................30Movement........................................................................................................................30Population Structure and Demographics......................................................................... 31Discussion...............................................................................................................................37Distribution and Habitat.................................................................................................. 37Movement........................................................................................................................40Population Structure and Demographics......................................................................... 41Conclusions.............................................................................................................................433 CONCLUSIONS.................................................................................................................... 45LIST OF REFERENCES...............................................................................................................47BIOGRAPHICAL SKETCH.........................................................................................................52

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6 LIST OF TABLES Table page 2-1 Behavior of Sternotherus odoratus at site of capture, North Fork of the Wh ite River, Ozark County, Missouri..................................................................................................... 302-2 Distance traveled by recaptured Sternotherus odoratus from the North Fork of the White River, Ozark County, Missouri, 2004-2007............................................................312-3 Mean carapace length (CL), plastron le ngth (PL) and weight (WT) of Sternotherus odoratus in the North Fork of the White River, Ozark County, Missouri, 2005 2007....322-4 Test for difference in male versus female carapace length of Sternotherus odoratus in the North Fork of the White Ri ver, Ozark County, Missouri, 2005-2007..................... 322-5 Test for difference in male versus female weight of Sternotherus odoratus in the North Fork of the White River, Ozark County, Missouri, 2005-2007...............................322-6 Test for difference in male ve rsus female plastron length of Sternotherus odoratus in the North Fork of the White River, Ozark County, Missouri, 2005-2007.........................322-7 Sex ratios and chi-square statistics fo r Sternotherus odoratus, North Fork of the White River, Ozark County, Missouri, 2005-2007............................................................33

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7 LIST OF FIGURES Figure page 1-1 Distribution of the Stinkpot, Sternotherus odoratus..........................................................101-2 Distribution of Sternotherus odoratus in Missouri............................................................152-1 Stations 0-50 of the 4.7 km research sect ion of the North Fork of the White River, Ozark County, Missouri.....................................................................................................192-2 Distribution of Sternotheru s odoratus within the 4.7 km study section of the North Fork of the White River, Ozark County, Missouri............................................................232-3 Size class distribution by carapace length for Sternotherus odoratus captured in the 4.7 km research section of the North Fork of the White River, Ozark county Missouri.............................................................................................................................33

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8 Abstract of Thesis Presen ted to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Master of Science SPATIAL ECOLOGY AND DEMOGRAP HICS OF A POPULATION OF Sternotherus odoratus (TESTUDINES: KINOSTERNIDAE) IN AN OZARK STREAM By Joseph J. Tavano August 2008 Chair: Max A. Nickerson Major: Interdisciplinary Ecology From 1969 through 2004, the turtle community of a 4.7 km section of the North Fork of the White River, Ozark County, Missouri wa s studied periodically. The Stinkpot, Sternotherus odoratus was found in all years in low abundance. From 2005 through 2007, I continued this survey to determine the distribution and rela ted parameters, movement patterns, population structure, and demogr aphic parameters of S. odoratus within the re search section. Distribution was similar from 1969 through 2006, with expansion to the terminal station of the research section in 2007. Tu rtle distribution was positively correlated with dense submerged patches of mud and rooted vegetation. Moveme nt between captures from 2004 to 2007 averaged 198 m for males and 102 m for females, with a ra nge of 0 m to 740 m. There was no difference between male and female carapace lengths or wei ght. There was a significant difference in male and female plastron lengths with females being larger Sex ratios were slightly female biased but not significantly so. My study provides insight into the effects of land use change and habitat alteration on aquatic ecosystems. Continued habitat alteratio n, by affecting the behavior and abundance of species, may have consequences not only for S. odoratus but for the entire community.

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9 CHAPTER 1 INTRODUCTION Turtles date back in the foss il record 210 million years to the late Triassic, pre-dating the dinosaurs (Gaffney, 1990). Tu rtles have evolved several unique adaptations, the most characteristic being the shell that encases the body. Extant turtles, or chelonians, are long-lived organisms, and may have the longest life spans of any extant vertebrate (Ernst et al., 1994). Giant tortoises have been known to live 120 to 15 0 years, and the oldest known box turtle lived 123 years (Pritchard, 1967). There are approximately 300 species of cheloni ans. Turtles can be found worldwide in terrestrial, aquatic, and marine habitats except at high latitudes or altitudes (Pough, 2004). Turtles, and river turtles in particular, can co mprise a significant portion of the biomass of ecosystems they inhabit (Congdon et al., 1986; Moll and Moll, 2004). They serve as predators, frugivores, seed dispersers, and scavengers, as we ll as prey for crocodilians and some birds and mammals (Moll, 1990; Ernst et al., 1994; De Lima et al., 1997; Ford & Moll, 2004). These roles act as important linkages both w ithin and between terrestrial a nd aquatic food webs (Moll and Moll, 2004). Turtles also play a vital role in nut rient cycling, maintenanc e of water quality, and energy flow (Moll and Moll, 2004). Moreover, turt le species with specia lized diet or habitat requirements can serve as indicators of habitat quality (E rnst et al., 1994). Despite their unique characteristics and th e important impacts they have on their ecosystems, turtles remain largely understudied. Many turtle populations are believed to be in decline (Moll, 1986; Ernst et al., 1994; Buhlman and Gibbons, 1997; van Dijk et al., 2000), yet not enough is known about the natural and life hi stories of many species, nor have there been sufficient long-term population studies to s upport that claim (Congdon et al., 1993; Dodd and Franz, 1993; Moll and Moll, 2004). Chelonians face many threats to survival including

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10 collection for the pet trade, utilization as a food resource, and ha bitat loss and alteration (Ernst et al., 1994; Buhlman and Gibbons, 1997; van Dijk et al., 2000; Moll and Moll, 2004). The Stinkpot, Sternotherus odoratus (Latreille) The Stinkpot, Sternotherus odoratus is a relatively sm all aquatic turtle found throughout North America from Maine to Florida, and west to Wisconsin and Texas (Ernst et al., 1994; Figure 1-1). Although it is consid ered to be a chiefly nocturnal species (Ernst, 1986), some studies have found peak activity periods to be b imodal. April to September activity periods in both Oklahoma and Pennsylvania have been reported between 0400 h to 1100 h and 1700 h to 2100 h (Mahmoud, 1969; Ernst, 1986). Peak activity for the rest of the year was between 1000 h and 1600 h (Mahmoud, 1969; Ernst, 1986). Figure 1-1. Distribution of the Stinkpot, Sternotherus odoratus. Reprinted from Ernst, C.H., J.E. Lovich, and R.W. Barbour. 1994. Turtles of the United States and Canada. Smithsonian Institution Press, Washi ngton D.C. and London. P. 142. Used with permission of the author.

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11 Sternotherus odoratus are omnivorous. Diet analyses reveal consumption of earthworms, leeches, clams, snails, crayfish, insects, minnows, tadpoles, algae, plants, seed, seed pods, fish eggs, and carrion (Lagler, 1943; Mahmoud, 1969; Er nst et al., 1994; Palmer and Braswell, 1995; Davis, 1996; Ford, 1999). Diets of Oklahoma Sti nkpots consisted primarily of insects, snails, and aquatic vegetation (Mahmoud, 1969) Snails also represented a si gnificant part of the diet of S. odoratus in Wisconsin (Vogt, 1981), Mi chigan (Lagler, 1943), and Florida (Bancroft et al., 1983). Sternotherus odoratus serve as prey for numerous species. Eggs are eaten by snakes ( Lampropeltis, Cemophora ) raccoons ( Procyon lotor), skunks ( Mephitis mephitis ), and herons (Ernst et al., 1994), and adults and ju veniles are preyed upon by raccoons, foxes ( Vulpes vulpes ), American Alligators ( Alligator mississippiensis ), Common Snapping Turtles (Chelydra serpentina), Bald Eagles ( Haliaeetus leucocephalus ), Cottonmouths ( Agkistrodon piscivorous ), water snakes (Nerodia sp.), Largemouth Bass ( Micropterus salmoides ), and wading birds (Clark, 1982; Delaney and Abercrombie, 1986; Pritchar d, 1989; Ernst et al., 1994; Mitchell, 1994; Palmer and Braswell, 1995). Although many species of freshw ater turtles exhibit pronounced sexual size dimorphism, sexual dimorphism in S. odoratus is not as pronounced. Tink le (1961) reported that sexual dimorphism in S. odoratus body size is only seen in southe rn populations. This observation is supported by Bancroft et al. (1983) and Meshaka (1988) who found that females in Florida have longer carapace lengths (CL) than males. In contrast, Edmonds and Brooks (1996) found males in a population in Ontario to have significantly longer carapace lengths than females. They hypothesized that this dimorphism was a result of selection for larger male body size in response to combat related competition for mates (E dmonds and Brooks, 1996). Aside from body size,

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12 tails of mature males are thicker and longer than that of females, and males develop elevated scale patches on the inner hind legs (Risley, 1930). Size at maturity varies with location and sex. In Michigan, Ri sley (1933) and Tinkle (1961) found that males reached maturity in th ree or four years at 60-70 mm carapace length (CL). Risley (1933) found that females mature d between ages nine and eleven at 80 mm CL, while Tinkle (1961) found mature females between tw o and seven years old, also at 80 mm CL. Mitchell (1988) in Virginia repo rted seeing male secondary sex ch aracteristics at two years and 51 mm CL, and females at 4 years and 66 mm CL. Mahmoud ( 1969) saw mature males at 65 mm CL between four and seven years of ag e and females matured between 65 mm and 85 mm CL at five to eight years of age in Oklahoma. Sex ratios for populations vary widely, but generally follow the pattern of being malebiased in northern populations and female-biase d in southern populations (Iverson and Meshaka, 2006). Mahmoud (1969) and Mitchell (1988) both found sex ratios not significantly different from 1:1 in Oklahoma and Virginia, respectively. An Ontario populat ion consisted of 72% males (Edmonds and Brooks, 1996), and Dodd (1989) reported a sex ratio of 1:2.8 males to females in Alabama. A range of explanations have been proposed to acco unt for the disparity in sex ratios, including differenti al mortality of nesting females (Edmonds and Brooks, 1996; Iverson and Meshaka, 2006), sampling bias as a result of trapping tech nique or behavioral differences (Dodd, 1989; Edmonds and Brooks 1996; Iverson and Meshaka, 2006), and temperature-dependent sex determination of ha tchlings (Edmonds and Brooks, 1996; Iverson and Meshaka, 2006). S ternotherus odoratus can be long-lived. Ernst (1986) observed two individuals in his field study, a male and a female, who were known to be 27 and 28 years old at the date of last

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13 capture, and a specimen at the Philadelphia Zoo li ved for 54 years, 9 months (Snider and Bowler, 1992). Sternotherus odoratus can be found in most any body of freshwater, including ponds and lakes, rivers and streams, ur ban lakes, reservoirs, and othe r man-made impoundments, although Iverson and Meshaka (2006) report a preference fo r permanent bodies of water with little or no current. While S. odoratus has been found on rocky and grav elly substrates (Mahmoud, 1969; Ernst et al., 1994; Palmer and Braswell, 1995), th is species is most commonly found in the softbottomed littoral zones of len tic systems with abundant subm erged vegetation (Kingsbury, 1993). Thermoregulatory basking in S. odoratus typically occurs while submerged in shallow water or floating at the surface, or while partiall y submerged atop lily pads or mats of algae or emergent vegetation (Ernst, 1986; Ernst et al., 1 994; Iverson and Meshaka, 2006). Basking also occurs on logs, stumps, or trees overhanging the water, with some individuals climbing as high as three meters above the waters surf ace (Vogt, 1981; Ernst, 1986; Nickerson, 2000). Several studies have examined movement of S. odoratus Most studies reveal a pattern of a larger home range in males than female s (Mahmoud, 1969; Ernst, 1986; Edmonds, 1998). Average distance traveled between captures by males and females ranges from 69 m and 51 m, respectively, in an Oklahoma stream (Mahm oud, 1969) to 2052 m and 828 m, respectively, in an Ontario lake (Edmonds, 1998). Most research on S. odoratus has been carried out in closed bodies of water such as lakes or reservoirs, with several ex ceptions. Ernst (1986) sampled a small creek in Pennsylvania together with marsh, floodplain pool, and mill pond habitats. Mahmoud (1969) studied S. odoratus in a small stream (average depth and widt h, approximately 1 m and 3 m, respectively)

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14 in Oklahoma, and Meylan et al. (1992), studied the turtle community of Rainbow Run in Marion County, Florida, with an emphasis on two kinoste rnid turtles, the Loggerhead Musk Turtle, Sternotherus minor and S. odoratus Sternotherus odoratus in Missouri Sternotherus odoratus is found statewide in Missouri except in the northern and northwestern third of the state (Johnson, 2000; Figure 1-2). The speci es is most commonly encountered in slow-m oving sections of larger rivers and streams in the Ozarks, the sloughs and backwaters of the Bootheel, and rivers in the northeast part of th e state (Johnson, 2000). Missouris smallest turtle species, S. odoratus typically range in size from 51 mm to 115 mm carapace length (Johnson, 2000), with the Mi ssouri record specimen measuring 117 mm carapace length (Powe ll et al., 1982). Research on S. odoratus in Missouri has been restricted mainly to two studies (Davis, 1996; Ford, 1999) that investigated reproduction, diet, and demogra phics. Davis (1996) studied the effects of cold water effluent on S. odoratus diet and reproduction in a reservoir in southwest Missouri. Ford (2000) gathered data on fora ging ecology and demographics of a population of S. odoratus in a lake created by the damming of a river, also in southwest Mi ssouri. Ford (1999) found that there was seasonal and se xual variation in the diet of S. odoratus in Missouri, with females eating more grass seeds, Heliosoma snails, crayfish, mayf lies, beetles, isopods, Ludwigia seeds, clams, and overall animal matter, and males consuming more nuts, water striders, and lepidopterans. Seed consumption in September was higher than for June-August, larger volume of vegetative material was consumed in July an d August compared to June, and more crayfish, snails, clams, and insects were consumed in June.

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15 Figure 1-2. Distribution of Sternotherus odoratus in Missouri (Johnson, 2000). Range map for Sternotherus odoratus reproduced from "The Amphibians and Rep tiles of Missouri." p. 182. Copyright 2000 by the Conservation Commission of the State of Missouri. Used with permission. From 1969 through 1980, M. A. Nickerson conducted a mark-recapture study of the turtle community in a 4.7 km section of the North Fork of the White River (NFWR), Ozark County, Missouri (Pitt, 2005). Pitts (2005) analyses revealed that S. odoratus was present in the research section, but in very low numbers. Pitt (2005) reassessed the turtle community of NFWR in 2004 with a focus on the Northern Map Turtle, Graptemys geographica historically the most abundant turtle in this commun ity (Nickerson, unpubl. data). Sh e found that whereas the river and surrounding terrestrial habita t had undergone significant changes since Nickersons surveys from 1969 through 1980, the number of S. odoratus captured was similar (Nickerson, unpub. data; Pitt, 2005).

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16 Justification and Objectives Although several ecological studies of S. odoratus have been conducted across its geographic range, little data exist on the life history of S. odoratus in one of its common habitatsthe river ecosystem. Likewise, there has been only one published long-term study of a S. odoratus population (Ernst 1986). T his study will add to the body of knowledge concerning S. odoratus by addressing these two cruc ial issues, as well provide additional data for future comparisons. Objectives of my study were to map the historical and curre nt distribution of Sternotherus odoratus in the 4.7 km study section of the North Fork of the White Rive r; identify habitat variables positively and negatively associated with current Sternotherus odoratus distribution in NFWR; determine movement patterns of the NFWR Sternotherus odoratus population; assess population structure and demographics of Sternotherus odoratus in NFWR.

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17 CHAPTER 2 SPATIAL ECOLOGY AND DEMOGRAP HICS OF A POPULATION OF Sternotherus odoratus (TESTUDINES: KINOSTERNIDAE) IN AN OZARK STREAM Introduction The Stinkpot, Sternotherus odoratus (Latreille), is a small aqua tic turtle f ound statewide in Missouri except in the northern and northwester n third of the state (Johnson, 2000). Periodic surveys of the turtle assemblage of the North Fork of the White River, Ozark County, Missouri from 1969 through 1980 (Nickerson, unpubl. data) and in 2004 (Pitt, 2005) revealed the presence of a small but consistent number of S. odoratus Pitt (2005) also found an increase in the likely habitat of S. odoratus within the research section. W ith this in mind, I conducted a markrecapture study of S. odoratus in this same research section over the summers from 2005 through 2007, to compare current and historical distribu tion, determine microhabita t factors associated with current S. odoratus distribution, analyze movement patterns, and to assess population structure and demographics. Materials and Methods Study Site The study area consists of a 4.7 km s ection of the North Fork of the White River (NFWR), Ozark County, Missouri. This stretch of river includes fast-flowing riffles, deep, calm pools, and long, straight, relatively shallow runs. Depths range from appr oximately 2.5 m (non-flood stage) to less than 50 cm (Nickerson and Mays, 1973). The substrate is comprised of extensive cher t-gravel beds, exposed limestone, sandstone, and dolomite bedrock, and numerous rocks and boul ders, both submerged and emergent, ranging in size from approximately dinner plate size to the size of small automobiles (Nickerson and Mays, 1973). Patches of mud or sand can be f ound along the river banks where the current is slow or the banks are eroding. Water clarity is generally high, although siltation has increased

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18 significantly since construction of an upstream high-water bridge in 1995 (Missouri Department of Conservation (MDC), 2001). Still pools and sl ow moving stretches collect sediments and silt which can cloud the water for more than 20 minutes when disturbed (personal observation). NFWR is primarily surrounded by oak-hickory forest with numerous shrubs and grasses (see Nickerson & Mays, 1973 for sp ecies list). The upstream e nd of the research section includes an extensively cleare d private campground on the west bank and an MDC campsite and boat ramp access point on the east bank (Pitt, 20 05). Clearing within the study section also occurs downstream at two canoe ranches/cam pgrounds, two private residences, and an MDC campground at the end of the st udy section (Pitt, 2005). More than 100 aquatic plant species occur in the NFWR drainage (Nickerson & Mays 1973 for list) with the most common species in the study section being Justicia americana, Water Willow; Ceratophyllum demersum, Coontail; and Myriophyllum heterophyllum and M. pinnatum Water Milfoil. Additionally, there are mosses and a species of l ong, fibrous algae from the genus Cladophora. The long, fibrous algae become very abundant midto late-summer and cling to submerged and emergent vegetation, rocks, and logs, forming extensive floating mats (Pitt, 2005). Other chelonian species found within the research section are the Northern Map Turtle, Graptemys geographica, the Red-Eared Slider, Trachemys scripta the Spiny Softshell, Apalone spinifera Common Snapping Turtle, Chelydra serpentina, and the River Cooter, Pseudemys concinna I followed the same methodology set forth in in itial turtle surveys by Nickerson and Mays in 1969 (unpubl. data), Nickerson in 1980 (unpubl. data), and Pitt in 2004 (2005) who reassessed the turtle community with a focus on G. geographica The 4.7 km section of river was divided into fifty-one 92 m-long stations (stations 0-50; Figure 2-1). The research section was surveyed every other day from mid-June to midAugust 2005 through 2007, between 0900 h and 1800 h,

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19 Figure 2-1. Stations 0-50 of the 4.7 km re search section of the North Fork of the White River, Ozark County, Missouri. Whitewater riffles are located in or between st ations 2-3, 9-10, 17, 29-30, and 38-39.

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20 weather permitting. Summer rainfall in this region occurs most often in the form of brief but often intense thundersto rms (Nickerson and Mays, 1973). Nick erson and Mays (1973) reported that in 1969 water clarity was diffi cult to disrupt, and they were able to work the day after a heavy rainfall (e.g., 5 cm). On 1 July 2007, between 6 cm and 9 cm of rain fell at the study site (National Climatic Data Center 2008), reducing underwat er visibility to less than 50 cm for 5 days (personal observation). Capture Methods Surveys were conducted via snorkeling and hand capturing, paired with a novel capture technique (see below). Trapping with hoop or f yke nets is the common me thod of capturing S. odoratus (Plummer, 1979), and use of hoop traps was attempted in this study. Traps were set between 1800 h 0900 h several times and yielded only captures of non-target species (e.g., crayfish). Trapping proved to be unsuitable fo r use in this study. NFWR is a popular river for recreation, receiving anywhere from 10 to 1000 recreational users/day in the summer. Theft or other disruption of traps, as we ll as removal of trapped turtles by recreational river users were concerns. Additionally, trap-related mortality was a concern, since traps would need to be placed where they would be fully submerged to avoid being seen by the public. Trap mortality of turtles is not uncommon (Barko et al., 2004), with one researcher having 41 S. odoratus drown in a trap within 24 hours (see Dodd, 1989). In a pilot study, S. odoratus was primarily encountered basking atop mats of floating emergent vegetation consisting of Myriophyllum sp. and Ceratophyllum demersum which were too dense to move through, and which would releas e a cloud of silt if touc hed, reducing visibility to < 10 cm. Neither the silt substrate nor the vege tation mats were present in the research section during historical surveys.

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21 The novel capture method involved one researcher si tting in the rear of an eight foot, flatbottomed john boat with the other re searcher standing in the water hol ding the front of the boat. The latter researcher would pus h the boat along the edge of the vegetation mats until stinkpots were sighted. The researcher holding the boat would then quickly push the boat forward onto and over the mat and the researcher in the boat would pick the turtles off the surface. The researcher pushing the boat through the water wa lked with his head submerged while wearing a facemask. The boat was pulled back upstream fo llowing the use of the novel technique and the area was surveyed while snorkeling in order to st ay consistent with th e other survey method. This method proved highly effective for both locating and capturing st inkpots in this study. Marking Methods Each captured turtle was numbered with nail polish (AM Cosm etics, Inc., North Arlington, NJ). The polish marks allowed for visual identifi cation from a distance if the turtle was able to escape before recapture and minimized frequent, unnecessary recaptures and possibility of stress that may come with handling. A clear topcoat wa s also applied to assure the polish would not come off over the course of the season. Turtles that were large enough to be sexed in 2006 and 2007 were injected with a passive integrat ed transponder (PIT) tag (Destron-Fearing Corporation, So. St. Paul, MN) to identify recapture s in subsequent years. Tags were injected into the anterior inguinal region next to the bridge of the carapace following the protocol of Buhlmann and Tuberville (1998). The injec tion site was prepped by cleaning with 70% isopropyl alcohol. The hole was covered with New-Skin liquid bandage (Medtech Laboratories Inc., Jackson, WY). Data Collection Each turtle was weighed to the nearest 0.1 g using an Ohaus Scout Pro digital scale. I measured straight-lin e carapace length (SCL) to th e nearest millimeter using calipers. I measured

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22 plastron length (PL) to the nearest millimeter usi ng a tape measure. Turtles were sexed visually when possible based on secondary sexual characte ristics. Mature male Stinkpots have longer and thicker tails than females, as well as scale patches on the inner hind le gs (Ernst et al., 1994). Water depth, substrate type and depth, behavior, and distance from shore were recorded at the site of each capture, as well as G PS coordinates or station number. Data Analysis GPS coordinates taken at captu re sites were entered into ArcGIS 9.2 (ESRI). Distance between captures was determ ined using the measur e tool. A geo-referenced map of the fiftyone 92 m study sections was created by tr acing over a USGS 7.5 minute quadrangle topographical map in ArcGIS using the edit tool. Turtles from the historical and current data were placed in the stations in whic h they were captured. Chi-square ( 2) goodness-of-fit tests were performed to determine if sex ratios differed from the expected value of 1:1. I performed Mann-Whitney Rank Sum tests to determine if there were any significant di fferences in CL, PL, or weight between sexes. Statisti cal analyses were performed at the = 0.05 level using SigmaStat 2.0 (Jandel Corporation 1995). Small sample sizes in 1969, 1970, 1980, and 2004 prevented more rigorous statistical comparisons. Results Distribution Distribution of S. odoratus in the research s ection from 1969 through 2007, displayed as num ber of S. odoratus captured per station, is shown in Figures 2-2A-G. Distribution was similar for all years, with the exception of S. odoratus present at the downs tream end (station 50) of the research section in 2007. No Sternotherus odoratus were encountered in any year in stations 3, 5-15, 17, or 30-49.

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23 A Figure 2-2. Distribution of Sternotherus odoratus within the 4.7 km study s ection of the North Fork of the White River, Ozark County, Missouri. A) 1969. B) 1970. C) 1980. D) 2004. E) 2005. F) 2006. G) 2007.

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24 B Figure 2-2. Continued

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25 C Figure 2-2. Continued

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26 D Figure 2-2. Continued

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27 E Figure 2-2. Continued

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28 F Figure 2-2. Continued

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29 G Figure 2-2. Continued

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30 Habitat Mean water depths at ca pture sites from 2005 through 2007 were 74.8 cm, 77.9 c m, 62.4 cm, and 76.0 cm for males, females, juveniles, and all pooled, respectively. From 1969 through 1980 Sternotherus odoratus were most often observed baskin g on logs (13/28 observations) or walking on the river bottom (12/28 obs ervations). From 2004 through 2007, S. odoratus were most often observed basking on emergent ve getation or algal mats (Table 2-1). Sternotherus odoratus was found on all substrate types, but was onl y found over sand, rock, or gravel if there was also dense vegetation present. Table 2-1. Behavior of Sternotherus odoratus at site of capture, North Fork of the White River, Ozark County, Missouri. Number of Observations Behavior 2004-2007 1969-1970 Basking on vegetation or algal mat 133 2 Basking on log 6 13 Swimming 2 1 On bottom 7 12 Total 148 28 Movement Most recaptured turtles were found either with in their original cap ture station or the adjacent station (Table 2 -2). The mean dist ance traveled by recaptured females was 102 m (n=17) and by males 198 m (n=10). One female had no measurable change in movement after 348 days. The longest movement recorded was by a male that traveled downstream approximately 780 m in 401 days. This included a movement of 440 m in 49 days. The longest movement by a female was 525 m downstream in 380 days. No turtle was captured on the opposite side of the river from the side where it was originally captured. Most movement was in the direction of upstream to downstream.

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31 Table 2-2. Distance tr aveled by recaptured Sternotherus odoratus from the North Fork of the White River, Ozark County, Missouri, 2004-2007. Turtle ID Sex Station Distance Tr aveled (m) Days Between Capture 1 Female 50 10 34 2 Female 29 <5 25 3 Female 29 40 25 4 Male 24 6 7 5 Female 24 14 15 6 Female 23 16 1 7 Male 23 10 6 8 Male 2328 530 18 9 Male 2223 72 356 10 Female 22 23 80 328 11 Female 22 10 19 11 Female 22 1 12 Male 1924 340 352 12 Male 2429 440 49 13 Female 18 24 470 720 14 Unknown 0.5 1 15 Female 24 6 16 Male 23 70 26 17 Female 23 21 100 772 18 Female 27 24 300 15 19 Male 20.523.5 270 333 20 Female 23 348 21 Female 23 21.75 100 731 22 Female 23 22 50 374 23 Female 24 28 525 380 24 Female 24 7 25 Female 24 23 100 12 26 Male 2422 180 31 27 Male 24 45 368 28 Male 0.5 20 408 Population Structure and Demographics A total of 112 different turtles was ca ptured from 2004 through 2007 with 28 turtles recaptured a total of 35 tim es. Mean CL, PL and weight from 2005-2007 are reported in Table 2-3. There were no statistically significant differences between male and female carapace lengths or weight in any year (T able 2-4, 2-5). There were statis tically significant differences in male versus female plastron length in 2007 and fo r the years pooled (Table 2-6). Sex ratios did

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32 not differ from the expected ratio of 1:1 (Table 2-7). Size class distributions by carapace length for each year are shown in Figures 2-3A-G. Table 2-3. Mean carapace length (CL), plastron length (PL) and weight (WT) of Sternotherus odoratus in the North Fork of the White Ri ver, Ozark County, Missouri, 2005 2007. Year Sex Mean CL (mm) Mean PL (mm) Mean WT (g) 2005 M F 97 89 71 69 162.4 139.3 2006 M F 103 97 70 75 180.5 164.2 2007 M F 91 95 62 74 131.3 172.2 All Years M F 96 94 67 73 155.2 159.5 Table 2-4. Test for difference in male versus female carapace length of Sternotherus odoratus in the North Fork of the White Rive r, Ozark County, Missouri, 2005-2007. Year Statistic P 2005 T = 201.0 0.312 2006 T = 262.0 0.597 2007 T = 181.5 0.569 All Years T = 1454.0 0.860 Table 2-5. Test for difference in male versus female weight of Sternotherus odoratus in the North Fork of the White River, Ozark County, Missouri, 2005-2007. Year Statistic P 2005 T = 200.0 0.332 2006 T = 258.0 0.928 2007 T = 153.0 0.107 All Years T = 1392.0 0.685 Table 2-6. Test for difference in male versus female plastron length of Sternotherus odoratus in the North Fork of the White Rive r, Ozark County, Missouri, 2005-2007. Year Statistic P 2005 T = 182.5 0.804 2006 T = 207.0 0.131 2007 T = 142.0 0.034* All Years T = 1253.5 0.035* Denotes significance at the =0.05 level.

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33 Table 2-7. Sex ratios and chi-square statistics for Sternotherus odoratus North Fork of the White River, Ozark County, Missouri, 2005-2007. Year Ratio males:females Chi-square P-value (2-tailed) 2005 1:1.82 2.065 0.151 2006 1:1.27 0.265 0.607 2007 1:1.38 0.516 0.472 All Years 1:1.43 2.306 0.129 Yates correction factor for continuity was applied to all cases to compensate for small sample sizes. A Figure 2-3. Size class distri bution by carapace length for Sternotherus odoratus captured in the 4.7 km research section of the North Fork of the White River, Ozark county Missouri. A) 1969. B) 1970. C) 1980. D) 2004. E) 2005. F) 2006. G) 2007. 0 1 2 3 4 5 6 7 8 9 10 0-3940-4950-5960-6970-7980-8990-99100-109110-119 120 Size Class (mm)Frequency Males Females Juveniles

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34 B C Figure 2-3. Continued 0 1 2 3 4 5 6 7 8 9 10 0-3940-4950-5960-6970-7980-8990-99100-109110-119 120 Size Class (mm)Frequency Males Females Juveniles 0 1 2 3 4 5 6 7 8 9 10 0-3940-4950-5960-6970-7980-8990-99100-109110-119 120 Size Class (mm)Frequency Males Females Juveniles

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35 D E Figure 2-3. Continued 0 1 2 3 4 5 6 7 8 9 10 0-3940-4950-5960-6970-7980-8990-99100-109110-119 120 Size Class (mm)Frequency Males Females Juveniles 0 1 2 3 4 5 6 7 8 9 10 0-3940-4950-5960-6970-7980-8990-99100-109110-119 120 Size Class (mm)Frequency Males Females Juveniles

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36 F G Figure 2-3. Continued 0 1 2 3 4 5 6 7 8 9 10 0-3940-4950-5960-6970-7980-8990-99100-109110-119 120 Size Class (mm)Frequency Males Females Juveniles 0 1 2 3 4 5 6 7 8 9 10 0-3940-4950-5960-6970-7980-8990-99100-109110-119 120 Size Class (mm)Frequency Males females Juveniles

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37 Discussion Distribution and Habitat The distribution of S. odoratus in the research section ha s remained virtually unchanged since 1969, with the primary difference being th e presence of the species in the farthest downstream station (station 50) of the section in 2007. Sternotherus odoratus is found in three distin ct areas: the west stream margins of statio ns 0-3 and 50, and the east stream margin from station 18-29. These three areas are separated by long stretches of river with gravel and bedrock substrates as well as by whitewater riffles, which both probably serve as barriers to movement of S. odoratus. No individual was observed to move betw een these areas in either a single season or between years. This obser vation, together with the pres ence of potential barriers to movement, indicates that there ar e probably three subpopulations of S. odoratus within the research section. In addition to being the only areas of the res earch section that contain thick muddy substrates, they are the only areas that al so contain dense stands of rooted underwater vegetation. These stands were considerably smaller or absent in 1969-1970, 1980 (Nickerson, personal comm.), and 2004 (personal observati on) than from 2005-2007. There are several likely explanations for the observed increase in vegetation. A flood event occurred just prior to the field season in 2004 that re gistered a daily mean discharge of over 15,000 cubic feet per second (cfs) (United States Geological Survey (USGS), 2008). This flood may have scoured submerged vegetation from the stream bed as well as displaced turtles downstream, both of which returned to the re search section in 2005 in the absen ce of any subsequent flood events. Nutrient loading is another likely cause of the increase in both aquatic ve getation and earlier and larger seasonal blooms of long fibrous algae. The research section of NFWR flows through a narrow valley, and atop bluffs on both sides of th e river are cow pastures. In addition, much residential development has occurred on the river upstream of the research section, and NFWR is

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38 heavily used in the summer for canoeing, kayaking, and tubing. All of these activities could be adding significant levels of nitrogen and phosphorus to the river, re sulting in increased growth of aquatic vegetation. Water qual ity testing upstream of, in, a nd downstream of the research section in summer 2007 revealed levels of total and fecal coliform bacteria that exceed those recommended for full body contact (Pitt, personal comm.). These high coliform counts indicate that human or animal waste is entering the rive r, and most likely causing an increase in nutrient levels. This could explain the increase in aquatic vegetation. More vegetation, and thus potential habitat, is probably why more S. odoratus were found in 2005-2007 than in historical surveys. While Ernst et al. (1994) repor t finding an individual in a gravel-bottomed stream in Arkansas, no S. odoratus were observed in open water on rock or gravel substrates in NFWR. Several individuals were captu red on emergent mats of Myriophyllum sp. and Ceratophyllum demersum that were rooted in and growing over rock and gravel. These areas were in close proximity to (~10 m) areas with mud substrate. These data indicate that presence of rooted vegetation, as opposed to soft-bottom substrates are more important to the distribution of S. odoratus in NFWR. Taking into account the preferred microhabitat of S. odoratus from this study as well as that reported in the literature (i.e., so ft bottom, emergent vegetation) and the area of that habitat in the research section, I estimate that S. odoratus could inhabit approximately 1.5 hectares, or 6% of the total area of the research section. This figure is arrived at by multiplying the approximate distance vegetation mats extend out from the banks (11 m) times the length of the number of stations occupied by vegetation mats in 2007 (1380 m) It is possible that S. odoratus utilizes the rock and gravel-bottomed sectio ns of river not included in this estimate, particularly during nighttime hours when surv eys were not conducted, but they were not

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39 observed doing so in this or previous studies of NFWR. Future studies of the NFWR should include nocturnal samp ling to determine if S. odoratus uses these widely available habitats. In contrast to Carr (1952) and Ernst et al. (1994) who reported that aerial basking (basking on objects out of the water) was uncommon in S. odoratus Nickerson (2000) found that North Fork individuals in 1969 and 1970 were most often observed basking aerially on logs or trees (15 observations), while aquatic basking was obser ved only three times. From 2004 to 2007, there were only six observations of aerial basking, whil e observations of partially submerged basking on mats of aquatic vegetation totaled 133. Over the period of Nickersons sampling, sycamores, willows, and other large trees were observed at the stream banks extending out over the river providing ample basking sites for turtles, as well as refugia in the exposed underwater root masses. These conditions no longer exist in the research section. Between surveys in 1980 and 2004, several massive floods occurred in the NF WR. Flooding in February and November of 1985 registered daily mean discharge of 32,000 cfs and 45,000 cfs, respectively (peak flow: 108,000 cfs and 133,000 cfs, respectively) at the dow nstream USGS monito ring station (USGS, 2008). Floods in September and November of 1993 registered daily mean discharge of 25,100 cubic feet per second (cfs) and 32,600 cfs, re spectively (peak flow: 52,500 cfs and 71,200 cfs, respectively) (USGS, 2008). The non-flood discha rge of NFWR is less than 2500 cfs (Nickerson et al., 2007). Flooding in NFWR between 3700 cfs and 5100 cfs in 1970 is known to have removed a massive sycamore tree from the research section (Nickerson et al., 2007). The floods of 1985 and 1993 likely washed out many more trees and may have led to the observed change in S. odoratus basking behavior. In addition, Pitt (2005) noted the general lack of basking sites in 2004 compared to historical studies. G. geographica by far the most abundant chelonian species in the research section, may be out-competing S. odoratus for a limited number of woody or

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40 rocky basking sites. Basking plays an importa nt thermoregulatory role in digestion and egg development, as well as aids in parasite removal. Boyer ( 1965) found that turtles basking on partially submerged substrates (such as vegetation or algal ma ts) were able to raise body temperature above that of the air or the water. Sternotherus odoratus in the NFWR may be able to thermoregulate effectively in this way, but apparently are una ble to receive the benefit of parasite removal, demonstrated by the prevalen ce of leech infestation on captured turtles. Movement Moveme nt of S. odoratus observed in this study is similar to patterns observed in other systems. Mahmoud (1969) found that the averag e distance traveled be tween captures was 66 m and 38 m for males and females, respectively, in less than 100 days and 69 m and 51 m for males and females, respectively, in greater th an 100 days. Ernst (1986) reported average movements of 117 m and 90 m for males and fema les, respectively. The average distance traveled between captures for males and female s in this study, 198 m and 102 m, respectively, follow the same pattern of longer distances tr aveled by males. Iverson and Meshaka (2006), analyzing data from several studies, speculate d that local conditions, rather than geography, dictate movement of S. odoratus This seems to be the case in NFWR. Both males and females were recorded traveling distances greater than 500 m, as well as not traveling any measurable distance. All recorded movements occurred within and may have been confined to the preferred microhabitat: submerged patches of rooted vegeta tion. A small-scale experiment intended as a pilot study was conducted during this survey to examine barriers to movement in S. odoratus. Four adults were displaced to the opposite bank of the river from where they were initially captured, a distance of approximately 50 m. Between the initial capture site and release site was a 1.5 m 2.5 m deep channel with a gravel/bedrock subs trate. To find the preferred habitat, turtles would need to cross the river or travel approximately 300 m downstream on the same side

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41 of the river. Displacement occurred in August 2006, and the turtles were not recaptured in 2007. Both the initial capture sites and the release sites, includ ing preferred and non-preferred microhabitat, were surveyed. While definitive results were not obtained from this side experiment, similar experiments could be done in rivers like the North Fork using radio telemetry to track the released turt les to answer this question. Population Structure and Demographics Small sam ple sizes combined with few recap tures during each sampling period prevented the determination of any reli able population estimates for S. odoratus in the research section. The sex ratios for 2005-2007 were all female-biase d, but none were significantly different from the expected ratio of 1:1 (Table 2-7). Ford (1999) also reported a slightly female-biased population in Missouri, and female-biased populati ons have also been reported for Michigan (Risley, 1933), Illinois, (Tinkle, 1961), Al abama (Dodd, 1989, Prassak et al., 1992), Louisiana (Rayburn et al., 1989), and Florida (Meshaka 1988, Aresco, 2005). Riselys (1933) high proportion of females in the population was attri buted to sampling bias (Gibbons, 1970 in Bury, 1979). Likewise Dodd (1989) stated that sampling over a wider area or over different portions of the reproductive season may have yielded a different sex ratio in Alabama. Females in the NFWR may have been captured more often than males because of increased activity related to nesting in the early-to-mid summer when sampli ng primarily occurred. Males would likely be encountered more often in the fall due to their search for mates. An examination of size class based on CL reve als that all size classes are fairly well represented and evenly distributed. The exceptions to this are the hatchling size class (under 40 mm CL) in all years and the distribution for the year 1980. Hatchlings were likely missed in sampling due to their small size and cryptic nature Hatchlings are most likely inhabiting the shallowest areas of the river that are not amenab le to sampling via snorkeling. In addition, eggs

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42 in Missouri are laid from June through August, and take two to three months to hatch (Johnson, 2000). Eggs may not have hatched during the field season that may explain the absence of hatchlings in samples. The 1980 data stand out in that only the largest size classes are represented (over 90 mm CL). While sampling for all other years occurred in the early to midsummer, 1980 sampling occurred late-summer to fall. Larger size classes may have been more conspicuous during this time of year due to movements related to the fall mating season. Juveniles were captured in every year except 19 80, suggesting that there is recruitment. The most striking detail that comes from the size clas s distribution graphs is the large increase in the number of turtles captured between 2004 a nd 2005-2007. As discussed above, 2005 saw an increase in the area of mats of emergent vegeta tion, as well as an increase in the amount of time the mats were present. In 2004, mats of vegeta tion or algae did not a ppear until early August, whereas from 2005-2007, these mats were present from the start of the field season in early June. The newly available habitat may have drawn S. odoratus from other parts of the river. In addition, drought conditions in the area may have caused S. odoratus to migrate into the study section from nearby cow ponds, creeks or backwaters. It is also possi ble that in 2004, more S. odoratus were present in th e research section but highly cr yptic, spending daytime hours buried in mud or hiding in muskrat or otter dens wh en the protection of dense submerged vegetation was not available. When the vegetation was present from 2005-2007, these turtles may have become more apparent by basking and feeding in the relative safety of the vegetation mats. It was not surprising that di morphism was not found between male and female carapace length, as dimorphism has been rarely reported in the species. The significant difference in plastron length may confer fitness benefits to ma les with small plastrons. Ford (1999) also found female S. odoratus in Missouri with larger plastron lengths than males, while the carapace

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43 lengths were similar. The biol ogical significance of this, cited by Ford in Gibbons and Lovich (1990), is that males with smaller plastrons may have a copulatory advantage through greater flexibility. Since greater plas tron lengths in females were no t observed in each year, further investigation of this pheno menon in NFWR is warranted. Male S. odoratus in the NFWR reach sexual maturity when CL ranges between 61 mm and 70 mm. The smallest male that could be identi fied based on secondary sexual characteristics was 70 mm. This individual was init ially captured in 2006 with a CL of 61 mm and sex could not be determined. It was recaptured 408 days later and was clearly a male based on secondary sex characteristics. Its age could not be determined accurately because of obscured plastral annuli, but the size at maturity is similar to that found by Mahmoud (1969) for males in Oklahoma (65 mm CL). Females in the NFWR were sexable between 57 mm CL and 65 mm CL. Most studies have found that female S. odoratus mature at larger sizes and gr eater ages than males (Risley, 1933; Tinkle, 1961; Mitchell, 1988; Edmonds, 1998). The findings on size at maturity in this study could be an artifact of small sample size, so future studies should involve some histological examination of specimens to more accurately determine the age and size at maturity of S. odoratus in the NFWR. Conclusions It is clear th at while there currently are more S. odoratus within the 4.6 km research section than were found in historical st udies, the number is still small, even without accu rate population estimates. The increase in number of turtles capt ured in recent years corresponds to the increase in available habitat. While si gnificant habitat alteration has o ccurred in and around the research section in the form of development, land cl earing, siltation, and ba sking site removal, distribution of S. odoratus within the research section has change d little in the last 38 years, with the species confined to three distinct areas.

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44 This study provides insight into the effects of land use change and habitat alteration on aquatic ecosystems. The NFWR has changed from a relatively pristine, remote river where few people were encountered during sampling periods in 1969 and 1970, to a heavily used recreation destination in 2007. Continued ha bitat alteration, by affecting th e behavior and abundance of species, may have consequences not only for S. odoratus but for the entire community. These data can inform future studies by providing a long-term pe rspective on a turtle species, as well as baseline data against which future surveys can be compared.

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45 CHAPTER 3 CONCLUSIONS Pitt (2005) and Nickerso n et al. (1984) doc umented dramatic changes in the NFWR watershed between 1969 and 2004, including large-scale land and riparian zone clearing, development, the removal of bask ing sites, nutrient loading, and increased recreational use. These factors appear to be benefiting the Sternotherus odoratus populations of the North Fork through an increase in mud, sedimentation, and growth of submerged aquatic vegetation. Many other ecologically generali zed chelonians have been show n not only to tolerate but flourish in polluted or degraded habitats. Molls (1977) study on the polluted Illinois River revealed that Common Snapping Turtles, Spiny So ftshells, False Map Turtles, and sliders were thriving while Yellow Mud Turtle s, Blandings Turtles, and Sm ooth Softshells were greatly reduced or absent. Likewise, Moll and Moll (20 04) note that several species, such as the Malayan Box Turtle, and the Black Pond Turtle have been more successful in impacted habitats than in more natural habitats (Moll and Khan bin Momin Khan, 1990 in Moll and Moll, 2004). Presumably, if generalists tend to benefit fr om degraded conditions, specialists likely would not. Pitt (2005) documented a decline in G. geographica a mollusk specialist, within the NFWR research section between 1969 and 1980, as well as a failure to rebound from the 1980 level by 2004. She attributed as least part of this failure to degraded habitat conditions. In addition, the generalist Trachemys scripta elegans became established in the research section over this time period, and Pitt (2005) determined that its distribution had expanded between 1980 and 2004. My study should not prove to be the end of re search in NFWR, as additional questions arose over the course of my res earch. While I did not encounter S. odoratus on rock or gravel substrates in open water, I sugge st that future studies in NF WR vary sampling both seasonally

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46 and temporally (nocturnal and diurnal) to determ ine variation in habitat use. Radio telemetry could also be employed to determine nocturnal habitat use. I suggest that the phenomenon of NFWR S. odoratus males having smaller PLs than females should be confirmed, as this phenomenon was not found to occur in 2005 or 2006. Likewise, I suggest that the age and size at maturity of NFWR S. odoratus should be confirmed, as findings in my study were based on just a few individuals. This could be accomplished by x-raying individuals in the lab for presence of ova or follicles, or by harvesting some individuals for hi stological examination. Finally, while my study has shown that the increase in S. odoratus abundance is correlated with an increase in rooted underwater vegetation, I suggest that the ca usative factors contributing to the increase in S. odoratus abundance be examined. Manipulative experiments involving vegetation and substrate, as well as examination of reproductive rate, recruitment, and migration from outside the study section could point to the factors causing the increase in S. odoratus abundance. My study provides further evidence of how gene ralist species can increase in abundance when habitats become degraded. Studies such as this, while important in a broad ecological context, will also add to the knowledge of Urban Ecology, a growing field with increasing importance as growing human populations expand into less populated, rela tively undisturbed areas.

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47 LIST OF REFERENCES Aresco, M.J. 2005. The effect of sex-specific terrestrial movem ents and roads on the sex ratio of freshwater turtles. Bi ological Conservation 123:37-44. Bancroft, G.T., J.S. Godley, D.T. Gross, N.N. Rojas, D.A. Sutphen, and R.W. McDiarmid. 1983. The herpetofauna of Lake Conway: species accounts. U.S. Army Corps of Engineers, Miscellaneous Papers, A-83-5. Army Engineers Waterways Experiment Station, Vicksburg, MS. Barko, V.A., J.T. Briggler, and D.E. Ostendorf. 2004. Passive fishing techniques: a cause of turtle mortality in the Mississippi River. Journal of Wildlif e Management 68:1145-1150. Boyer, D.R. 1965. Ecology of the baski ng habit in turtles. Ecology 46:99-118. Buhlmann, K.A., and J.W. Gibbons. 1997. Imperiled aquatic reptiles of the Southeastern United States: historical review and current conserva tion status. In G.W. Benz and D.E. Collins, (eds.), Aquatic Fauna in Peril: The Southeas tern Perspective. Special Publication 1, pp. 201-232. Southeastern Aquatic Research In stitute, Lenz Design and Communications, Decatur, GA. Buhlmann, K.A., and T.D. Tuberville. 1998. Use of passive integrated transponder (PIT) tags for marking small freshwater turtles. Ch elonian Conservation and Biology 3:102-104. Bury, R.B. 1979. Population ecology of freshwat er turtles. In M. Harless and H. Morlock, (eds.), Turtles: Perspectives and Research, pp. 571-602. Wiley, New York. Carr, A.F. 1952. Handbook of turtles. The turtle s of the United States, Canada, and Baja California. Cornell University Press, Ithaca, NY. Clark, W.S. 1982. Turtles as a food source of nes ting bald eagles in the Chesapeake Bay region. Journal of Field Ornithology 53:49-51. Congdon, J.D., J.L. Greene, and J.W. Gibbons. 1986. Biomass of freshwater turtles: a geographic comparison. American Midland Naturalist 115:165-173. Congdon, J.D., A.E. Dunham, and R.C. van Loben Sels. 1993. Delayed sexual maturity and demographics of Blandings turtles ( Emydoidea blandingii): implications for conservation and management of long-lived organisms. Conservation Biology 7:826-833. Davis, S.D. 1996. Reproduction and diet of Kinosternon odoratum inhabiting a cold water reservoir in Southwest Missouri. Unpubl. Ma sters Thesis. Sout hwest Missouri State University, Springfield. Delaney, M.F., and C.L. Abercrombie. 1986. Am erican alligator food habits in northcentral Florida. Journal of Wildlife Management 50:348-353.

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48 De Lima, A.G., W.E. Magnusson, and V.L. Da Costa. 1997. Diet of the turtle Phrynops rufipes in Central Amazonia. Copeia 1997:216-219. Dodd, C.K. 1989. Population structure and biomass of Sternotherus odoratus (Testudines: Kinosternidae) in a northern Alab ama lake. Brimleyana 15:47-56. Dodd, C.K., and R. Franz. 1993. The need for status information on common herpetofaunal species. Herpetological Review 24:47-50. Edmonds, J.H. 1998. Population ec ology of the stinkpot turtle ( Sternotherus odoratus) in Georgian Bay, Ontario. Unpubl. Masters Thes is. University of Guelph, Guelph, Canada. Edmonds, J.H., and R.J. Brooks. 1996. Demography, sex ratio, and sexual size dimorphism in a northern population of common musk turtles ( Sternotherus odoratus ). Canadian Journal of Zoology 74:918-925 Ernst, C.H. 1986. Ecology of the turtle, Sternotherus odoratus, in southeastern Pennsylvania. Journal of Herpetology 20:341-352. Ernst, C.H., J.E. Lovich, and R.W. Barbour. 199 4. Turtles of the United States and Canada. Smithsonian Institution Pre ss, Washington D.C. and London. Ford, D. 1999. Foraging ecology and demography of Sternotherus odoratus in a southwestern Missouri population. Unpubl. Ma sters Thesis. Southwest Missouri State University, Springfield. Ford, D.K., and D. Moll. 2004. Seasonal and sexual variation in foraging patterns in the stinkpot, Sternotherus odoratus in southwestern Missouri. Journal of Herpetology 38:296-301. Gaffney, E.S. 1990. The comparative osteology of the Triassic turtle Proganochelys Bulletin of the American Museum of Natural History 194:1-263. Gibbons, J.W. 1970. Sex ratio in turtles. Researches in Population Ecology 12:252-254. Gibbons, J.W., and J.E. Lovich. 1990. Sexual dimorphism in turtles with emphasis on the slider turtle (Trachemys scripta ). Herpetological Monographs 4:1-29. Iverson, J.B., and W.E. Meshaka. 2006. Sternotherus odoratus common musk turtle or stinkpot. In P. A. Meylan, (e d.), Biology and Conservation of Florida Turtles. Chelonian Research Monographs 3:207-223. Johnson, T.R. 2000. The Amphibians and Reptiles of Missouri, second edition. Missouri Department of Conservation, Jefferson City, MO. Kingsbury, B.A. 1993. Habitat use by the common musk turtle, Sternotherus odoratus in Crooked Lake, Indiana. Abstract, Annual mee tings of the Indiana Academy of Sciences, Purdue University, West Lafayette, IN.

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49 Lagler, K.F. 1943. Food habits and economic relati ons of the turtles of Michigan with special reference to fish management. Am erican Midland Na turalist 29:257-312. Mahmoud, I.Y. 1969. Comparative ecology of the kinosternid tur tles of Oklahoma. Southwestern Naturalist 14:31-66. Meshaka, W.E. 1988. Reproductive characteristics of South Florida Sternotherus odoratus and Kinosternon baurii (Testudines: Kinosternidae). Proceedings of the Arkansas Academy of Sciences 42:1988. Meylan, P.A., C.A. Stevens, M.E. Barnwell, and E.D. Dohm. 1992. Observations on the turtle community of Rainbow Run, Marion County, Florida. Florida Scientist 55:219-228. Missouri Department of Conservation. 2001. North Fork River watershed inventory and assessment. URL: http://www.mdc.mo.gov/fish/w atershed/northfrk/contents/. Accessed 20 April, 2008. Mitchell, J.C. 1988. Population ecology and li fe histories of the freshwater turtles Chrysemys picta and Sternotherus odoratus in an urban lake. Herpet ological Monographs 4:40-61. Mitchell, J.C. 1994. The Reptiles of Virginia. Smithsonian Instit ution Press, Washington, D.C. Moll, D. 1977. Ecological investigations of tu rtles in a polluted ecosystem: the Central Illinois River and adjacent flood plain lakes. Unpubl. Ph. D. Diss. Illinois State University, Normal. ----1986. The distribution, stat us, and level of exploitation of the freshwater turtle Dermatemys mawei in Belize, central. Biological Conservation 35:87-96. ----1990. Population sizes and foraging ecology in a tropical freshwater stream turtle community. Journal of Herpetology 24:48-53. Moll, D., and E.O. Moll. 2004. The Ecology, Exploitation, and Conservation of River Turtles. Oxford University Press, New York. Moll, E.O., and Mohd Khan bin Momin Khan. 1990. Turtles of Taman Negara. Journal of Wildlife and Parks X:135-138. National Climatic Data Center. 2008. Clim atological data, Missouri, July 2007. URL: http://www1.ncdc.noaa.gov/pub/or ders/8BDB9903-ED09-F97C-446668FA82AE6E86.PDF. Accessed 4 May, 2008. Nickerson, M.A. 2000. Testudines: Sternotherus odoratus (Common Musk Turtle): Aerial basking. Herpetologi cal Review 31:238-239. Nickerson, M.A., and C.E. Mays. 1973. The hellb enders: North American giant salamanders. Milwaukee Public Museum Publications in Biology and Geology 1:viii + 106 pp.

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50 Nickerson, M.A., C.E. Mays, M.D. Tohulka, and H.E. Barton. 1984. Changing chelonian populations in a spring-fed Ozark river. Program Abstract, Jo int Meeting of the American Society of Ichthyologists and Herpetologist s/Society for the Study of Amphibians and Reptiles, p. 165. University of Oklahoma, Norman. Nickerson, M.A., A.L. Pitt, and M.D. Prysby. 2007. The effects of flooding on Hellbender salamander, Cryptobranchus alleganiensis (Daudin 1803), populations. Salamandra 43:111-117. Palmer, W.M., and A.L. Braswell. 1995. Reptiles of North Carolina. University of North Carolina Press, Chapel Hill, NC. Pitt, A.L. 2005. Reassessment of the turtle comm unity in the North Fork of White River, Ozark County, Missouri. Unpubl. Masters Thesis. University of Florida, Gainesville. Plummer, M.V. 1979. Collecting and marking. In M. Harless and H. Morlock, (eds.), Turtles: Perspectives and Research, pp. 45-60. Wiley, New York. Pough, F. H., R. M. Andrews, J. E. Cadle, M. L. Crump, A. H. Savitzky, and K. D. Wells. 2004. Herpetology. Pearson Edu cation, Inc., Upper Saddle River, NJ. pp. 97-111. Powell, R., K.P. Bromeier, N.A. Laposha, J.S. Parmerlee, and B. Miller. 1982. Maximum sizes of amphibians and reptiles from Missouri. Transactions of the Missouri Academy of Science 16:99-106. Prassak, S.L., J.B. Sauge, and C. Guyer. 1992. Skewed sex ratios and sexual dimorphism in a population of musk turtles ( Sternotherus odoratus ). Society for the Study of Amphibians and Reptiles, 35th Annual Meeting Abstracts, p. 75. Pritchard, P.C.H. 1967. Living Turtles of the World. TFH Publications, Neptune City, NJ. Pritchard, P.C.H. 1989. The alligator snapping turtle: Biology and conservation. Milwaukee Public Museum, Milwaukee, WI. Rayburn, L.A., C. Fontenot, and J. Palis. 1989. Population ecology of the stinkpot turtle, Sternotherus odoratus in an urban pond. Proceedings of the Louisianna Academy of Science 52:70. Risley, P.J. 1930. Anatomical differen ces in the sexes of the musk turtle, Sternotherus odoratus (Latreille). Papers of th e Michigan Academy of Scien ce, Arts and Letters 11:445-464. Risley, P.J. 1933. Observations on the na tural history of the common musk turtle, Sternotherus odoratus (Latreille). Papers of the Michigan Academy of Science, Arts and Letters 17:685-711. Snider, A.T., and J.K. Bowler. 1992. Longevity of reptiles and amphibians in North American collections, second edition. Society for the Study of Amphibians and Reptiles, Herpetological Circular 21:1-40.

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51 Tinkle, D.W. 1961. Geographic variation in reproduction, size, sex ratio, and maturity of Sternotherus odoratus (Testudinata: Chelydr idae). Ecology 42:68-76. United States Geological Survey. 2008. Peak stream flow for Misso uri USGS 07057500 North Fork River near Tecumseh, MO. URL: http://nwis.waterdata.usgs.gov/mo/nwis/peak?site_no=07057500&agency_cd=USGS&for mat=html. Accessed 9 May, 2008. van Dijk, P.P., B.L. Stuart, and A.G.J. Rhodin. 2000. Asian turtle trade: Proceedings of a workshop on conservation and trade of freshwater turtles and tortoises in Asia. Chelonian Research Monographs No. 2. New York Tu rtle and Tortoise Society, New York. Vogt, R.C. 1981. Natural History of Amphibians and Reptiles of Wisconsin. Milwaukee Public Museum, Milwaukee, WI.

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52 BIOGRAPHICAL SKETCH Joseph J. Tavano was born 1978 in Hyannis, Massachusetts. He grew up in the town of Harwich, Massachusettes on Cape Cod where he graduated from Harwich High School in 1996. He earned a B.A. in iology fr om the University of Vermont in 2005. He began graduate school at the University of Florida in 2006 working toward an M.S. in nterdisciplinary cology. Upon graduating, Mr. Tavano plans to pursue a career in teaching or as a Wildlife Biologist.