Determination of Dispersal Patterns and Characterization of Important Habitats for Lake Sturgeon Restoration in the Upper Tennessee River System

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University of Tennessee, Knoxville TRACE: Tennessee Research and Creative Exchange

Masters Theses Graduate School

12-2015

Determination of Dispersal Patterns and Characterization of Important Habitats for Lake Sturgeon Restoration in The Upper Tennessee River System

Christina Grace Saidak University of Tennessee - Knoxville, [email protected]

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Recommended Citation Saidak, Christina Grace, "Determination of Dispersal Patterns and Characterization of Important Habitats for Lake Sturgeon Restoration in The Upper Tennessee River System. " Master's Thesis, University of Tennessee, 2015. https://trace.tennessee.edu/utk_gradthes/3604

This Thesis is brought to you for free and open access by the Graduate School at TRACE: Tennessee Research and Creative Exchange. It has been accepted for inclusion in Masters Theses by an authorized administrator of TRACE: Tennessee Research and Creative Exchange. For more information, please contact [email protected]. To the Graduate Council:

I am submitting herewith a thesis written by Christina Grace Saidak entitled "Determination of Dispersal Patterns and Characterization of Important Habitats for Lake Sturgeon Restoration in The Upper Tennessee River System." I have examined the final electronic copy of this thesis for form and content and recommend that it be accepted in partial fulfillment of the equirr ements for the degree of Master of Science, with a major in Wildlife and Fisheries Science.

J. Larry Wilson, Major Professor

We have read this thesis and recommend its acceptance:

Brian Alford, Mark A. Cantrell, Mike Jones

Accepted for the Council: Carolyn R. Hodges

Vice Provost and Dean of the Graduate School

(Original signatures are on file with official studentecor r ds.) Determination of Dispersal Patterns and Characterization of Important Habitats for Lake Sturgeon Restoration in the Upper Tennessee River System

A Thesis Presented for the Master of Science Degree The University of Tennessee, Knoxville

Christina Grace Saidak December 2015

Acknowledgements

I owe my deepest gratitude to my major advisor, Dr. J. Larry Wilson, for the opportunity to pursue an advanced degree in Fisheries Science, and for all of his support and guidance.

Without his assistance and dedication every step of the way, this thesis would not have been accomplished. I want to thank you, Doc, for your belief in me and for your patience and understanding these last few years, not to mention your eagerness to get me out the door!

It is also my pleasure to express a special thanks to the rest of my committee including

Mr. Mark Cantrell, Dr. Brian Alford, and Dr. Mike Jones. I worked with Mr. Cantrell during my summer internship with the U.S. Fish and Wildlife Service. He has been a major component in the completion of my thesis. His eagerness and enthusiasm for my project made me even more excited about the project. During the course of my study, Mr. Cantrell offered guidance and encouragement in all aspects of the project. He spent days and days on the river with me and many hours in the car traveling from one study area to another. Mark, I want to offer you my deepest thanks and appreciation for your continued leadership, support, and friendship.

Dr. Alford has been an invaluable member of my committee. He was always available to talk and offer advice and guidance during my project. I can’t express my gratitude enough, Dr.

Alford. Even though I have not had the opportunity to work with Dr. Jones, his impact on my project is evident throughout this thesis. I appreciate his eagerness and willingness to help during my project.

Completing my thesis required more than academic support, and I have many, many people to thank for listening, and, at times, having to tolerate my constant blabbering about Lake

Sturgeon over the past few years. I cannot begin to express my gratitude and appreciation for their support. Mr. Parker Hurst, Mr. Keith Garner, Mr. Carlos Echevarria, Mr. Chad Shirey, and iii

Mr. Brian Hickson have been unwavering in their support during the time I have spent at the

University of Tennessee working on my project. For many memorable hot days and cold evenings out on the river, I must thank you all. I would also like to thank all the people (too many to list here) who helped me collect and analyze data in the snow, the rain, the sleet, the storms, and the beautiful days.

Most importantly, none of this could have happened without the support of my family.

My sister Rebecca Ann offered encouragement through phone calls and cards – despite my own limited devotion to correspondence. With her own brand of humor, Rebecca has always supported me in all my crazy endeavors. And, to my Aunt June– it would be an understatement to say that, as a family, we have experienced some ups and downs in the past years but every time I was ready to quit, you did not let me, and I am forever grateful. This thesis stands as a testament to your unconditional love and encouragement.

Abstract

Lake Sturgeon, Acipenser fulvescens, are one of the slowest to reach sexual maturity and longest-lived freshwater fish species in North America. These fish are a species of special concern by the U.S. Fish and Wildlife Service, a vulnerable species by the American Fisheries

Society (Jelks et al. 2008), and a threatened species in Tennessee (Chiasson et al. 1997; Williams et al. 1989). They have been reintroduced into the Upper Tennessee River system since 2000.

Since December 2013, 49 Lake Sturgeon have been implanted with ultrasonic acoustic transmitters, and 26 fixed-station receivers installed throughout the Upper Tennessee River

System to monitor their movement. The objectives of this study were: 1) to determine dispersal and movement patterns of reintroduced Lake Sturgeon in the Upper Tennessee River system, 2) to identify water quality characteristics of seasonally important habitats, 3) to compare temperature and dissolved oxygen at summer refugia areas of known sturgeon concentrations, with other unused habitats, 4) to identify and assess potential spawning habitats in the Upper

Tennessee River system, and 5) to determine if dams inhibit upstream and downstream movements.

Lake Sturgeon implanted with acoustic transmitters were detected and monitored throughout the study area in the Upper Tennessee (RM 427-632), Clinch (RM 0-5), Hiwassee

(RM 5-501.9), Holston (RM 0-52.2), and French Broad Rivers (RM 0-32.3). There was a higher concentration of fish aggregating in Fort Loudoun Reservoir and smaller numbers in Watts Bar and Chickamauga Reservoirs. Movements varied with the majority of the fish traveling <5 km during the study period. Areas with aggregations of fish (>2 fish) were designated as “core use”.

During the study, there were 1,130,809 individual transmitter detections recorded.

The temperatures and dissolved oxygen (DO) levels recorded fell within acceptable ranges for Lake Sturgeon with the exception of the summer season in Chickamauga Reservoir.

Temperatures rose to >30 C and the DO levels dropped to <2 mg/L June through August 2014.

Gaining a better understanding of the factors affecting Lake Sturgeon recruitment and survival will be critical in designing restoration or reintroduction programs in the upper Tennessee River system and areas like it.

Table of Contents

Chapter 1: Introduction 1

Chapter 2: Literature Review 5

Morphology 6

Biology and Habitat 6

Movement 10

Chapter 3: Methods 13

Study Area 13

Upper Tennessee River 13

Methodology 17

Trotlines 17

Transmitter Implantation 17

Biotelemetry 20

Data Analyses 22

Biotelemetry Data Analyses 22

Water Quality Data Analyses 23

Chapter 4: Results and Discussion 25

Biotelemetry 25

Water Quality 33

Chapter 5: Summary and Recommendations 37

Future Research Recommendations 39

Literature Cited 41

Appendices 53

Vita 148

vii

List of Tables

Table 1. Table of data collected for all Lake Sturgeon outfitted with acoustic 55

transmitters during the study.

Table 2. Table of aging scheme for reintroduced Lake Sturgeon in the Upper Tennessee 57

River system courtesy of The Southeastern Lake Sturgeon Working Group.

Table 3. Table of fixed station Vemco receiver array in the Upper Tennessee River 58

system.

Table 4. Table of potential spawning sites for Lake Sturgeon below dams and in tailrace 59

areas in the Upper Tennessee River system.

Table 5. Table of minimum home ranges for Lake Sturgeon on Fort Loudoun 60

Reservoir.

Table 6. Table of minimum home ranges for Lake Sturgeon on Watts Bar Reservoir. 61

Table 7. Table of minimum home ranges for Lake Sturgeon on Chickamauga 62

Reservoir.

Table 8. Table depicting the overall minimum home range size for Lake Sturgeon 63

outfitted with acoustic tags on the Tennessee River, as determined by detection

at fixed array of acoustic receivers. Acoustic receivers were located at dams,

large tributaries, and known Lake Sturgeon habitats, to detect gross movement

patterns, not at regular intervals.

Table 9. Average daily water temperatures 2013-2015 measured at Fort Loudoun 64

Reservoir, Watts Bar Reservoir, and Chickamauga Reservoir on the Upper

Tennessee River compared to Fox River , Wisconsin.

viii

List of Figures

Figure 1 Morphology of the Lake Sturgeon (Courtesy of University of Michigan). 66

Figure 2 Map of the study area which includes the Upper Tennessee River (RM 427 67

– 632), Clinch River (RM 0 - 5), Hiwassee River (RM 0 - Holston River

(RM 0 – 52.2), and French Broad River (RM 0 – 32.3).

Figure 3 Photograph depicting eggs collected from wild brood stock on the Wolf 68

River in Wisconsin with collaboration between U.S. Fish and Wildlife

Service, the Warm Springs National Fish Hatchery, and the Wisconsin

Department of Natural Resources.

Figure 4 Map of Lake Sturgeon stocking locations on the Upper Tennessee River 69

system. Lake Sturgeon have been stocked as young-of-the-year and

fingerlings at Nance’s Ferry on the Holston River, and at Seven Islands on

the French Broad River.

Figure 5 Photograph of scute removal on fingerling Lake Sturgeon at the hatchery 70

prior to stocking in the Upper Tennessee River System

Figure 6 Photograph of wild-caught hatchery-reared Lake Sturgeon in the Upper 71

Tennessee River system on surgery table with MS-222 sedation.

Figure 7 Photograph of wild-caught hatchery-reared Lake Sturgeon in the Upper 72

Tennessee River study area on surgery table with incision open to determine

gender and sexual maturation by examination of gonads before implantation

of acoustic transmitter.

Figure 8 Photograph of VEMCO acoustic transmitters in a 70% alcohol bath ready to 73

be surgically implanted at a 45° angle into a wild-caught hatchery reared

Lake Sturgeon in the Upper Tennessee River system.

Figure 9 Photograph of sutures after implantation of acoustic transmitter into wild- 74

caught hatchery-reared Lake Sturgeon in the Upper Tennessee River study

area using a single synthetic, non- absorbable, ETHICON (Ethibond Extra

Purple Braided, 4.0 Metric) suture with an interrupted criss-cross suture and

a double knot.

Figure 10 Photograph of wild-caught hatchery reared Lake Sturgeon in the Upper 75

Tennessee River system in recovery tank after surgery to implant acoustic

transmitter before re-release into river.

Figure 11 Manual tracking reintroduced Lake Sturgeon movements by boat using 76

VEMCO VR100 with directional and omni-directional hydrophones in the

Upper Tennessee River system

Figure 12 Map of fixed acoustic receiver stations in the Upper Tennessee River 77

system extending from the Tennessee River downstream of Nickajack Dam

(RM 424.4) to the first dam at the Clinch, French Broad, and Holston

Rivers.

Figure 13 Side Scan Sonar image of Lake Sturgeon in Watts Bar Reservoir taken with 78

boat mounted Humminbird sonar.

Figure 14 Map of water quality sites in the Upper Tennessee River study area. 79

Figure 15 Photograph of Hydrolab sonde in Watts Bar Reservoir on U.S. Fish and 80

Wildlife Service research buoy at RM 573.4.

Figure 16 Photograph of Hydrolab sonde on U.S. Fish and Wildlife research buoy in 81

Fort Loudoun Reservoir at RM 628.7.

Figure 17 Photograph of Hydrolab sonde on U.S. Fish and Wildlife research buoy in 82

Chickamauga Reservoir at RM 487.3.

Figure 18 Map of “core use” areas in the Upper Tennessee River study area based on 83

current movement data

Figure 19 Map illustrating four possibilities for fish passage used by Lake Sturgeon 84

#26638 at the time that it moved through the Fort Loudoun dam in 2014.

Figure 20 Floating bar chart describing lockages for Fort Loudoun Dam from March 85

20, 2014, to April 4, 2014, during the time in which Lake Sturgeon #26638

passed through the dam.

Figure 21 Photograph of Lake Sturgeon below Chickamauga Dam April 20, 2015 86

(courtesy of Dan Walker University of Tennessee).

Figure 22 Graph illustrating Lake Sturgeon outfitted with acoustic transmitters that 87

also reported depth used various depths across the winter and spring

seasons. Lake Sturgeon #15954 was detected at deeper areas during

January and February, and in shallower areas 4 to 6 ft (1.2 to 1.8 m) for

extended times during late March, i.e., possibly at the sand bar area across

the river and just downstream of Blythe Ferry boat ramp, and/or the shallow

sand bar at the west end of Hiwassee Island. Both areas were within the

detection range of the fixed receiver at the bluff at RM 499.

Figure 23 Graph illustrating Lake Sturgeon outfitted with acoustic transmitters that 88

also reported depth used various depths across the winter and spring

seasons. Lake Sturgeon #15956 was detected at deeper areas during

December and January, while it was detected at 6 to 16 ft (1.8 to 4.9 m)

regularly during April and May.

Figure 24 Graph illustrating Lake Sturgeon outfitted with acoustic transmitters that 89

also reported depth used various depths across the winter and spring

seasons. Lake Sturgeon #15957 was detected at deeper areas during

December and January, while it was detected at 3.5 to 20 ft (1.1 to 6.1 m)

regularly during April and May.

Figure 25 Scatter graphs for Lake Sturgeon outfitted with transmitters with depth 90

sensor in Chickamauga Reservoir based upon river mile.

xii

Figure 26 Graph illustrating average daily water temperature comparisons from 2013- 91

2015 for Fort Loudoun, Watts Bar, Chickamauga Reservoirs in Tennessee

and the Fox River in Wisconsin with preferred low and high temperatures

for optimum Lake Sturgeon growth.

Figure 27 Seasonal temperature comparisons for Watts Bar, Chickamauga, and Fort 92

Loudoun Reservoirs and the Fox River for the winter (December 22 to

March 21) 2014.

Figure 28 Seasonal temperature comparisons for Watts Bar, Chickamauga, and Fort 93

Loudoun Reservoirs and the Fox River for the spring (March 22 to June 21)

2014.

Figure 29 Seasonal temperature comparisons for Watts Bar, Chickamauga, and Fort 94

Loudoun Reservoirs and the Fox River for the summer (June 22 to

September 21) 2014.

Figure 30 Seasonal temperature comparisons for Watts Bar, Chickamauga, and Fort 95

Loudoun Reservoirs and the Fox River for the fall (September 22 to

December 21) 2014.

xiii

Chapter 1

Introduction

Lake Sturgeon (Acipenser fulvescens) is considered a species of special concern by U.S.

Fish and Wildlife Service) (USFWS), a vulnerable species by the American Fisheries Society

Endangered Species Committee in all states where they occur (Jelks et al. 2008), and a threatened species in Tennessee (Chiasson et al. 1997; Williams et al. 1989). Since the 1960’s, population densities have declined dramatically throughout its range. Factors contributing to declining numbers of Lake Sturgeon include: overfishing, overharvesting of eggs (roe) and fillets, construction of hydroelectric dams that cut off migration routes for spawning, and water pollution and channelization that destroy spawning and nursery habitat (Etnier and Starnes 1993;

Galarowicz 2003). Restoration of this species is hindered by a lack of knowledge regarding spatial ecology and habitat use during all life stages (Altenritter et al. 2013).

Lake Sturgeon typically migrate to spawning grounds April through June once the water temperature reaches 12-15 C. Lake Sturgeon spawn periodically (annually or biannually for males and every 3 to 9 years for females) therefore yearly migrations do not occur for all individuals (Bruch and Binkowski 2002; Lyons and Kempinger 1992; Priegel and Wirth 1971;

Sheidegger 2012). However, due to construction of dams, short-term movement and long-term migration patterns of Lake Sturgeon have been restricted (Galarowicz 2003). Hydroelectric dams may also alter both water temperature and flow patterns which could impact timing and location of spawning events (Bain et al. 1988; Sheidegger 2012). Restrictions on migration could potentially lead to a reduction in survival and recruitment due to a loss in preferred spawning, resting, and feeding substrate (Knights et al. 2002; WDNR 2000).

The natural flow regimes of the French Broad and Holston Rivers were altered by the construction and operation of both the Douglas and Cherokee Dams in the 1940’s. The dissolved oxygen levels in the tailwater areas of these dams were also lowered, making these areas unsuitable for Lake Sturgeon spawning. Before the construction of dams and other barriers to movement, Lake Sturgeon were abundant throughout the large streams of the upper and middle

Mississippi River Basin, the Great Lakes and Hudson Bay drainages and the upper Coosa River

(Scott 1951). Old records, i.e., recreational catch reports, creel surveys, and anecdotal evidence suggests that Lake Sturgeon were present in the southern bend of the Tennessee River in

Alabama (Agassiz 1854) and Kuhne (1939) indicated that they were present but uncommon in the upper Tennessee and Cumberland rivers. Brimley (1946) reported them in the French Broad

River near Hot Springs, North Carolina, and Etnier and Starnes (1993) stated that there were scattered reports throughout the lower Tennessee River. In the 1960’s sightings were reported in

Fort Loudoun Reservoir and occasionally in Douglas and Watts Bar Reservoirs as well as the lower Little Tennessee River (now Tellico Reservoir).

Low dissolved oxygen levels were alleviated at both Douglas and Cherokee Dams in

1991 when The Tennessee Valley Authority (TVA) implemented the Reservoir Releases

Improvement/Lake Improvement Plan (Huddleston 2006). This plan improved minimum downstream flows by turbine pulsing in the Holston, French Broad, and Upper Tennessee Rivers

(Huddleston 2006). With improved dissolved oxygen levels in the tailwaters and other water quality improvements implemented under the auspices of the Clean Water Act of 1973, biologists believed that Lake Sturgeon could be reintroduced into the Upper Tennessee River

System (Huddleston 2006). The University of Tennessee (UTK) along with TWRA, USFWS, and the Wisconsin Department of Natural Resources (WDNR) agreed in 1995 to restore Lake

Sturgeon to their historic range in the Upper Tennessee River system, thus resulting in the formation of the Tennessee River Lake Sturgeon Working Group, which is now called the

Southeastern Lake Sturgeon Working Group (SELSWG 2014).

The Lake Sturgeon Reintroduction Program (LSRP) originated in 1998 with a long-term goal of removing Lake Sturgeon from the State endangered species list by re-establishing a self- sustaining population of Lake Sturgeon in the Upper Tennessee River system (SELSWG 2014).

In 2000, a pilot study of forty-one Lake Sturgeon hatched from eggs collected in 1998 from the

Wolf, Yellow, and Fox Rivers, Wisconsin, were implanted with acoustic transmitters and released into the French Broad River. Martin and Layzer (2001) indicated that these fish dispersed and were subsisting in the Upper Tennessee River system, which led to the implementation of a larger-scale effort rearing Lake Sturgeon in three National Fish Hatcheries

(NFHs) operated by the USFWS as well as at the Tennessee Aquarium Research Institute

(TNARI). Eggs and milt were provided by WDNR from wild brood stock in the Wolf, Yellow, and Wisconsin Rivers in Wisconsin (SELSWG 2014). Since July of 2000, approximately

160,000 young-of-the-year and fingerling Lake Sturgeon have been reintroduced into the French

Broad River system and below Douglas and Cherokee Dams.

The first movement study for the reintroduced Lake Sturgeon was conducted in 2003

(Huddleston 2006). Twenty juvenile Lake Sturgeon reared at Chohutta Fish hatchery in Georgia were implanted with radio transmitters and released into the French Broad River at Seven Islands

State Birding Park in Kodak, Tennessee. This study demonstrated that dispersal and movement patterns for these Age 2 fish were highly variable (Huddleston 2006).

There are a substantial amount of data available on Lake Sturgeon populations across

Canada and in northern U.S. river systems; however, there is limited information on populations

3 in the southeastern U.S. rivers. Research on Canadian and northern U.S. populations has enabled managers to identify “prime” sturgeon habitat, but what may be considered suitable habitat in those areas may be different for Lake Sturgeon populations in the Upper Tennessee River system, at the southern-most extent of the species’ range. Researchers speculate that due to the warmer water temperatures in the Tennessee River, the reintroduced sturgeon may mature faster and reproduce at an earlier age than the populations in Canadian and northern U.S. rivers.

However, more information is required to better understand dispersal patterns and seasonal movement as it relates to changes in water temperature and dissolved oxygen, as well as locating suitable spawning areas.

Today, biologists need to evaluate the success of the Upper Tennessee River Lake

Sturgeon Restoration Project by monitoring seasonal movement and dispersal patterns. The objectives of this study were: 1) to determine dispersal and movement patterns of reintroduced

Lake Sturgeon in the Upper Tennessee River system, 2) to identify water quality characteristics of habitats identified from movement studies as seasonally important, 3) to compare temperature and dissolved oxygen at summer refugia areas of identified sturgeon concentrations, 4) to identify and assess potential spawning habitats in the Upper Tennessee Basin, and 5) to determine if dams inhibit upstream and/or downstream movements. Gaining a better understanding of the factors that may affect Lake Sturgeon recruitment and survival will be critical to ensuring the success of restoration and reintroduction programs for Lake Sturgeon in the Upper Tennessee River.

Chapter 2

Literature Review

The Lake Sturgeon (Acipenser fulvescens) is part of a family of fish (Acipenseridae) that has existed for more than 150 million years, and is often referred to as a living fossil. Lake

Sturgeon are a fecund, potadromous species. The Lake Sturgeon is one of 27 sturgeon species worldwide (Huddleston 2006). The family’s unique set of life history characteristics, morphology, and habitat preferences have made them very susceptible to anthropogenic factors that impede migration, destroy habitat, and overharvesting of adults (Burr and Warren 2014;

Wilson and McKinley 2004). Lake Sturgeon are one of the slowest to reach sexual maturity and longest-lived freshwater fish species in North America. They have an intermittent spawning cycle process (Schram et al. 1999) with males reaching sexual maturity between 15 to 20 and females between 24 to 26 years of age (Becker 1983; Sheidegger 2012; WDNR 2000). Males may live up to 55 and females to over 80 with some records of individuals caught at of 152 years of age (Becker 1983; Sheidegger 2012). Population estimates for Lake Sturgeon range from only 33 individuals in a river system to over 34,000 individuals (Wilson and McKinley 2004).

Lake Sturgeon populations in many areas have declined dramatically since the 1960’s due to several factors. Throughout their range, populations have been impacted by overfishing, over-harvesting of eggs (roe) and fillets, construction of hydroelectric dams that cut off migration routes for spawning, and water pollution and channelization that destroy spawning and nursery habitat (Etnier and Starnes 1993; Galarowicz 2003). Lake Sturgeon once thrived in the

Tennessee River system; however, the last sighting, prior to reintroduction, was recorded in the

1961 (Etnier and Starnes 1993). They are currently listed as an endangered species in Tennessee

(Williams et al. 1989). Due to their life history characteristics and the other factors which have 5 led to their extirpation, Lake Sturgeon and similar species cannot return to their historical home ranges without help from a reintroduction or rehabilitation program (George et. al., 2009;

Huddleston 2006).

Morphology

Lake Sturgeon were described by Etnier and Starnes (1993) as having 9 to 17 dorsal plates, 19 to 42 lateral plates, 35 to 40 dorsal fin rays, and 25 to 30 anal fin rays. Their body color varies from light brownish gray to dark brown or black with some mottling, with juveniles having dark spots that fade as they mature. They have a elasmobranch-like heterocercal tail and five rows of sharp, bony scutes along their body which dull with age and are used for protection against predators when they are young (Wallus 1990). They have a protractible, toothless mouth with thick fleshy lips located under their shovel-like rostrum with four barbles, located anterior to the mouth that are lined with taste buds that they use to locate food in the substrate. Located on the rostrum are pores with ampullary organs that they use electroreception to detect weak electrical fields emitted by prey items (Miller 2004; New and Bodznick 1985; Teeter et al. 1980)

(Figure 1). Lake Sturgeon are carnivores in all life stages switching from zooplankton as larvae.

Crayfish, mollusks, aquatic insects, fishes, and other available benthic organisms are utilized by juveniles and adults (Jackson et al 2002; Wilson and McKinley 2004).

Biology and Habitat

Lake Sturgeon growth (weight and length) is fastest during a protracted juvenile stage but slows dramatically as they reach sexual maturity. This growth pattern is due to an unbalanced distribution of energy to somatic growth during the juvenile years (Beamish et al. 1996; Peterson et al. 2007). There is little information on the life history characteristics for Lake Sturgeon in the southern most extent of their range, but the information available indicates that warm water

6 temperatures show an increase in growth rates accompanied by earlier size and/or age-at- maturation and smaller maximum sizes (Fortin et al. 1996; Noakes et al. 1999). Based on this information, biologists speculate that Lake Sturgeon populations in the Tennessee River system should be smaller in maximum size, have a shorter lifespan, and should spawn earlier and more frequently than northern populations (Etnier and Starnes 1993; Power and McKinley 1997).

Water chemistry, and its physical property, is an important factor in the survival of Lake

Sturgeon populations. Water temperature and dissolved oxygen (DO) content are the most important water quality parameters that affect Lake Sturgeon at all life stages and year classes.

Temperature influences metabolic rates, growth, and species distributions (Lyons and Stewart

2014). Lake Sturgeon occur in areas with a multitude of temperature variations but the thermal preferences and tolerances are not well known.

All fish exist within a “thermal niche” that is defined by optimal conditions for growth, lethal limits and behavioral preferences (Lyons and Stewart 2014; Magnuson et al. 1979).

Laboratory testing on juvenile Lake Sturgeon indicate that the species survives and grows best in cool summer water temperatures similar to those favored by salmonid and cottid species and warmer temperatures like those preferred by centrarchid, ictalurid, catostomid, and percid species (Lyons and Stewart 2014). Generally, cool water fish species prefer water temperatures near 20-22 C with lethal temperatures at or near 31-35.1 C (Lyons et al. 2009; Lyons and Stewart

2014; Wilkes 2011). Lake Sturgeon can tolerate high and low water temperatures but they perform optimally at temperatures <25 C (77 F) (SELSWG 2014). Optimum spawning temperatures for Lake Sturgeon range from 9 to 20 C (46.4 to 68 F) (Burr and Warren 2014).

Cooler temperatures are required for proper egg development and to trigger ovulation in females

(Webb et al. 1999). Embryo and larval development is optimal between 11 to 20 C (52 to 68 F)

(Burr and Warren 2014). Although activity and growth of juvenile sturgeon increases with warmer temperatures, mortalities in sturgeon due to temperature have been documented in the

Mississippi River at temperatures >28 C (82 F) (Phelps et al. 2010). They are also more susceptible to low DO concentrations at higher water temperatures than other fish species.

Lake Sturgeon are benthic fish that regularly come into contact with hypoxic areas (areas with low DO), and they are less tolerant of hypoxia than most other fish species (Secor and

Gunderson 1998). Singer et al. (1990) indicated that Lake Sturgeon may use anaerobic metabolism during anoxic conditions producing alternative end products, thus avoiding lactic acidosis to ensure survival. Basically they utilize amino acids as oxidative substrates as opposed to sugars. Lake Sturgeon begin to suffer at DO levels < 6 mg/L and egg survival requires even higher dissolved oxygen (Secor and Gunderson 1998).

Lake Sturgeon are lithophilic riverine spawners, depositing eggs directly upon hard or rocky shorelines at depths between 0.5 to 4.7 m (1.6 to 15.4 ft) in swiftly moving water (Bruch and Binkowski 2002; Manny and Kennedy 2002; Wilson and McKinley 2004) with velocities between 12 and 22 cm/s and an optimal velocity of 16 cm/s (Peake 2006). Water temperature affects both the timing and length of spawning events for Lake Sturgeon. Spawning takes place from late April to early June and is cued by water temperatures between 8.3 to 16 C (46.9 to 60.8

F) (Bruch and Binkowski 2002; Burr and Warren 2014; Wilson and McKinley 2004). Lake

Sturgeon have an intermittent spawning cycle process with females spawning every 3 to 9 years and males every 2 to 3 years (Auer 1999; Kempinger 1996) with male Lake Sturgeon reaching sexual maturity between 15-20 and females between 24 to 26 years of age (Becker 1983;

Sheidegger 2012; WDNR 2000;). This reproductive strategy with late maturation and

8 intermittent spawning, has become a limiting factor in the reintroduction and rehabilitation of this species.

Males arrive at the spawning sites 1 to 2 days before the females when water temperatures reach 8.3 to 16 C (46.9 to 60.8 F). Spawning lasts 1 to 4 days and is diel; spawning is both polygamous and polyandrous (both sexes spawning with several individuals during a season), thus maximizing genetic diversity of offspring (Bruch and Binkowski 2002; Burr and

Warren 2014; Peterson et al. 2003). During spawning, a phenomenon known as porpoising occurs. This is a courtship behavior where sturgeon begin a spectacle referred to as “tail walking”. While “tail walking”, sturgeon thrust their heads above the water as if they are walking on their tails while swimming upstream. During porpoising sturgeon will sporadically jump completely out of the water. Porpoising occurs up to 14 days before spawning starts

(Bruch and Binkowski 2002). Once the females arrive at a spawning site, they are “green” which means they are not yet ovulating. However, after 1 to 2 hours ovulation begins. Once an ovulating female is detected by pheromones (Burr and Warren 2014) in the area, males will quickly approach either side of the female and begin nosing her sides and abdomen before pounding thunderously on her sides and abdomen, causing her to expel eggs in 5 second bursts, followed by the releasing of sperm (Burr and Warren 2014; SELSWG 2014). Active spawning has been reported when water temperature are between 8.3 to 21.1 C (46.9 to 70 F) (Burr and

Warren 2014).

Sturgeon have lower fecundity compared to other fish species with similar reproductive strategies. Lake Sturgeon have an average relative fecundity, with females capable of producing between 4,000-7,000 eggs per pound of fish during a single spawning event (USFWS 2014).

Lake Sturgeon are broadcast spawners expelling eggs coated with a greyish white sticky coating

9 which adheres to underlying substrate (Burr and Warren 2014; Bruch and Binkowskie 2002). A female Lake Sturgeon will participate in several spawning events until she has expelled all of her mature eggs with 240 second resting periods between bouts (Burr and Warren 2014).

Lake Sturgeon are potamodromous, living their entire lives in large freshwater rivers and lakes at depths between 4 to 9 m (13.1 to 29.5 ft) over a variety of substrates including silt, sand, clay, mud, rubble, and bedrock (Burr and Warren 2014; Kynard et al. 2000; SELSG 2013).

Dams are one of the main factors in the decline of sturgeon populations in the Southeast (Burr and Warren 2014; USFWS 2004). The shorelines below dams on the Tennessee River have been rip-rapped with large rocks which provide large interstitial spaces for spawning sturgeon. The constructions of wing-dams along the rest of the river to aid in barge navigation are also presumed to be additional areas with suitable spawning habitat for Lake Sturgeon (Huddleston

2006). However, even with adequate spawning substrate, tailwater releases in tributary reservoirs create high flows and high bottom velocities that can impede spawning. Dams have negatively impacted both the quality and quantity of spawning habitat and upstream spawning migrations in the Tennessee River but navigation locks do allow for some limited upstream movement (Knights et al. 2002). If Lake Sturgeon do successfully spawn below dams, post- spawning adult and juvenile fish will be at risk of entrainment and death from turbines during downstream migrations (SELSWG 2014).

Movement

Sturgeon movements are based on feeding optimization and reproductive success. Lake

Sturgeon move upstream to spawn and downstream movements are usually initiated to find better feeding grounds or to return to an area of summer or winter refugia (Auer 1996). Lake

Sturgeon exhibit both random and non-random movements. Upstream movements by spawning

10 individuals are offset by downstream movements of early life-history stages which redistribute individuals throughout the species range (SELSWG 2014). Movements by non-spawning Lake

Sturgeon are highly variable and can be dictated by thermal regimes and available forge (Auer

1996; Kynard 1997). Several studies have shown that most individuals within a population move randomly within a home range of 10 to 14 km (6.2 to 8.7 miles); however some fish exhibited unidirectional movements indicative of emigration (Peterson et al. 2007; Priegel and

Wirth 1974).

Previous studies conducted in the Mississippi River Basin, Mississippi and the Sturgeon

River, Wisconsin have suggested high site fidelity in adult individuals within rivers (Auer 1999;

Knights et al. 2002). Spawning migrations have been documented to cover nearly 500 km (311 miles) with a daily average of >22 km/day (13.7 miles/day) (Wilson and McKinley 2004).

Movements by non-spawning and early life stage individuals are extensive and highly variable.

Daily movements for non-spawning individuals has been documented at <0.2 km/day (<0.12 miles/day) to a maximum of <5 km/day (<3.1 miles/day); however, some individuals have exhibited extensive movements of >500 km (>311 miles/day) (Burr and Warren 2014; Wilson and McKinley 2004).

Sturgeon movements are strongly correlated to changes in water temperature.

Downstream movements from spawning areas to areas of summer resting or “summer refugia” by recently spawned or non-spawning adults and sub-adults have indicated that individuals don’t start to move until water temperatures approach 13 C (55.4 F) (McKinley et al. 1998). While there are exceptions as evidenced by a current feeding study at UTK (2015), sturgeon typically do not feed while in these summer refugia areas (Burr and Wilson 2004; Heise et al. 2005; Moser and Ross 1995) therefore they do not demonstrate much movement during this time. Once water 11 temperatures decline in the fall, their migration to deep >19.6 ft (>6 m), low-velocity, overwintering pools begins (Rusak and Mosindy 1997). Fish stay in these pools until water temperatures rise to 10 to 12 C (50-53.6 F) in the spring, which signals their migration to spawning areas.

Miranda and Killgore (2013) identified patterns in fish benthic distributions in the

Mississippi River and found that during the 14-year study, fish exhibited non-random depth distributions that varied by species and seasonally. They also found that most species used a wide range of depths with larger fish occurring in deeper water during the highest periods of the annual cycle. Depth is a key characteristic of aquatic ecosystems and an important factor in structuring fish assemblages (Miranda and Kilgore 2014). Fish tend to use habitats that are physiologically convenient in terms of DO, water temperature, and current (Mathews et al. 2004;

Miranda and Killgore 2013). There is an optimum depth range within the water column for abiotic and biotic conditions for each ontogenetic stage of fish; this optimum shifts seasonally and by ontogenetic life stage.

Chapter 3

Methods

Study Area

Upper Tennessee River

All figures and tables relating to this study are located in Appendices 1 and 2 of this text.

The study area on the Upper Tennessee River system (UTR) begins at the confluence of the

Holston and the French Broad Rivers. From there, the UTR travels southwesterly while absorbing additional tributaries of the Little, the Little Tennessee, and the Clinch Rivers; farther down the Hiwassee and Ocoee Rivers enter the UTR before reaching Chattanooga, Tennessee.

The river then enters the Tennessee River Gorge as it winds around the lower corner of the

Cumberland Plateau, reaching the lower portion of the Sequatchie Valley.

River miles (RM) will be used in this discussion as a substitute for longitudinal position along the Tennessee River and its tributaries, as depicted on navigation charts and USGS quadrangle maps, rather than kilometers (km). The study area extends from RM 424.7 of the

Tennessee River to RM 652.0 (Figure 2). Specifically, it is comprised of the Holston River from

Cherokee Dam (HRM 52.2) downstream to it confluence with the French Broad River (RM 652),

32.3 river miles of the French Broad River from Douglas Dam downstream to its convergence with the Holston River and 114 river miles upstream from Douglas Dam, 48 river miles of the

Holston River from Cherokee Dam downstream to their confluence forming the Tennessee

River, 50 river miles of the Tennessee River upstream of Fort Loudoun Dam (RM 530), and

Watts Bar Reservoir upstream of Watts Bar Dam (RM 530) with the lower reaches of the

Hiwassee River and Nickajack Reservoir. The UTR System includes a total of 174 miles (280 km) of riverine and lacustrine habitat. 13

The natural character of the Tennessee River has been altered by a series of impoundments, constructed by the Tennessee Valley Authority (TVA) from the late 1930s to the

1960s (Starnes and Etnier, 1986). These impoundments maintain downstream flows and are referred to as “flow through” or run-of-the-river impoundments. Therefore, river- like flows still persist on the upper portion of the reservoirs although the substrate has been altered extensively.

These areas once had an abundance of rocky gravel shoals; however, after the construction of the impoundments, the substrate and flows were greatly impacted, which in turn has obstructed Lake

Sturgeon from their critical habitat by curtailing movement and migration patterns.

Chickamauga Lake is a reservoir along the Tennessee River created when the

Chickamauga Dam was completed in 1940. The lake stretches from Watts Bar Dam at RM

529.9 to Chickamauga Dam at RM 471.0 making the lake 58.9 miles (94.8 km) in length. The

Hiwassee River empties into Chickamauga Lake at Hiwassee Island, just north of the Highway

60 Bridge at RM 500. Chickamauga Lake is immediately downstream from Watts Bar Lake and immediately upstream from Nickajack Lake. Actual lake levels vary due to weather conditions and power needs. There have been several commercial fishermen repots of Lake Sturgeon captured as by-catch in gill nets over the years. Acoustic transmitters were placed into nine Lake

Sturgeon in this area.

Watts Bar Lake is a reservoir on the Tennessee River created by Watts Bar Dam as part of the Tennessee Valley Authority system. Located about midway between Chattanooga and

Knoxville, the lake begins as the Tennessee River below Fort Loudoun Dam and stretches 72.4 miles (116.5 km) to Watts Bar Dam near Spring City. The Clinch River connects to the main channel of the lake at RM 568 near the Southwest Point in Kingston. The TVA’s Kingston

Stream Plant is just upriver from the convergence of the Clinch and the Tennessee Rivers.

In December 2008, the Kingston Fossil Plant spilled 1.1 billion US gallons of coal fly ash slurry into the Emory River, Watts Bar Reservoir, and the Clinch River when an ash dike ruptured. A study conducted by the Environmental Toxicology and Chemistry laboratories at

Appalachian State University in January of 2009 showed significantly elevated levels of toxic metals (including arsenic, copper, barium, cadmium, chromium, lead, mercury, nickel, and thallium) in samples of slurry and river water (Anonymous 2009). This is an area of water and sediment quality concern for Lake Sturgeon. Based upon movement data collected, there are regular aggregations of Lake Sturgeon in that area; VEMCO acoustic transmitters were implanted into 21 individual fish in this water body.

Fort Loudoun Lake is a reservoir in East Tennessee on the Upper Tennessee River. It extends approximately 50 miles (80 km) from the confluence of the Holston and French Broad

Rivers near Knoxville to Fort Loudoun Dam in Lenoir City. The reservoir is connected by the

Tellico Canal to Tellico Reservoir on the Little Tennessee River. Water is diverted through the canal to Fort Loudoun for power production. Tellico Canal also offers commercial barge access to Tellico without the need for a lock. The average depth is 22.3 feet (6.8 m), with a maximum depth of 86 feet (26.1 m), and an average width of 1,968 feet (0.6 km). A large number of Lake

Sturgeon have been caught and reported on this water body by anglers, and previously by commercial fishermen. There are currently 19 individual fish with VEMCO acoustic transmitters implanted aggregating in Fort Loudoun.

The French Broad River gradients are low to moderate and water is hard and relatively high in productivity. The substrate is mostly limestone bedrock, fine cherty gravel, and silty sand (Starnes and Etnier 1986). Depths below Douglas Dam range from less than 3.3 ft (1.0 m)

15 in shoal areas to 49.2 ft (15 m) in deep pools; however, water levels vary up to 9.8 ft (3 m) depending on discharges from Douglas Dam.

The Little River is a tributary to the Tennessee River which drains a 380-square-mile

(980 km²) area containing some of the most remarkable scenery in the southeastern United

States. There have been numerous angler reports recounted over the years below the Rockford

Dam. This is an area of interest as it has been noted as being a potential spawning area.

The Little Pigeon River is a tributary to the Tennessee River located entirely within

Sevier County, Tennessee. It rises from a series of streams which flow together on the dividing ridge between the states of Tennessee and North Carolina inside the boundary of the Great

Smoky Mountains National Park. The river is subdivided with three separate tributaries: East,

Middle, and West. The confluence of the two forks is at Sevierville; from there, the stream continues to flow northward until its confluence with the French Broad River just downstream from Douglas Dam. The area below Blalock’s mill dam is an area of interest as it has been noted as being a potential spawning area.

Nickajack Reservoir extends 46 rm upstream from Nickajack Dam to Chickamauga Dam.

When the Tennessee River leaves Chickamauga Dam, it flows through Chattanooga and forms

Nickajack Reservoir. Nickajack enters the Tennessee River Gorge 12 miles downstream of

Chattanooga. Nickajack Lake is filled with sloughs and outflow from dozens of underwater springs and caverns with the maximum depth reaching 145 ft (14.2 m). No Lake Sturgeon were captured or tagged here during the current study.

Methodology

Trotlines

Trotlines were deployed during the 2-year study 2013 to 2014, in the fall and winter months from November through December. Trotlines were used for the sampling because they target benthic fish such as catfish and sturgeon. Trotlines can be deployed in a variety of depths and habitats in the Tennessee River. Trotlines were 350 ft (107 m) in length and had 50, J-style hooks (size 2/0 to 4/0) attached to the main line by shorter dropper lines or “trots” spaced every 5 to 7 ft (1.5 to 2 m). A swivel was used to attach the trots to the main line using a half hitch knot, this connection is called the "staging." Hooks were then baited with 2.5 cm diameter chunks of cut buffalo fish (Catostomidae: Ictiobus spp.).

The swivels are necessary because Blue Catfish (Ictalurus furcatus), Channel Catfish

(Ictalurus punctatus), and Flathead Catfish (Pylodictis olivaris) are common by-catch and are notorious for twisting off hooks. Baited hooks were deployed on the bottom to improve the chances of a sturgeon successfully engulfing the hook. Trotlines were set oblique to the shore

(neither parallel nor perpendicular). Anchors weighing approximately 15 lbs each were attached to the line on the buoy end with a smaller weight anchor attached to the downstream end to maintain bottom position in the channel. Lines were left overnight for approximately 15 hours and “run” (retrieved) the following morning; and with limited exceptions were set oblique to shore.

Transmitter Implantation

Forty-nine Lake Sturgeon (1+ year classes) were captured primarily by trotline and were surgically implanted with acoustic transmitters (VEMCO 69 mHz acoustic transmitters, Amirix

Systems Inc., Nova Scotia, Canada) in the fall of 2013 and 2014 (Table 1). These fish were of

17 various ages, originally hatched at Warm Springs National Fish Hatchery in Warm Springs,

Georgia, from eggs collected from wild brood stock in the Wolf River, Wisconsin (Figure 3).

Hatchery fry were stocked as young-of-the-year (YOY) or as yearlings released into the French

Broad and Holston Rivers annually since 2000 as part of the Lake Sturgeon Reintroduction

Program (SELSWG 2014) (Figure 4). These subject fish were at liberty 2 to 12 years prior to transmitter implantation, enough time to establish wild movement patterns and randomly mix with the untagged population. There were twelve V13-1x (11.0 g.) transmitters each with an average battery life of 1,200 days, thirteen V16-4x (24.4 g.) transmitters each with an average battery life of 1,106 days, twenty V16-6x (34.2 g.) transmitters with an average battery life of

2,331 days, and four V16P-4x (36.0 g.) equipped with depth with an average battery life of 1,213 days. All transmitters were set with a 90 second delay between code transmissions. Transmitter weights did not exceed the maximum recommended 2% body weight of the fish (Bridger and

Booth 2003).

Prior to transmitter implantation, each captured fish was weighed to the nearest 0.5 lb

(0.2 kg) and both a total length and a fork length measurement was taken to the nearest millimeter and recorded. Fish were scanned for both coded wire tags (CWT) and passive integrated transponder tags (PIT) with frequencies of either 125 mHz, 130 mHz, or 134 mHz.

Readers used during this monitoring (e.g., Biomark 601 PIT tag reader) were capable of detecting all frequencies and were set to read as a decimal. If no PIT tag was detected, one was inserted just posterior to the dorsal fin on the right hand side; the ID number for the tag was recorded. Fish year class was determined by counting the missing scutes on either the left or right sides of the fish and recorded following the SELSWG (2014) aging protocol (Figure 5,

Table 2). Since 2005, tissue samples have been collected by taking a small fin clip from the left

18 pectoral fin on all captured fish. These tissue samples are collected, stored, and analyzed at numerous distinguishing microsatellite loci to document the baseline genetic makeup of the population, particularly with regard to parental relatedness and parental assignment of post- stocking recaptures (SELSWG 2014).

When Lake Sturgeon were caught, they were held in oxygenated live wells and transported to a main surgery boat where transmitters were implanted following procedures from

Warm Springs National Fish Hatchery, Warm Springs, Georgia, and modified from Wagner

(2011). Just prior to the surgical procedure, all surgical instruments were sterilized in a 70% alcohol bath. Fish were prepared for surgery by being anaesthetized using a bath solution of 100 mg/L of tricaine methano-sulfonate (MS-222); this solution was also used to irrigate the gills during surgery to ensuring sedation. Upon reaching a sedated anesthesia stage (i.e., slow movement and breathing reduced) fish were removed from the solution and placed on a moist surgery table.

A tube supplying fresh water and a low dose of MS-222 over the gills was placed in the mouth of the fish to maintain respiration (Figure 6). A sterile surgical packet containing all surgical instruments and supplies would be used to make a 0.4 to 0.8 inch (10 to 20 mm) incision. The incision site for implanting the transmitter, 1.6 to 2.4 inches (40 to 60 mm) anterior to the pelvic fins, or 3 to 5 scutes forward from the pelvic fin, was disinfected with betadine before the incision was made, after the incision was made, and again after the incision was closed. A retractor was used to open the incision to examine internal gonads for determining gender and maturation (Figure 7). A transmitter, sterilized in a 70% alcohol bath, was inserted into the body cavity at a 45º angle (Figure 8). The incision was closed using a single

19 synthetic, non- absorbable, ETHICON (Ethibond Extra Purple Braided, 4.0 Metric) suture and a double knot (Figure 9).

After a final application of betadine to the closed incision site, a thin coating of superglue was applied to ensure closure of the suture site. Post-surgery fish were held in an aerated recovery tank until they had completely recovered from anesthesia before release (Figure 10).

The surgical procedure and total holding time required no more than 15 to 20 minutes (including anesthesia induction, surgery and recovery). During the study there were more fish caught than those that were outfitted with transmitters. Some fish caught were deemed to have lower fitness

(small fish with low body weight) than expected, these fish were released after measurements were recorded and PIT tagging completed. Transmitters were not placed into unhealthy, stressed, or pre-spawning fish.

Biotelemetry

The relocation of tagged Lake Sturgeon was accomplished by: (1) roving, active tracking with a manual receiver, and (2) downloading data from an array of 27 fixed receivers and one receiver located on a working barge. Manual tracking by boat was conducted 3 to 5 days per week depending on water and weather conditions using a VR100 ultrasonic receiver (Amirix

Systems Inc., Nova Scotia, Canada) (Figure 11). There were two attempts at manually monitoring any diel patterns during the study. The VR100 detects all VEMCO coded transmitters, as well as collecting depth telemetry data from the implanted depth transmitters.

The VR100 was used with the VH165 omni-directional hydrophone and the VH110 (50 to 85 kHz) directional hydrophone. Locations for deploying water quality sondes in the study area was determined by analyses of sturgeon movements during manual tracking.

Twenty-seven fixed station VR2W-69kHz receivers (Amirix Systems Inc., Nova Scotia,

Canada) were strategically placed throughout the UTR based upon the rivers bathymetry, hydrology, and current Lake Sturgeon movement data. Acoustic transmitters are affected by a variety of water conditions; water quality, weather conditions, biological and man-made noise, mooring design, and bottom type/geomorphology of the area. A range test was completed for each receiver to estimate the receiver’s ability to decode transmitters at various distances. A range test tag with a fixed time delay of 15 seconds was placed in the water column at known distances from the receiver for 5 minutes with the expectation of receiving a minimum of 20 detections within a 5-minute period. A range test was performed each time a receiver was deployed and at various ranges.

The VR2W’s were usually bottomed moored. The VR2W’s were usually cabled to a tree, root, stump, or root off the bank by a stainless steel cable, the cable was stretched out parallel to the shore with the VR2W cabled at the other end with a small weight, the VR2W then had a small buoy saddled to the center so that the hydrophone floated up right off the river bottom with no shade area. This type of bottom mooring ensured that the receiver had an unobstructed “view” of any tagged fish that may swim within the expected acoustic range. The

VR2W receivers used a lithium 3.6 volt industrial D‐cell battery. Approximate battery life was

12 to 15 months.

Figure 12 indicates a map of the fixed array of receivers which extend from the

Tennessee River downstream of Nickajack Dam (RM 424.4) to the first dam at the Clinch,

French Broad, and Holston Rivers (Table 3). The receivers detected acoustic tagged fish at dams, tributaries, and other areas of known concentrations of Lake Sturgeon. These receivers were downloaded twice per month. The VR2W receivers are capable of identifying all VEMCO

69 kHz coded transmitters, providing robust and reliable information on the migration and movement patterns of the study species over great distances. Each time a VR2W receiver was downloaded, basic water quality data (DO, temperature, pH, and conductivity) were recorded using an onboard Hydrolab® DS5X multiparameter sonde (Hydrotech ZS Consulting,

Roundford, Texas). Data from three stationary Hydrolab sondes were also collected every three months, with battery changes occurring every 2 to 3 weeks.

When a fish was detected during manual tracking, the GPS location, the fish’s unique transmitter number, and water quality characteristics were recorded. The water column depth was recorded using a boat-mounted sonar unit (Humminbird 1199CI HD Series, Techsonic

Industries, Eufaula, Alabama) equipped with internal GPS side-imaging unit. If a Lake Sturgeon was detected by the sonar unit, a picture of the sonar image was taken and saved (Figure 13).

Data Analyses

Biotelemetry Data Analyses

Data collected from VR2W receivers were downloaded into the VEMCO User

Environment (VUE) Software version 2.1.3. This software was used for initialization, configuration, and data uploading from VEMCO acoustic monitoring receivers. This software also combined data from multiple receivers into a single integrated database. VUE was used for passive monitoring with the VR2W receivers. VUE software was used to download from receivers, time correct, filter, query, convert sensor data, and to graphically display data.

Detections collected in VUE were listed in order of date and were viewed by reciver and/or ID code.

A VR100 was used for active tracking of tagged fish. The VR100 data were downloaded into the VR100 PC Host software (VR100HS V3.2.1). These programs allowed the data

22 collected from each fish to be analyzed both individually and collectively. Data collected were summarized in the Microsoft Excel ™ (Microsoft Office 2010, Microsoft Corporation,

Redmond, Washington) program and then imported and analyzed using ArcGIS 10.0™

(Environmental Systems Research Institute, 2010, Badlands, California). Data analyses were performed on all 49 fish.

Individual maps were created for each fish (Appendix 3) and used to determine: 1) movement and dispersal patterns of reintroduced Lake Sturgeon in the Upper Tennessee River,

2) locations of seasonally important habitats, and 3) locations of potential spawning areas.

Tables were created illustrating distances moved by fish between original capture site and receiver detections for each tagged fish using the Microsoft Excel™ program. Core use areas and areas of seasonally important habitats were determined by fish aggregations and by using the

Presence/Absence tool in the VUE software. Smith and Clugston (1997) determined that aggregations were determined to be an area where >2 fish were present at the same time.

Water Quality Data Analyses

Three Hydrolab sondes were deployed in May 2014, each anchored to a research buoy with a 160 lb central anchor. A fourth Hydrolab sonde was kept on the boat to be used during manual tracking with the VR100. Each time a transmittered fish was detected the time, GPS location, depth, and water quality parameters were recorded. The fourth Hydrolab was also used to re-calibrate and switch out fixed station Hydrolab sondes as needed. Batteries were replaced and all sondes were re-calibrated following the manufacturer’s instructions; sensors and units were also cleaned and repaired if necessary following the manufacturers guidelines before each re-deployment for further water quality monitoring.

Figure 14 depicts a map of the three fixed station Hydrolab DS5X multiparameter sondes which were placed in specific locations in the study area based on the current movement data:

Watts Bar Reservoir at RM 573.4 (Figure 15), Fort Loudoun Reservoir at RM 628.7 (Figure 16), and above Hiwassee Island at RM 487.3 (Figure 17). These sondes were equipped with DO and temperature sensors with a central brush for long-term unattended monitoring. Turbidity was not included in this study because current research suggests that turbidity does not play a vital role in the daily feeding and movement of Lake Sturgeon since they are benthic feeders with poor eyesight.

Temperature and DO data were also collected from The Tennessee Valley Authority

(TVA) River Operations Center for Fort Loudoun, Chickamauga, and Watts Bar Reservoirs,

Tennessee (courtesy of Mr. Arthur Ramsey), and from the US Geological Survey (USGS) for the

Fox River, Wisconsin. Water quality parameters from the Hydrolab multiparameter sondes were downloaded and analyzed using Hydras 3LT® Windows® software (Hydrotech ZS Consulting,

Roundford, Texas) then graphed in the Microsoft Excel ™ program. The least-square means of temperature among the three reservoirs and the Fox River were then compared on a seasonal basis. SAS Enterprise Guide v. 6.1 was used to analyze the temperature data with a generalized linear model in the PROC GENMOD procedure. A Poisson distribution and a log-log link were assumed.

Chapter 4

Results and Discussion

Biotelemetry

Core use areas, or holding areas, are areas where aggregations of fish have been detected during the summer and fall months. These core use areas were detected in bends of the river, at the head of submerged islands, in the tailwaters, or at the confluences of tributaries (Knights et al. 2002), and typically >16.4 ft (>5 m) deep (Erickson et al. 2002) in the lower portions of the river (McKinley et al. 1998). An area with an aggregation of at least 2 fish was considered a core use area. Lake Sturgeon in the UTR system demonstrated a similar trend in distinct core use areas (Figure 18). These locations were identified and monitored continuously; water quality parameters were recorded regularly at these sites. Knights et al. (2002) found that Lake Sturgeon on the Mississippi River exhibited core use in bends of the river, confluences of tributaries, and in the tailwaters. Huddleston (2006) described a similar trend in juvenile Lake Sturgeon in the

French Broad River.

Huddleston (2006) found that movements of juvenile Lake Sturgeon in the French Broad

River were correlated to temperature and discharges from Douglas Dam; juvenile fish moved upstream during periods of increased flow and downstream during periods of decreased flow.

Kynard (1997) found that in southern rivers, some long distance summer migrations of Shortnose

Sturgeon may have reflected greater energy resources due to warmer water. Pre-spawning Lake

Sturgeon in Wisconsin also exhibited a migration from downstream core use areas to deep overwintering pools (Bruch and Binkowski 2002).

Overwintering pools were also identified and monitored during this study; water quality parameters were recorded regularly at these sites as well. Overwintering pools were described 25 by Rusak and Mosindy (1997) as deep >19.7 ft (>6 m) slow flow areas where fish aggregations had been detected. Pools in the study area of the Tennessee River range up to 90 ft (27 m) in outer bends adjacent to limestone bluffs. While Rusak and Osindy (1997) found Lake Sturgeon remained in these overwintering pools until they were triggered to migrate to adjacent spawning areas by rising water temperatures in the spring, sub-adult fish in this study remained in these areas year-round, moving short distances possibly for feeding.

Abundance of food and thermal regimes predict movement for non-spawning potamodromous species (Kynard 1997). Upstream migrations occur in the fall, winter, and spring for spawning adults with downstream movements occurring in the summer and fall

(Warren and Burr 2014). Downstream movements were usually associated with locating abundant food sources and areas with favorable water quality and habitat (Auer 1996; Sullivan et al. 2003).

All 49 Lake Sturgeon outfitted with acoustic transmitters were detected again after implantation of transmitters. The VUE software documented 1,130,809 individual transmitter detections during this study. Upon implantation of transmitters, gender, and sexual maturity were determined. The sex ratio of the 49 fish used in this study was approximately 5:1:2, 63% being female, 12% of unknown gender, and 23 % male. All fish outfitted with transmitters during this study were determined to be in Stage I gonadal development (Wildhaber et al. 2006), except two that appeared to be in Stage II; therefore, none of the fish in this study were determined to be close to spawning age or sexual maturity. Based on this information, along with the movement data collected during the study, there were no gross movement patterns that would suggest an upstream spawning migration. Some Lake Sturgeon in this study did make

26 substantial upstream and downstream relocations but these movements cannot be attributed to spawning.

Sturgeon species vary in diel movements (Warren and Burr 2014). Gulf Sturgeon

(Acipenser oxyrinchus desotoi) show strong diel movement patterns, moving more at night during all seasons except summer (Sulak and Clugston 1999; Wrege et al. 2012), whereas

Shortnose Sturgeon exhibit no diel movement patterns (Kynard et al. 2007). During the present study, two continuous movement tracking efforts were conducted in order to document diurnal movement patterns. During this portion of the study, fish were detected and then manually tracked continuously once for approximately 6.5 hours and again for approximately 12 hours to evaluate diurnal movement and/or forging patterns. The first diurnal monitoring effort was on

July 16, 2014, on Ft. Loudoun Reservoir. Monitoring started at Georges Creek, RM 630, beginning at 3:00 pm and ending at 9:00 pm EST. Fish ID #26647 was the only fish detected and it was found at a depth of 60 ft (18.3 m) in the channel. Water quality data and depth were recorded by the onboard Hydrolab sonde. There was no movement detected from this fish during this monitoring effort.

The second diurnal monitoring effort was on July 30, 2014, on Watts Bar Reservoir.

Monitoring started at the James Ferry TWRA boat ramp at RM 569, beginning at 8:30 am and ending at 9:00 pm EST. Five fish, ID #26638, #28533, #29935, #28530, and #26642, were detected and monitored continuously during this effort. Water quality and depth data were recorded in each location where a fish was detected, and periodically throughout the day. The monitored fish did not display any distinct diel movement pattern and remained relatively sedentary during the monitoring periods. Telemetry data indicated that both juvenile and adult

Lake Sturgeon preferred some areas over others and generally aggregate in certain locations. In

27 general, sturgeon movement patterns fluctuated between species, populations, and even individuals. The exceedingly variable movement patterns demonstrated by Lake Sturgeon could be attributed to factors which include: environmental avoidance (i.e. changes in water temperature and dissolved oxygen levels >6 mg/L), discharge from dams, foraging, or innate behaviors and reproductive success. Early life history stages of sturgeon species instinctively migrated downstream beginning with the larval drift soon after hatching, with the distance traveled during drift based on water velocities, discharges below dams, and location in the water column (Auer and Baker 2002; McLeod et al. 1999).

Acoustic receivers were placed near dams and large tributaries during this study to locate potential spawning sites. During this study, one fish, #26638, was detected passing through the

Fort Loudoun Dam on Fort Loudoun Reservoir. Figure 19 illustrates four ways in which fish

#26638 could have traversed the dam: 1) through the Loudoun Powerhouse - through the turbines - downstream only, 2) through the Loudoun Dam Spillway - during gate openings - downstream only, 3) through the Loudoun Lock - during vessel lockage - upstream or downstream, or 4) through the Tellico Dam - during gate openings - downstream only. Figure 20 is a floating bar chart describing the lockages from March 20, 2013, to April 4, 2014. This fish could also have passed through the lockage. This fish went undetected for several weeks before passing the next receiver downstream of the dam at RM 598. It then traveled downstream to the

Clinch River and is presumably now in the Emory River at RM 5 since passing through the dam.

Although the Corps of Engineers reported one lockage during that timeframe, it cannot be determined if that fish passed through the lock or through the turbines.

Dams can inhibit the migration routes of many fish species. Disruption of natural flows downstream may inhibit upstream migration cues for spawning adults (Cooke et al. 2002), while

28 post-spawning adults and juvenile Lake Sturgeon are at risk of entrainment and death from turbines during downstream migrations (Jager et al. 2007). Dredging and channelization above and below dams for navigation destroys both river and estuary habitat for Lake Sturgeon

(Kynard 1997).

While dams are barriers to fish movement, these fish have demonstrated that they are able to move past dams, at least downstream. While there have been numerous documented occurrences of tagged sturgeon passing through navigation locks, countless numbers of sturgeon are delayed and even prohibited from reaching spawning or preferred feeding areas. Fish

#26638 remained in the tailwater area near the dam for several weeks before moving farther downstream, it is unclear if the fish was trying to move back upstream through the dam once it had passed downstream.

In April of 2015, anglers reported two Lake Sturgeon below Chickamauga Dam (Figure

21). The two fish were reported in the area for several hours and looked to be in good physical condition, with one fish averaging 3 to 4 ft in length and the other around 2.5 ft in length.

Typical spawning behaviors were not observed and the fish were maneuvering independently of each other. The area below Chickamauga is rip-rapped and the temperature of the water at the time of the observation was 17 C (62.6 F). Potential spawning sites below dams and tailrace areas were monitored throughout the study (Table 4).

Table 5 indicates the minimum home ranges measured for all 19 Lake Sturgeon outfitted with acoustic tags at Fort Loudoun Reservoir on the Tennessee River, as determined by detection at fixed array of acoustic receivers. The home ranges on Fort Loudoun Reservoir ranged from

3.6 to 23.1 river miles (X = 8.41 river miles, n = 18; this data excludes fish #26638 because it moved to another reservoir). Fish #26638 was captured at RM 628.9 on Fort Loudoun and has

29 traveled to RM 5 on the Emory River (Appendix 3. Pages 98-100). This fish had a total tracked range of approximately 64 river miles, and is the only fish that successfully navigated through or past a dam during this study.

Table 6 documents the minimum home ranges measured for all 21 Lake Sturgeon outfitted with acoustic tags at Watts Bar Reservoir on the Tennessee River, as determined by detection at fixed array of acoustic receivers. The home ranges on Watts Bar Reservoir ranged from 2 to 47.5 river miles (X = 9.77 river miles, n = 21). Fish #28529 was captured at RM

573.28; it traveled both upstream and downstream of its capture location, however, the most interesting movement was its downstream movement to Watts Bar Dam at RM 530. This fish then moved back upstream as far as RM 576 (Appendix 3, Page 129).

Table 7 illustrates the minimum home ranges measured for seven of the nine Lake

Sturgeon outfitted with acoustic tags at Chickamauga Reservoir on the Tennessee River, as determined by detection at fixed array of acoustic receivers. The home ranges on Chickamauga

Reservoir ranged from 5 to 10 river miles (X = 7.18 river miles, n = 7). Two of the fish tagged in

December 2014, have not yet been detected again since their release so they are not included in

Table 7. Table 8 shows the overall minimum home range size for Lake Sturgeon outfitted with acoustic tags on the Tennessee River, as determined by detection at fixed array of acoustic receivers. Acoustic receivers were located at dams, large tributaries, and known Lake Sturgeon habitats, to detect gross movement patterns, not at regular intervals.

Fish #29936 and #29937 were captured at RM 501.8 in November 2013 and have not been detected again, #29936 not since March 29, 2013, and #29937 not since May 10, 2013

(Appendix 3, Pages 147-148). Chickamauga Reservoir is currently the only reservoir in the study area where commercial fishing is still allowed. One speculation as to why these fish have

30 not been detected again is that they had been removed due to commercial fishing bycatch. If this situation occurred, they were either illegally harvested or died due to rising water temperatures and the stress of being netted for an extended period.

Curtis et al. (1997) described similar movements in Shovelnose Sturgeon

(Scaphirhynchus platorynchus) on the Mississippi River with some fish having restricted home ranges while others had extensive home ranges >1 km. Huddleston (2006) found these same patterns during her study of juvenile Lake Sturgeon dispersal in the French Broad River. She found that while some fish remained sedentary others were very widely dispersed within the study area after release.

The Tennessee River is a heavily impacted river where anthropogenic changes have altered temperature, depth, discharge, and other related habitat characteristics; therefore, understanding the distributions of benthic fish species like Lake Sturgeon along various depth gradients to minimize species impacts in a heavily altered river is very significant. All sturgeon species have developed morphological adaptations to deep-water habitats such as a ventral protractile mouth, small eyes, and compact elongated bodies (Miranda and Killgore 2013).

Allen et al. (2007) indicated that groups of Shovelnose Sturgeon and Pallid Sturgeon

(Scaphirhynchus albus) used deep areas significantly more, and medium and shallow areas less, based on availability in an experimental river study. Biotic factors such as prey availability and competition also play a crucial role in depth selection by fish species (Eckmann and Imbrock,

1996; Mous et al. 2004).

During the sampling season in December 2014, three fish at Chickamauga Reservoir were outfitted with acoustic transmitters which were equipped with depth sensors. The Lake

Sturgeon that were outfitted with depth sensors were detected using various depths across the

31 winter and spring seasons. Lake Sturgeon #15954 was a very healthy male that was commercial bycatch during a scheduled December 2014 trotline sampling season on Chickamauga Reservoir.

This fish (the largest fish captured since the reintroduction began) had a total length (TL) of 51 in (4.3 ft), fork length (FL) of 48.5 in (4 ft), and a weight of 32 lbs (14.5 kg). Fish #15954 was detected in deep reaches of the reservoir at depths up to 65.7 ft (20 m) during January and

February, while it was detected at shallower depths ranging from 4 to 16 ft (1 to 5 m) for extended times during late March, likely at the sand bar area across the river and just downstream of Blythe Ferry boat ramp, and/or the shallow sand bar at the west end of Hiwassee

Island, both areas within the detection range of the fixed receiver on the bluff at RM 499 (Figure

22).

Lake Sturgeon #15956 at Chickamauga Reservoir was also outfitted with an acoustic transmitter that reported depth. This fish was a smaller female with a TL of 44.5 in (3.7 ft), FL of 42.5 in (3.5 ft), and weighed 12.57 lbs (5.7 kg). Similar to fish #15954, fish #15956 was detected at depths up to 66.7 ft (20.3 m) during December and January, while it was detected at depths ranging from 6 to 16 feet regularly during April and May (Figure 23).

Fish #15957 at Chickamauga Reservoir was the third Lake Sturgeon outfitted with an acoustic transmitter equipped with a depth sensor during the 2014 sampling season. This fish was a male with TL of 42 in (3.5 ft), FL of 39 in (3.25 ft), and weighed 11.7 lbs (5.3 kg). Fish

#15957 illustrated the same depth pattern as both Lake Sturgeon #15954 and #15956. This fish was also detected deepest during December and January, reaching depths of 70.1 ft (21.4 m) during December and January, while it was detected at depths ranging from 3.5 to 20 ft (1.1 to

6.1 m) regularly during April and May (Figure 24).

In general, the Lake Sturgeon outfitted with acoustic transmitters equipped with a depth sensor, used the maximum depths in the vicinity of each fixed receiver. Figure 25 shows scatter graphs of each location that the Lake Sturgeon outfitted with depth transmitters were detected.

Based on bathymetry maps and soundings with boat-mounted sonar, the maximum depth within the listening range of the receiver at RM 497 was 55 ft (16.8 m), 48 ft (14.6 m) at RM 499, 71 ft

(21.6 m) at RM 502, and 48 ft (14.6 m) at RM 505.

Vannote et al. (1980) and Junk et al. (1989) highlighted that studies on fish distributions in rivers have mainly concentrated on longitudinal and latitudinal spatial distributions rather than depth distributions because most rivers are fairly shallow. However, the Tennessee River is generally deeper compared with other lentic systems where Lake Sturgeon occur. This is due to the fact that the Tennessee River is impounded and has been profoundly altered for commercial navigation and hydroelectric power. Fish have been found to hold up in the deeper bends in the

Tennessee River than in the shallower areas. Understanding depth distributions in altered rivers is necessary to better understand the habitat requirements of fish communities.

Water Quality

Figures 27-30 compare thermal characters of the study area to the Fox River, Wisconsin, the source of the broodstock used for the Tennessee River restoration project. During the months of warmer water temperatures and lower dissolved oxygen levels, current research on Lake

Sturgeon indicates that the fish primarily resided in deep summer pools during the daylight hours when the water temperature is highest, then foraged for food in very shallow shoals during the night when water temperature was coolest. Water quality measurements are important during these diurnal and nocturnal movement patterns because they provide water quality variables that can be compared to both day and night water conditions in areas where the sturgeon aggregate,

33 seasonal water quality comparisons can also be made using these data. Field observations with juvenile Lake Sturgeon indicate that, in general, the species survive and grow best at ‘cool’ summer water temperatures (Lyons and Stewart 2014).

Lake Sturgeon prefer water temperatures <25 C (77 F), however, they can tolerate much warmer temperatures (SELSWG 2014). As water temperatures rise during the warmer summer months, pockets of cool water in areas where water is not as mixed or where an underwater spring bubbles up through the substrate can provide areas of “summer refugia” for Lake

Sturgeon. These areas of refugia can sustain aggregations of sensitive species (Sedell et al.

1990). The deep overwintering pools where the transmittered fish have remained since capture in November may be areas of groundwater flow or cool springs.

While activity and growth of juvenile sturgeon increase with warmer temperatures, Lake

Sturgeon become stressed at temperatures >28 C (>82.4 F); Lyons and Stewart (2014) defined these temperatures as ‘marginal,’ with temperatures at or above 30 C (86 F) defined as

‘stressful’. Most water temperatures in the Upper Tennessee River study area where Lake

Sturgeon aggregate were found to be neither ‘marginal’ nor ‘stressful’. For most of the study area, the maximum daily water temperatures ranged between 19 to 26 C (66.2 to 78.8 F) however, the average maximum daily water temperatures found on Chickamauga Reservoir regularly exceeded the ‘marginal’ and ‘stressful’ temperature tolerances for Lake Sturgeon. The average maximum daily water temperatures on Chickamauga Reservoir regularly rose >30 C (86

F) from June to until October, when temperatures started to decline. Table 9 reflects the average daily water temperatures for the study area at Fort Loudoun, Watts Bar, and Chickamauga

Reservoirs in Tennessee, and compared to the Fox River in Wisconsin, where some of the broodstock for the Tennessee restoration were collected. Figures 26 illustrates average daily

34 water temperature comparisons from 2013 to 2015 for Fort Loudoun, Watts Bar, and

Chickamauga Reservoirs and the Fox River with preferred low and high temperatures for optimum Lake Sturgeon growth.

Aside from temperature, the most important water quality parameter that affects Lake

Sturgeon is DO. Dissolved oxygen refers to the amount of oxygen that is dissolved in water at a given temperature and given atmospheric pressure. As water temperatures increase, the concentration (mg/L) of DO in a volume of water decreases. The decrease in DO increases metabolic stress in fish which, subsequently, may increase their susceptibility to disease. Lake

Sturgeon perform optimally at DO levels >6 mg/L with oxygen saturations of at least 70 to 80%.

Dissolved oxygen concentrations will vary throughout the day due to oxygen production by aquatic plants and changes in water temperature. In order to accurately depict this variation, DO measurements need to be collected frequently to ensure that daily fluctuations are not missed.

The timing of DO sample collections is important. Early in the morning, DO is usually low because aquatic plants do not produce oxygen at night and do not begin active photosynthesis until sunrise. DO concentrations usually peak in the afternoon and decline in the evening

(Huddleston 2006).

Dissolved oxygen levels were collected during the summer of 2014 during a short-term focal project comparing water quality parameters in “core use” areas in the study area. When comparing the daily maximum temperatures and DO data, Fort Loudoun Reservoir had the highest DO and lowest temperatures relative to Chickamauga and Watts Bar Reservoirs. In general, DO levels in the TVA reservoirs are greatly influenced by physical factors including retention time and volume (Ruane and Hauser 1991). All of the mainstem TVA reservoirs, including this study area, have a similarly short (12 to 15 days) retention time, with weak

35 stratification except during very low flows. Chickamauga Reservoir becomes inundated with a thick layer of aquatic vegetation during the late spring and summer months. This thick vegetation could also affect the locally low DO levels in Chickamauga Reservoir, both by production of DO through photosynthesis in the upper photaic epilimnon, and reduction of DO by decaying vegetation in deeper areas. The vegetation is so thick at times that it may even inhibit movement and feeding of Lake Sturgeon which may also explain to why two tagged fish in Chickamauga Reservoir have moved out of the area.

Chapter 5

Summary and Recommendations

Lake Sturgeon are considered a species of special concern by the USFWS, a vulnerable species by the American Fisheries Society in states where they occur, and a threatened species in

Tennessee. Lake Sturgeon have been reintroduced into the Upper Tennessee River system since

2000 but currently there are limited data on movement or habitat preferences. A multi-agency partnership was formed in 1998 to re-introduce Lake Sturgeon into the Upper Tennessee River system. Since the initial re-introductions in 2000, approximately 160,000 young-of-the-year and juvenile Lake Sturgeon have been reintroduced into the Holston and French Broad Rivers. A telemetry study of 49 Lake Sturgeon was conducted in the Upper Tennessee River system to determine dispersal and movement patterns, identify water quality characteristics of seasonally- important habitat, to compare temperature and dissolved oxygen at summer refugia areas of known sturgeon concentrations with other unused habitats and to identify, map and assess potential spawning habitats in the Upper Tennessee Basin. Results of the study are as follows:

1. There were 1,130,809 individual transmitter detections documented during this study.

2. The sex ratio of the 49 fish used in this study was approximately 5:2, 63% female, 23%

male, with 12% of unknown gender.

3. Of the 49 Lake Sturgeon outfitted with acoustic transmitters, two individuals were

tracked for a limited time, and went missing from Chickamauga Reservoir, with the last

known locations documented in March, and May 2014, respectively (two were not

detected again after release). Tagged individuals were scattered throughout the study

area with higher concentrations captured/tagged, then relocated in Fort Loudoun and

Watts Bar Reservoirs. 37

All individuals showed movement, but only a few individuals traveled away from

their original capture sites. Movements of all 49 individuals varied, with the majority of

the fish detected traveling <5 km (<3.1 miles), during the study period. There were no

notable differences in distance traveled by individuals based on location (i.e., by

reservoir). There were no noticeable diel movement patterns. There were no movements

that could be attributed to spawning migrations.

4. Water quality parameters were similar across the study area. The temperatures and DO

levels recorded were within acceptable ranges for Lake Sturgeon, with the exception of

the summer season at Chickamauga Reservoir. On Chickamauga Reservoir, temperatures

increased and DO levels dropped dramatically during June through October.

5. “Core use” areas and other seasonally important habitats were identified based on fish

aggregations. These areas were located in bends of the river mostly below the Little

River on Fort Loudoun Reservoir, along the Paint Rock Refuge and Seven Islands area

on Watts Bar Reservoir, and centered around the confluence of the Hiwassee River

around Hiwassee Island on Chickamauga Reservoir. It is recommended that these areas

continue to be monitored and that the monitoring include water quality monitoring

specifically DO and temperature.

6. While no spawning events were recorded, potential spawning areas were located based

on water quality parameters, angler reports, and substrate composition. Potential

spawning sites were identified at rocky natural shoal areas and in rocky substrate near

dams, both on mainstem UTR and large tributaries. It is recommended that continued

monitoring include water quality monitoring, especially temperature and dissolved

oxygen.

Future Research Recommendations

Based on the results of this study, and other observations, I recommend future research into these areas and issues:

a. A study to determine prey/forage availability in seasonally-important habitat and core-use

areas would be useful in determining potential areas within the Upper Tennessee River

system where Lake Sturgeon aggregations are likely to occur. It is recommended that

underwater videography and side-scan sonar be used in analyses of these areas (Flowers

and Hightower 2015).

b. An additional study to be given consideration is one focusing on the effects of heavy

metals and sediment pollutants on Lake Sturgeon at all life stages. Given the imperiled

status of Lake Sturgeon and the history of pollution and heavy metal contamination in the

current study area, this information would be very useful in determining management

strategies for understanding future reproductive success and juvenile survival in some

areas.

c. Additional consideration should be given to studying the effects of dredging in areas

where Lake Sturgeon core use areas have been identified. Commercial vessel passage

and navigation is very prominent in this study area. Lake Sturgeon tended to concentrate

in the navigation channel regardless of flow and temperature which makes them

susceptible to entrainment during channel dredging (Boysen and Hoover 2009; Hoover et

al. 2011). Disposal of dredged sediments into deep water areas during maintenance

could also negatively impact prime sturgeon foraging and loafing habitats.

d. Understanding the importance of prey and habitat availability for juvenile and adult Lake

Sturgeon distribution in the UTR which is the southernmost extent of the species range,

has important implications for conservation management. A basic knowledge of specific

habitat conditions and macroinvertebrate densities that affect Lake Sturgeon distribution

could be used to identify areas within the UTR system where restorations efforts can be

more focused.

Therefore, a telemetry study combined with macroinvertebrate sampling and gut

content collections from reintroduced Lake Sturgeon of all life stages has important

management implications. Linking the distribution of aquatic macroinvertebrates in the

UTR to the occurrence of Lake Sturgeon would help biologists better understand the

importance of prey availability and physical habitat. Proper management of the species

likely involves river enhancements which would provide sturgeon with access to a wider

range of habitat conditions, the following: 1) natural variation in flow, 2) velocity, 3)

temperature, 4) better water quality (including DO), 5) extensive prey availability, 6)

habitat diversity, and 7) free-flowing segments of river providing suitable staging and

spawning sites. e. Further study of the potential for improvements to upstream passage at locks and dams to

provide access to likely spawning sites.

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APPENDICES

APPENDIX 1. Tables

54 Table 1. Lake Sturgeon outfitted with acoustic tags on the Tennessee River.

Year Weight Total Length Fork Length ID Capture Class (kg) Gender (cm) (cm) Code RM Fort Loudoun Reservoir 2006 1.68 U 70.5 61.5 26638 628.95 2007 1.80 F 79.3 70.4 26640 628.80 2005 1.40 F 68.0 60.0 26641 629.49 2005 5.33 F 75.5 26643 626.60 2008 1.68 F 76.0 67.5 26644 629.51 2005 2.35 F 83.0 72.0 26645 624.30 2004 1.63 F 72.7 63.4 26647 632.61 2005 5.33 F 75.5 26648 627.63 2005 1.90 F 77.7 69.2 26650 623.30 2007 9.52 F 63.2 55.4 26704 632.10 2008 1.96 F 78.0 26705 629.70 2005 1.13 F 66.4 57.3 26706 632.60 2003 1.59 M 74.2 65.8 27594 632.50 2007 1.15 F 66.0 75.5 27602 626.65 2006 1.30 U 69.4 60.2 27603 629.00 2005 1.48 M 72.0 63.8 27604 628.78 2007 1.18 F 66.5 58.0 27605 630.00 2004 2.10 F 91.0 83.0 29932 626.35 2011 2.06 F 75.5 67.5 26707 619.20 Watts Bar Reservoir 2005 2.77 F 81.2 73.5 26639 573.88 2007 2.44 F 80.8 71.4 26642 569.32 2004 3.20 F 95.0 85.0 26646 571.33 2004 2.86 M 85.2 75.0 26649 573.61 Unk 1.27 F 64.9 57.4 27595 573.68 2003 8.29 F 112.0 105.0 27596 568.90 2009 1.23 U 63.3 56.5 27600 574.55 2004 1.54 M 73.5 66.6 27601 574.84 Unk7 2.11 M 88.5 78.5 28528 568.80 2001 4.52 M 94.0 85.7 28529 573.28 2001 2.24 M 105.0 92.5 28530 570.51 2002 2.76 F 109.0 97.5 28531 570.50 2003 3.47 F 89.7 80.9 28532 572.76 2004 6.12 F 110.0 100.0 28533 568.30 2004 2.94 F 85.0 76.8 29933 574.46 55 Table 1. Lake Sturgeon outfitted with acoustic tags on the Tennessee River (continued).

Year Weight Total Length Fork Length ID Capture Class (kg) Gender (cm) (cm) Code RM Unk 2.70 M 81.0 72.0 29934 569.23 2002 6.00 F 107.0 95.0 29938 572.31 Unk 3.99 F 102.0 93.0 29939 574.00 2002 6.64 F 108.0 97.0 29940 569.60 2006 4.40 M 103.0 92.0 29942 572.57 Chickamauga Reservoir 2004 6.17 F 109.0 98.0 29936 501.75 2005 8.57 F 116.0 105.0 29937 501.89 2004 5.00 M 103.0 93.0 24521 499.80 Unk 6.38 U 107.0 97.0 24524 489.90 2004 14.52 M 130.0 123.2 15954 494.15 2002 5.70 F 113.0 108.0 15956 503.70 2003 5.30 M 107.0 99.1 15957 503.50

56 Table 2. Reintroduced Lake Sturgeon scute removal protocol (SELSWG 2014).

Year Class Mark 1999-2000 Coded Wire Tag (CWT) only 2001 2nd scute removed on right side, CWT 2002 3rd scute removed on left side 2003 4th scute removed on right side 2004 5th scute removed on left side 2004 5th and 6th scutes removed on left side (Yearling and older) 2005 6th scute removed on right side 2005 6th and 7th scutes removed on right side (Yearling and older) 2006 7th scute removed on left side 2007 8th scute removed on the right side 2008 1st and 2nd scutes removed on the left side 2009 1st and 2nd scutes removed on the right side 2010 3rd and 4th scutes removed on the left side 2011 5th and 6th scutes removed on the right side 2012 7th and 8th scutes removed on the left side 2013 7th and 8th scutes removed on the right side 2014 3rd and 4th scutes removed on the right side 2015 5th and 6th scutes removed on the left side 2016 1st and 2nd scutes removed on the left side 2017 1st and 2nd scutes removed on the right side 2018 3rd and 4th scutes removed on the left side 2019 5th and 6th scutes removed on the right side 2020 7th and 8th scutes removed on the left side

57 Table 3. Fixed array of acoustic receivers were located at dams on the Upper Tennessee River and at large tributaries.

RM Site Type Longitude Latitude TN RM 423.6 Nickajack Dam - downstream Lock & Dam -85.63388 35.01036 TN RM 466.0 Chickamauga Dam - Lock & Dam -85.27843 35.06135 downstream TN RM 497.0 Tennessee River Tag site -85.03171 35.38435 TN RM 499.0 Tennessee River Tag site -85.01599 35.40742 TN RM 502.0 Chickamauga Reservoir - reservoir -84.98001 35.44022 Hiwassee Island TN RM 505.0 Tennessee River Tag site -84.96904 35.46277 HW RM 5.0 Lower Hiwassee River Large tributary -84.94299 35.38092 TN RM 522 Watts Bar Dam - downstream Lock & Dam -84.80921 35.54486 TN RM 530 Watts Bar Dam - upstream Lock & Dam -84.78336 35.62458 TN RM 565 Watts Bar Marney Bluff Tag site, known -84.55467 35.83306 concentration TN RM 569 Watts Bar Reservoir - lower Tag site, known -84.53896 35.84441 Paint Rock Refuge concentration CL RM 1.5 Lower Clinch RM 1.5 Large tributary -84.52944 35.88167 CL RM 3.75 Clinch River RM 3.75 Large tributary -84.50182 35.89222 CL RM 13.0 Melton Hill Dam - downstream Lock & Dam -84.29863 35.88441 EM RM 5.0 Emory River at Little Emory R. Large tributary -84.48170 35.93773 TN RM 576 Watts Bar Reservoir - upper Tag site, known -84.45755 35.78735 Paint Rock Refuge concentration TN RM 599 Fort Loudoun Dam - Lock & Dam -84.27437 35.75681 downstream TN RM 610.5 Fort Loudoun Dam - upstream Lock & Dam -84.16215 35.79819 TN RM 633.6 Loudoun Reservoir Tag site, known -84.02069 35.87647 concentration TN RM 630 Fort Loudoun Reservoir - Tag site, known -84.00824 35.84780 Pellissippi Bridge concentration TN RM 651 Fort Loudoun Reservoir - Forks Large tributary -83.86354 35.95731 of the River LT RM 01 Tellico Canal Canal -84.24735 35.77688 LT RM 31 Little Tennessee River - Terminal Dam -84.08339 35.54170 Chilhowee Dam FB RM 16 French Broad River - Seven Stocking Site -83.68691 35.93704 Islands FB RM 32 French Broad - Douglas Dam Terminal Dam -83.54218 35.95984 LPG RM 0.5 Little Pigeon River Large tributary -83.58935 35.93179 HR RM 32 Holston River - Nance Ferry Stocking Site -83.63513 36.12586 HR RM 52 Cherokee Dam - downstream Terminal Dam -83.50770 36.16852 58 Table 4. Potential spawning sites – rocky natural shoal areas and rocky substrate near dams.

Mainstem Dams Douglas Dam tailrace Cherokee Dam tailrace Fort Loudoun Dam tailrace Watts Bar Dam tailrace Chickamauga Dam tailrace Nickajack Dam tailrace

Tributary Dams and Shoals Holston River at Nance Ferry Clinch River at Melton Hill Dam tailrace Little Tennessee River at Chilhowee Dam tailrace Tellico River at Tellico Plains Little Pigeon at Blalocks Dam Little River at Rockford Dam Emory River at Oakdale

59 Table 5. Minimum home range size of Lake Sturgeon outfitted with acoustic tags at Fort Loudoun Reservoir on the Tennessee River, as determined by detection at fixed array of acoustic receivers. Acoustic receivers were located at dams, large tributaries, and known Lake Sturgeon habitats, to detect gross movement patterns, not at regular intervals.

Total Fork Weight Gender Length Length ID Capture Tributary Home Year Class (kg) (cm) (cm) Code RM Upstream Downstream miles Range Fort Loudoun Reservoir 2006 1.68 U 70.5 61.5 26638 628.95 629.0 565.0 64.0 2007 1.80 F 79.3 70.4 26640 628.80 633.6 630.0 3.6 2005 1.40 F 68.0 60.0 26641 629.49 633.6 630.0 3.6 2005 5.33 F 75.5 26643 626.60 630.0 626.0 4.0 2008 1.68 F 76.0 67.5 26644 629.51 633.6 610.5 23.1 2005 2.35 F 83.0 72.0 26645 624.30 630.0 610.5 19.5 2004 1.63 F 72.7 63.4 26647 632.61 633.6 630.0 3.6 2005 5.33 F 75.5 26648 627.63 633.6 630.0 3.6 2005 1.90 F 77.7 69.2 26650 623.30 651.0 630.0 21.0 2007 0.95 F 63.2 55.4 26704 632.10 633.6 630.0 3.6 2008 1.96 F 78.0 26705 629.70 651.0 630.0 21.0 2005 1.13 F 66.4 57.3 26706 632.60 633.6 630.0 3.6 2003 1.59 M 74.2 65.8 27594 632.50 633.6 630.0 3.6 2007 1.15 F 66.0 75.5 27602 626.65 633.6 630.0 3.6 2006 1.30 U 69.4 60.2 27603 629.00 633.6 630.0 3.6 2005 1.48 M 72.0 63.8 27604 628.78 633.6 630.0 3.6 2007 1.18 F 66.5 58.0 27605 630.00 633.6 630.0 3.6 2004 2.10 F 91.0 83.0 29932 626.35 633.6 630.0 3.6 2011 2.06 F 75.5 67.5 26707 619.20 630.0 610.5 19.5

60 Table 6. Minimum home range size of Lake Sturgeon outfitted with acoustic tags at Watts Bar Reservoir on the Tennessee River, as determined by detection at fixed array of acoustic receivers. Acoustic receivers were located at dams, large tributaries, and known Lake Sturgeon habitats, to detect gross movement patterns, not at regular intervals. Total Fork Weight Gender Length Length ID Capture Tributary Home Year Class (kg) (cm) (cm) Code RM Upstream Downstream miles Range Watts Bar 2005 2.77 F 81.2 73.5 26639 573.88 576.00 569.0 7.0 2007 2.44 F 80.8 71.4 26642 569.32 569.0 565.0 1.5 5.5 2004 3.20 F 95.0 85.0 26646 571.33 576.0 565.0 1.5 12.5 2004 2.86 M 85.2 75.0 26649 573.61 576.0 569.0 7.0 Unk 1.27 F 64.9 57.4 27595 573.68 569.0 565.0 1.5 5.5 2003 8.29 F 112.0 105 27596 568.90 565.0 569.0 9.5 5.5 2009 1.23 U 63.3 56.5 27600 574.55 576.0 574.5 1.5 2004 1.54 M 73.5 66.6 27601 574.84 569.0 565.0 1.5 5.5 Unk7 2.11 M 88.5 78.5 28528 568.80 576.0 565.0 1.5 12.5 2001 4.52 M 94.0 85.7 28529 573.28 576.0 530.0 1.5 47.5 2001 2.24 M 105 92.5 28530 570.51 576.0 565.0 1.5 12.5 2002 2.76 F 109 97.5 28531 570.50 569.0 565.0 4.0 2003 3.45 F 89.7 80.9 28532 572.76 576.0 572.6 3.4 2004 6.16 F 110.0 100 28533 568.30 576.0 565.0 1.5 12.5 2004 2.99 F 85.0 76.8 29933 574.46 576.0 565.0 11.0 Unk 2.70 M 81.0 72.0 29934 569.23 576.0 569.2 6.8 2005 3.63 F 93.6 82.8 29935 574.54 576.0 569.0 7.0 2002 6.00 F 107.0 95.0 29938 572.31 576.0 565.0 1.5 12.5 Unk 3.99 F 102.4 92.5 29939 574.00 576.0 574.0 2.0 2002 6.64 F 107.5 97.0 29940 569.60 576.0 565.0 11.0 2006 4.40 M 103.0 92.0 29942 572.57 576.0 565.0 1.5 12.5 61 Table 7. Minimum home range size of Lake Sturgeon outfitted with acoustic tags at Chickamauga Reservoir on the Tennessee River, as determined by detection at fixed array of acoustic receivers. Acoustic receivers were located at dams, large tributaries, and known Lake Sturgeon habitats, to detect gross movement patterns, not at regular intervals.

Total Fork Weight Gender Length Length ID Capture Tributary Home Year Class (kg) (cm) (cm) Code RM Upstream Downstream miles Range Chickamauga 2004 6.17 F 109.0 98.0 29936 501.75 502.0 501.8 5.0 5.3 2005 8.57 F 116.0 105.0 29937 501.89 502.0 497.0 5.0 10.0 2004 5.00 M 103.0 93.0 24521 499.80 501.9 497.0 5.0 Unk 6.38 U 107.0 97.0 24524 489.90 505.0 497.0 8.0 2004 14.52 M 129.5 123.2 15954 494.15 505.0 497.0 8.0 2002 5.70 F 113.0 108.0 15956 503.70 505.0 497.0 8.0 2003 5.30 M 106.6 99.1 15957 503.50 505.0 499.0 6.0

62 Table 8. Overall minimum home range size of Lake Sturgeon outfitted with acoustic tags on the Tennessee River, as determined by detection at fixed array of acoustic receivers. Acoustic receivers were located at dams, large tributaries, and known Lake Sturgeon habitats, to detect gross movement patterns, not at regular intervals.

Mean Distance in River Miles Sample Size Reservoir ( rm) (n) Fort Loudoun 8.41 18 Watts Bar 9.77 21 Chickamauga 7.18 7 Overall 8.84 46

63 Table 9. Average daily water temperatures 2013-2015 measured at Fort Loudon Reservoir, Watts Bar Reservoir, and Chickamauga Reservoir on the Upper Tennessee River compared to Fox River , Wisconsin. The number of days that were within the biologically meaningful temperature ranges is reported as a percent of the study period.

Temperature Fort Loudon Watts Bar Chickamauga Fox River Range (°Celsius) Reservoir Reservoir Reservoir Wisconsin

Biologically Meaningful Temperature Ranges 12-17 12.01% 13.32% 26.09% 8.41% <12 36.46% 39.74% 13.86% 55.46% >17 51.53% 46.94% 60.04% 35.70% >28 0.00% 0.00% 25.98% 0.55% >30 0.00% 0.00% 13.54% 0.55%

Statistics Mean 16.03 15.06 20.76 10.87 Median 17.47 15.73 21.03 7.90 Min 3.59 1.66 7.65 0.00 Max 26.41 26.76 32.22 30.20 std dev 6.75 7.74 7.31 9.41

APPENDIX 2. Figures.

Figure 1. Morphology of Lake Sturgeon (Courtesy of the University of Michigan).

66 Figure 2. Map of the study area which includes the Upper Tennessee River (RM 427 – 632), Clinch River (RM 0 - 5), Hiwassee River (RM 0 - Holston River (RM 0 – 52.2), and French Broad River (RM 0 – 32.3).

67 Figure 3. Photograph depicting eggs collected from wild brood stock on the Wolf River in Wisconsin with collaboration between U.S. Fish and Wildlife Service, the Warm Springs National Fish Hatchery, and the Wisconsin Department of Natural Resources.

68 Figure 4. Map of Lake Sturgeon stocking locations for the Upper Tennessee River system. Lake Sturgeon have been stocked as young-of-the year and fingerlings at Nance’s Ferry on the Holston River, and at Seven Islands on the French Broad Rivers. 69 Figure 5. Photograph of scute removal on fingerling Lake Sturgeon at the hatchery prior to stocking in the Upper Tennessee River System. 70

Figure 6. Photograph of wild-caught hatchery-reared Lake Sturgeon in the Upper Tennessee River system on surgery table with MS-222 running over gills during surgery to implant acoustic transmitter.

Figure 7. Photograph of wild-caught hatchery-reared Lake Sturgeon in the Upper Tennessee River study area on surgery table with incision open to determine gender and sexual maturation by examination of gonads before implantation of acoustic transmitter.

Figure 8. Photograph of VEMCO acoustic transmitters in a 70% alcohol bath ready to be surgically implanted at a 45° angle into a wild-caught hatchery reared Lake Sturgeon in the Upper Tennessee River system. 73

Figure 9. Photograph of sutures after implantation of acoustic transmitter into wild-caught hatchery-reared Lake Sturgeon in the Upper Tennessee River system using a single synthetic, non- absorbable, ETHICON (Ethibond Extra Purple Braided, 4.0 Metric) suture with an interrupted criss-cross suture and a double knot. 74 Figure 10. Photograph of wild-caught hatchery reared Lake Sturgeon in the Upper Tennessee River system in recovery tank after surgery to implant acoustic transmitter before re- release into river. 75

Figure 11. Manual tracking reintroduced Lake Sturgeon movements by boat using VEMCO VR100 with directional and omni- directional hydrophones in the Upper Tennessee River system.

Figure 12. Map of fixed acoustic receiver stations in the Upper Tennessee River system extending from the Tennessee River downstream of Nickajack Dam (RM 424.4) to the first dam at the Clinch, French Broad, and Holston Rivers.

Figure 13. Side Scan Sonar image of Lake Sturgeon in Watts Bar Reservoir taken with boat mounted Humminbird sonar. 78

Figure 14. Map of water quality sites in the Upper Tennessee River study area. 79

Figure 15. Photograph of Hydrolab sonde in Watts Bar Reservoir on U.S. Fish and Wildlife Service research buoy at RM 573.4. 80

Figure 16. Photograph of Hydrolab sonde on U.S. Fish and Wildlife research buoy in Fort Loudoun Reservoir at RM 628.7.

Figure 17. Photograph of Hydrolab sonde on U.S. Fish and Wildlife research buoy in Chickamauga Reservoir at RM 487.3. 82 Figure 18. Map of “core use” areas in the Upper Tennessee River study area based on current movement data. 83 Figure 19. Map illustrating four possibilities for fish passage used by Lake Sturgeon #26638 at the time that it moved through the Fort Loudoun dam in 2014.

Figure 20. Floating bar chart describing lockages for Fort Loudoun Dam from March 20, 2014 to April 4, 2014 during the time in which Lake Sturgeon #26638 passed through the dam.

85 Figure 21. Photograph of Lake Sturgeon below Chickamauga Dam April 20, 2015 courtesy of Dan Walker University of Tennessee.

86 Figure 22. Lake sturgeon outfitted with acoustic transmitters that also reported depth used various depths across the winter and spring seasons. Lake Sturgeon #15954 was detected at deeper areas during January and February, while it was detected at 4 – 16 feet for extended times during late March, likely at the sand bar area across the river and just downstream of Blythe Ferry boat ramp, and/or the shallow sand bar at the west end of Hiwassee Island, both areas within the detection range of the fixed receiver at the bluff at RM 499. 87

Figure 23. Lake sturgeon outfitted with acoustic transmitters that also reported depth used various depths across the winter and spring seasons. Lake Sturgeon #15956 was detected at deeper areas during December and January, while it was detected at 6 – 16 feet regularly during April and May. 88 Figure 24. Lake sturgeon outfitted with acoustic transmitters that also reported depth used various depths across the winter and spring seasons. Lake Sturgeon #15957 was detected at deeper areas during December and January, while it was detected at 3.5 - 20 feet regularly during April and May.

RM 497 RM 499 0 10

20 30 40

Depth (feet) Depth 60 70

12/1 12/31 1/30 3/1 3/31 4/30 5/30 6/29 12/1 12/31 1/30 3/1 3/31 4/30 5/30 6/29

RM 502 RM 505 0 10

50 60

Depth (feet) Depth 70

80 12/1 12/31 1/30 3/1 3/31 4/30 5/30 6/29 12/1 12/31 1/30 3/1 3/31 4/30 5/30 6/29

Figure 25. Lake sturgeon outfitted with acoustic transmitters that also reported depth used the maximum depths in the vicinity of each fixed receiver. Based on bathymetry maps and soundings with boat-mounted sonar, the maximum depth within the listening range of the receiver at RM 497 was 55 feet, RM 499 was 48 feet, RM 502 was 71 feet, and RM 505 was 48 feet.

90 35

C) °

10 Temperature ( Temperature

0 Nov-12 Jun-13 Dec-13 Jul-14 Jan-15 Aug-15 Fort Loudoun Watts Bar Chickamauga Fox River, WI Preferred Temperature - low Preferred Temperature - high

Figure 26. Graph illustrating average daily water temperature comparisons from 2013-2015 for Fort Loudoun, Watts Bar, Chickamauga Reservoirs in Tennessee and the Fox River in Wisconsin with preferred low and high temperatures for optimum Lake Sturgeon growth. 91 Waterbody Least Squares Means Waterbody Estimate Standard Error z Value Pr > |z| Alpha Lower Upper CH 2.4955 0.01764 141.48 <.0001 0.05 2.4610 2.5301 FL 2.0329 0.02244 90.58 <.0001 0.05 1.9889 2.0768 FR 0.3128 0.05314 5.89 <.0001 0.05 0.2086 0.4169 WB 1.7115 0.02636 64.94 <.0001 0.05 1.6598 1.7631

Differences of Waterbody Least Squares Means Waterbody Waterbody Estimate Standard Error z Value Pr > |z| Alpha Lower Upper CH FL 0.4627 0.02855 16.21 <.0001 0.05 0.4067 0.5186 CH FR 2.1828 0.05599 38.98 <.0001 0.05 2.0730 2.2925 CH WB 0.7841 0.03171 24.72 <.0001 0.05 0.7219 0.8462 FL FR 1.7201 0.05769 29.82 <.0001 0.05 1.6070 1.8332 FL WB 0.3214 0.03462 9.28 <.0001 0.05 0.2536 0.3892 FR WB -1.3987 0.05932 -23.58 <.0001 0.05 -1.5150 -1.2824

Figure 27. Seasonal temperature comparisons for Watts Bar, Chickamauga, and Fort Loudoun Reservoirs and the Fox River for the winter (December 22 to March 21) 2014.

92 Waterbody Least Squares Means Waterbody Estimate Standard Error z Value Pr > |z| Alpha Lower Upper CH 3.1508 0.01246 252.96 <.0001 0.05 3.1264 3.1752 FL 2.8985 0.01413 205.12 <.0001 0.05 2.8708 2.9261 FR 2.5492 0.01692 150.67 <.0001 0.05 2.5160 2.5823 WB 2.8460 0.01451 196.20 <.0001 0.05 2.8176 2.8744

Differences of Waterbody Least Squares Means Waterbody Waterbody Estimate Standard Error z Value Pr > |z| Alpha Lower Upper CH FL 0.2523 0.01884 13.40 <.0001 0.05 0.2154 0.2892 CH FR 0.6016 0.02101 28.63 <.0001 0.05 0.5604 0.6428 CH WB 0.3048 0.01912 15.94 <.0001 0.05 0.2673 0.3422 FL FR 0.3493 0.02204 15.84 <.0001 0.05 0.3061 0.3925 FL WB 0.05244 0.02025 2.59 0.0096 0.05 0.01275 0.09213 FR WB -0.2968 0.02229 -13.32 <.0001 0.05 -0.3405 -0.2531

Figure 28. Seasonal temperature comparisons for Watts Bar, Chickamauga, and Fort Loudoun Reservoirs and the Fox River for the spring (March 22 to June 21) 2014.

93 Waterbody Least Squares Means Waterbody Estimate Standard Error z Value Pr > |z| Alpha Lower Upper CH 3.4006 0.01315 258.68 <.0001 0.05 3.3748 3.4264 FL 3.1620 0.01462 216.23 <.0001 0.05 3.1333 3.1906 FR 3.1609 0.01463 216.04 <.0001 0.05 3.1322 3.1896 WB 3.2050 0.01431 223.94 <.0001 0.05 3.1770 3.2331

Differences of Waterbody Least Squares Means Waterbody Waterbody Estimate Standard Error z Value Pr > |z| Alpha Lower Upper CH FL 0.2386 0.01966 12.13 <.0001 0.05 0.2001 0.2772 CH FR 0.2397 0.01967 12.19 <.0001 0.05 0.2011 0.2783 CH WB 0.1956 0.01943 10.06 <.0001 0.05 0.1575 0.2336

FL FR 0.001082 0.02069 0.05 0.9583 0.05 -0.03946 0.04163 FL WB -0.04306 0.02046 -2.10 0.0353 0.05 -0.08317 -0.00296 FR WB -0.04414 0.02047 -2.16 0.0310 0.05 -0.08426 -0.00403

Figure 29. Seasonal temperature comparisons for Watts Bar, Chickamauga, and Fort Loudoun Reservoirs and the Fox River for the summer (June 22 to September 21) 2014.

94 Waterbody Least Squares Means Waterbody Estimate Standard Error z Value Pr > |z| Alpha Lower Upper CH 2.9753 0.01674 177.69 <.0001 0.05 2.9425 3.0081 FL 2.8081 0.01821 154.25 <.0001 0.05 2.7724 2.8438 FR 2.0394 0.02674 76.27 <.0001 0.05 1.9870 2.0918 WB 2.7062 0.01916 141.26 <.0001 0.05 2.6686 2.7437

Differences of Waterbody Least Squares Means Waterbody Waterbody Estimate Standard Error z Value Pr > |z| Alpha Lower Upper CH FL 0.1672 0.02473 6.76 <.0001 0.05 0.1187 0.2157 CH FR 0.9360 0.03155 29.67 <.0001 0.05 0.8741 0.9978 CH WB 0.2691 0.02544 10.58 <.0001 0.05 0.2193 0.3190 FL FR 0.7687 0.03235 23.77 <.0001 0.05 0.7053 0.8321 FL WB 0.1019 0.02643 3.86 0.0001 0.05 0.05011 0.1537 FR WB -0.6668 0.03289 -20.27 <.0001 0.05 -0.7313 -0.6024

Figure 30. Seasonal temperature comparisons for Watts Bar, Chickamauga, and Fort Loudoun Reservoirs and the Fox River for the fall (September 22 to December 21) 2014.

APPENDIX 3. Detections of Lake Sturgeon with acoustic tags in Tennessee River 2013 – 2015 – Maps of re-locations.

Lake Sturgeon captured and tagged at Fort Loudoun Reservoir.

97 84°30'0"W 84°20'0"W 84°10'0"W 84°0'0"W E E E E E E E

E 36°0'0"N E E E E E E E E E E E E E E ")E E E E E E E E E E E E E E E E E E E E E E E E E E E E E ")E E E E E E E E E E E E E E EEE E E E E E E E E E ")E E E E E E E E E E E E kj ") E E E 35°50'0"N E E E ")E E E E E E E ")E E E E E E E E E E") E E E E E E E E E E E E E E E ") E E E E E E E E E E E E E

E E ID 26638 E kj Capture E F E ") E Acoustic Array 0 4

Kilometers 35°40'0"N E VR100 Miles E Bearcat 0 4 E E E

98 84°30'0"W 84°20'0"W E E E E E 911 ")E5 E E E E E 4 E 3 E E E E E E 2 E E E E 1 E E E 7 E E 3 6 E E E E E ") 4 E E 2 E 5 E E E E E E E E 1 E 5680 E EE 567 569 E ") 570 E 571 ") E 572 E 35°50'0"N 573 E

574 E 575 E 576 602 603 ")E 601 E E 577 E 604 E 582 583 600 E E E 589 E ")1 E 597 E 584 596 E 605 578 581 590 E 3 E E E E E 599 E 2 606 588 598 E E 585 E E E 579 580 E 591 595 ") 4 E E 587 E E E 586 E E 594 5 592 E E E 593 E 6 E ID 26638 7 kj Capture E F ") Acoustic Array 80 2 E VR100 Kilometers 9 Miles Bearcat 0 E 2 E

99 84°10'0"W 84°0'0"W E E 640 639 E E E 638 E E E 635 E E 637 E E 633 634 636 E E E E 632 E 631 E 617 627 628 E 618 622 E E E E 623 630 616 621 E E E E 629 619 624 626 E 615 E E E kj E 620 E 625 E 614 E 35°50'0"N 613 610 E E 611 612 609")E E 602 603 E 601 E E E 604 608 E E 600 ")1 597 E E E 605 3 E 607 599 E 2 606E 598 E E E E ") 4 E 5 E E

6 E

7 E

8 E ID 26638 9 kj Capture E F ") Acoustic Array 0 2 10 11 E Kilometers E VR100 Miles 12 E 13 Bearcat 0 2

E 35°40'0"N

100 84°30'0"W 84°20'0"W E E 2 1 E E E E 7 3 6 E E E E ") 4 E E 2 5 E E E E E 1 E

567 5680 E EE

566 E 569 E 570 E 564 565 571 E E E 572 E 35°50'0"N

573 E

574 kjE 575 E 576 E 577 582 E E 583 E 589 E 578 581 584 590 E E E E 588 E 585 579 580 E 591 E E E ID 26639 587 586 E kj Capture E F592 ") Acoustic Array 0 2 E Kilometers VR100 Miles Bearcat 0 2

101 84°0'0"W E

638 E

635 E

633 637 E 634 636 E ") E E

632 E

631 E

628 627 E 622 E E 623 630 E ")E kj 629 626 E 624 E E

625

E 35°50'0"N

ID 26640 kj Capture F ") Acoustic Array 0 1 Kilometers VR100 Miles Bearcat 0 1

102 84°0'0"W E

638 E

635 E

633 637 E 634 636 E ") E E

632 E

631 E

628 627 E 622 E E 623 630 E ")E

629 626 E kj 624 E E

625

E 35°50'0"N

ID 26641 kj Capture F ") Acoustic Array 0 1 Kilometers VR100 Miles Bearcat 0 1

103 84°0'0"W E

638 E

635 E

633 637 E 634 636 E E E

632 E

631 E

628 627 E 622 E E 623 630 E kj ")E 629 626 E 624 E E

625

E 35°50'0"N

ID 26643 kj Capture F ") Acoustic Array 0 1 Kilometers VR100 Miles Bearcat 0 1

104 84°0'0"W E

638 E

635 E

633 637 E 634 636 E ") E E

632 E

631 E

628 627 E 622 E E 623 630 E ")E

629 626 E kj 624 E E

625

E 35°50'0"N

ID 26644 kj Capture F ") Acoustic Array 0 1 Kilometers VR100 Miles Bearcat 0 1

105 84°10'0"W 84°0'0"W E

639 E

638 E 635 E 633 637 E 634 636 E E E 632 E 631 E

617 618 627 628 E E 622 E E E 623 630 616 E E E 621 ") E 629 619 626 E E 624 E 615 620 E E E kj 625 E

614 35°50'0"N E

613 610 E E 611 612 ")E E 609 E

608 E

605 E 607 E 606 E ID 26645 kj Capture F ") Acoustic Array 0 1 Kilometers VR100 Miles Bearcat 0 1

106 84°0'0"W E

638 E

635 E

633 637 E 634 636 E kj ") E E

632 E

631 E

628 627 E 622 E E 623 630 E ")E

629 626 E 624 E E

625

E 35°50'0"N

ID 26647 kj Capture F ") Acoustic Array 0 1 Kilometers VR100 Miles Bearcat 0 1

107 84°0'0"W E

638 E

635 E

633 637 E 634 636 E ") E E

632 E

631 E

628 627 E 622 E E 623 630 E kj ")E

629 626 E 624 E E

625

E 35°50'0"N

ID 26648 kj Capture F ") Acoustic Array 0 1 Kilometers VR100 Miles Bearcat 0 1

108 84°0'0"W

650 649 E 648 E 651 E E 652 ") E 647 645 E E 644 646 E E 641 E 643 642 E E 640 E

639 E

638 E 635 E 633 637 E 634 636 E ") E E 632 E 631 E 627 628 622 E E E 623 630 E ")E kj 629 624 626 E E E ID 26650 625 E kj Capture

F 35°50'0"N ") Acoustic Array 0 1 Kilometers VR100 Miles Bearcat 0 1

109 84°0'0"W E

638 E

635 E

633 637 E 634 636 E ") E E

632 kjE

631 E

628 627 E 622 E E 623 630 E ")E

629 626 E 624 E E

625

E 35°50'0"N

ID 26704 kj Capture F ") Acoustic Array 0 1 Kilometers VR100 Miles Bearcat 0 1

110 84°0'0"W

650 649 E 648 E 651 E E 652 ") E 647 645 E E 644 646 E E 641 E 643 642 E E 640 E

639 E

638 E 635 E 633 637 E 634 636 E ") E E 632 E 631 E 627 628 622 E E E 623 630 E ")E 629 kj 624 626 E E E ID 26705 625 E kj Capture

F 35°50'0"N ") Acoustic Array 0 1 Kilometers VR100 Miles Bearcat 0 1

111 84°0'0"W E

638 E

635 E

633 637 E 634 636 E kj ") E E

632 E

631 E

628 627 E 622 E E 623 630 E ")E

629 626 E 624 E E

625

E 35°50'0"N

ID 26706 kj Capture F ") Acoustic Array 0 1 Kilometers VR100 Miles Bearcat 0 1

112 84°10'0"W 84°0'0"W E

639 E

638 E 635 E 633 637 E 634 636 E E E 632 E 631 E

617 618 627 628 E E 622 E E E 623 630 616 E E E 621 ") E 629 619 626 E E 624 E 615 620 E E kj E 625 E

614 35°50'0"N E

613 610 E E 611 612 ")E E 609 E

608 E

605 E 607 E 606 E ID 26707 kj Capture F ") Acoustic Array 0 1 Kilometers VR100 Miles Bearcat 0 1

113 84°0'0"W E

638 E

635 E

633 637 E 634 636 E ") E E kj 632 E

631 E

628 627 E 622 E E 623 630 E ")E

629 626 E 624 E E

625

E 35°50'0"N

ID 27594 kj Capture F ") Acoustic Array 0 1 Kilometers VR100 Miles Bearcat 0 1

114 84°0'0"W E

638 E

635 E

633 637 E 634 636 E ") E E

632 E

631 E

628 627 E 622 E E 623 630 E kj ")E

629 626 E 624 E E

625

E 35°50'0"N

ID 27602 kj Capture F ") Acoustic Array 0 1 Kilometers VR100 Miles Bearcat 0 1

115 84°0'0"W E

638 E

635 E

633 637 E 634 636 E ") E E

632 E

631 E

628 627 E 622 E E 623 630 E ")E

629 626 Ekj 624 E E

625

E 35°50'0"N

ID 27603 kj Capture F ") Acoustic Array 0 1 Kilometers VR100 Miles Bearcat 0 1

116 84°0'0"W E

638 E

635 E

633 637 E 634 636 E ") E E

632 E

631 E

628 627 E 622 E E 623 630 E ")E

kj 629 626 E 624 E E

625

E 35°50'0"N

ID 27604 kj Capture F ") Acoustic Array 0 1 Kilometers VR100 Miles Bearcat 0 1

117 84°0'0"W E

638 E

635 E

633 637 E 634 636 E ") E E

632 E

631 E

628 627 E 622 E E 623 630 E kj")E 629 626 E 624 E E

625

E 35°50'0"N

ID 27605 kj Capture F ") Acoustic Array 0 1 Kilometers VR100 Miles Bearcat 0 1

118 84°0'0"W E

638 E

635 E

633 637 E 634 636 E ") E E

632 E

631 E

628 627 E 622 E E 623 630 E ")E

kj 629 626 E 624 E E

625

E 35°50'0"N

ID 29932 kj Capture F ") Acoustic Array 0 1 Kilometers VR100 Miles Bearcat 0 1

119

Lake Sturgeon captured and tagged at Watts Bar Reservoir.

120 84°30'0"W E E 10 E6 9 11 E 5 E E 12 E E 4 E E 3 E E E E 2 E E E 1 E E E 7 3 6 E E E ") 4 E E 2 5 E E E E 1 E

567 5680 E EE

566 E 569 562 E E 563 kj 570 E E 561 564 565 571 E E E E 572 E 35°50'0"N E 573 E E 574 E 575 E 576 ID 26642 E kj Capture 577 E 582 F ") E 583 Acoustic Array 0E 2 Kilometers VR100581 584 Miles 578 E E E Bearcat 0 2 E

121 84°30'0"W E 84°20'0"W E2 E 1 E E E E 7 3 6 E E E E ") 4 E E 2 5 E E E E ") E 1 E

567 5680 E EE

566 E 569 E 570 E 565 E 571 E kj 572 E 35°50'0"N

573 E

574 E 575 E 576 E 577 582 E E 583 E 589 E 584 596 578 581 590 E E E E E 588 E 585 579 580 E 591 595 E E ID 26646 E E 587 586 E kj Capture E 592 F ") Acoustic Array 0 1 E Kilometers 593 VR100 Miles E Bearcat 0 1

122 84°30'0"W E 84°20'0"W E2 E 1 E E E E 7 3 6 E E E E ") 4 E E 2 5 E E E E E 1 E

567 5680 E EE

566 E 569 E 570 E 565 571 E E 572 E 35°50'0"N

573 E

574 kj E 575 E 576 E 577 582 E E 583 E 589 E 584 596 578 581 590 E E E E E 588 E 585 579 580 E 591 595 E E ID 26649 E E 587 586 E kj Capture E 592 F ") Acoustic Array 0 1 E Kilometers 593 VR100 Miles E Bearcat 0 1

123 84°30'0"W E 84°20'0"W E2 E 1 E E E E 7 3 6 E E E E ") 4 E E 2 5 E E E E E 1 E

567 5680 E EE

566 E 569 E ") 570 E 565 571 ")E E 572 E 35°50'0"N

573 E

kj 574 E 575 E 576 ")E 577 582 E E 583 E 589 E 584 596 578 581 590 E E E E E 588 E 585 579 580 E 591 595 E E ID 27595 E E 587 586 E kj Capture E 592 F ") Acoustic Array 0 1 E Kilometers 593 VR100 Miles E Bearcat 0 1

124 84°30'0"W

7 8 E 10 E 6 E E 9 11 E 5 E E") 12 E E 4 E E 3 E E E E 2 E E E 1 E E 7 3 6 E E E ") 4 E E 2 kj 5 E E E ") E 1 E

567 5680 E EE

566 E 569 562 E E 563 ") E 570 E 561 564 565 571 E E ")E E 572 E 35°50'0"N 560 E 573 E 559 E ID 27596 574 kj Capture 558 E F E 575 ") Acoustic Array 0 1 E 576 E Kilometers VR100 Miles 577 582 BearcatE 0 E 1 583 E

125 84°30'0"W 84°20'0"W E E E E1 7 3 6 E E E E ") 4 E E 2 5 E E E E E 1 E

568567 0 EEE

566 E 569 E 570 E 565 571 E E 572 E 35°50'0"N

573 E

574 E kj 575 E 576 E 577 582 E E 583 E 589 E 584 596 578 581 590 E E E E E 588 E 585 579 580 E 591 595 E E E E 587 586 E ID 27600 E 592 kj Capture E F593 ") Acoustic Array 0 1 E Kilometers VR100 Miles Bearcat 0 1

126 84°30'0"W 84°20'0"W E E 2 1 E E E E 7 3 6 E E E E ") 4 E E 2 5 E E E E ") E 1 E

567 5680 E EE

566 E 569 E ") 570 E 564 565 571 E ")E E 572 E 35°50'0"N

573 E

574 kjE 575 E 576 E 577 582 E E 583 E 589 E 578 581 584 590 E E E E 588 E 585 579 580 E 591 E E E ID 27601 587 586 E kj Capture E F592 ") Acoustic Array 0 1 E Kilometers VR100 Miles Bearcat 0 1

127 84°30'0"W 84°20'0"W E E 2 1 E E E E 7 3 6 E E E E ") 4 E E 2 kj 5 E E E E ") E 1 E

567 5680 E EE

566 E 569 E ") 570 E 564 565 571 E ")E E 572 E 35°50'0"N

573 E

574 E 575 E 576 ")E 577 582 E E 583 E 589 E 578 581 584 590 E E E E 588 E 585 579 580 E 591 E E E ID 28528 587 586 E kj Capture E F592 ") Acoustic Array 0 2 E Kilometers VR100 Miles Bearcat 0 2

128 84°40'0"W 84°30'0"W E E E 2 E43 5 76 E ") E 1 E 5675680 E EE 566 E 569 562 563 ")E E E 570 561 564 565 E 571 E E ")E E 572 560 E 553 E 35°50'0"N E 554 E 573 E 552 559 E 555 E kj E 574 558 E 575 551 556 E E 576 E E 557 ")E 550 E 577 E E 549 578 E E 545 546 548 E E E 579 547 E 544 E E E 543 E 542 E 541 536 E E 537 540 535 E E E 538 539 534 E E E

533 E ID 28529

532 35°40'0"N E kj Capture F 531 ") Acoustic Array 0 2 E Kilometers 530 VR100 ")E Miles Bearcat 0 2 E

129 84°30'0"W 84°20'0"W E E 2 1 E E E E 7 3 6 E E E E ") 4 E E 2 5 E E E E ") E 1 E

567 5680 E EE

566 E 569 E ") 570 E 564 565 571 E ")E kj E 572 E 35°50'0"N

573 E

574 E 575 E 576 ")E 577 582 E E 583 E 589 E 578 581 584 590 E E E E 588 E 585 579 580 E 591 E E E ID 28530 587 586 E kj Capture E F592 ") Acoustic Array 0 2 E Kilometers VR100 Miles Bearcat 0 2

130 84°30'0"W 84°20'0"W E E 2 1 E E E E 7 3 6 E E E E 4 E E 2 5 E E E E E 1 E

567 5680 E EE

566 E 569 E ") 570 E 564 565 571 E ")E kj E 572 E 35°50'0"N

573 E

574 E 575 E 576 E 577 582 E E 583 E 589 E 578 581 584 590 E E E E 588 E 585 579 580 E 591 E E E ID 28531 587 586 E kj Capture E F592 ") Acoustic Array 0 2 E Kilometers VR100 Miles Bearcat 0 2

131 84°30'0"W 84°20'0"W E E 2 1 E E E E 7 3 6 E E E E 4 E E 2 5 E E E E E 1 E

567 5680 E EE

566 E 569 E 570 E 564 565 571 E E E 572 E 35°50'0"N

573 E

574 kjE 575 E 576 ")E 577 582 E E 583 E 589 E 578 581 584 590 E E E E 588 E 585 579 580 E 591 E E E ID 28532 587 586 E kj Capture E F592 ") Acoustic Array 0 2 E Kilometers VR100 Miles Bearcat 0 2

132 84°30'0"W 84°20'0"W E E 2 1 E E E E 7 3 6 E E E kjE ") 4 E E 2 5 E E E E ") E 1 E

567 5680 E EE

566 E 569 E ") 570 E 564 565 571 E ")E E 572 E 35°50'0"N

573 E

574 E 575 E 576 ")E 577 582 E E 583 E 589 E 578 581 584 590 E E E E 588 E 585 579 580 E 591 E E E ID 28533 587 586 E kj Capture E F592 ") Acoustic Array 0 2 E Kilometers VR100 Miles Bearcat 0 2

133 84°30'0"W 84°20'0"W E E 2 1 E E E E 7 3 6 E E E E 4 E E 2 5 E E E E E 1 E

567 5680 E EE

566 E 569 E ") 570 E 564 565 571 E ")E E 572 E 35°50'0"N

573 E

574 E kj 575 E 576 ")E 577 582 E E 583 E 589 E 578 581 584 590 E E E E 588 E 585 579 580 E 591 E E E ID 29933 587 586 E kj Capture E F592 ") Acoustic Array 0 2 E Kilometers VR100 Miles Bearcat 0 2

134 84°30'0"W 84°20'0"W 1 E

567 5680 E EE

566 E 569 E kj 570 E 565 571 E E 572 E 35°50'0"N

573 E

574 E 575 E 576 ")E 577 582 E E 583 E 589 E 584 596 578 581 590 E E E E E 588 E 585 579 580 E 591 595 E E E E 587 586 E E 594 592 E E 593 E ID 29934 kj Capture F ") Acoustic Array 0 2 Kilometers VR100 Miles Bearcat 0 2

135 84°30'0"W 84°20'0"W E E 2 1 E E E E 7 3 6 E E E E 4 E E 2 5 E E E E E 1 E

567 5680 E EE

566 E 569 E ") 570 E 564 565 571 E ")E E 572 E 35°50'0"N

573 E

574 E kj 575 E 576 ")E 577 582 E E 583 E 589 E 578 581 584 590 E E E E 588 E 585 579 580 E 591 E E E ID 29935 587 586 E kj Capture E F592 ") Acoustic Array0 2 E Kilometers VR100 Miles Bearcat 0 2

136 84°30'0"W 84°20'0"W E E E E 12 3 E4 E E E E E 2 E E E E 1 E E E 7 E E 3 6 E E E E E E ") 4 E 2 E 5 E E E E E E E E 1 E 5675680 E EE

566 569 E E ") 570 564 565 E 571 E ")E E 572 E 35°50'0"N kj573 E

574 E 575 E 576 602 603 ")E 601 E E 577 E 604 E 582 583 600 E E E 589 E 1 E 597 E 584 596 E 578 581 590 E 3 E E E E E 2 588 598 599 E 585 E E E 579 580 E 591 595 ") 4 E E 587 E E E 586 E E 594 5 592 E E E 593 E 6 E

7 ID 29938 E kj 8 Capture E F ") Acoustic Array 0 2 9 E VR100 Kilometers Miles10 11 E Bearcat 0 E 2 E 35°40'0"N

137 84°30'0"W 84°20'0"W E E 2 1 E E E E 7 3 6 E E E E 4 E E 2 5 E E E E E 1 E

567 5680 E EE

566 E 569 E 570 E 564 565 571 E E E 572 E 35°50'0"N

573 E

574 E kj 575 E 576 ")E 577 582 E E 583 E 589 E 578 581 584 590 E E E E 588 E 585 579 580 E 591 E E E ID 29939 587 586 E kj Capture E F592 ") Acoustic Array 0 2 E Kilometers VR100 Miles Bearcat 0 2

138 84°30'0"W 84°20'0"W E E 2 1 E E E E 7 3 6 E E E E 4 E E 2 5 E E E E E 1 E

567 5680 E EE

566 E 569 E ") kj 570 E 564 565 571 E ")E E 572 E 35°50'0"N

573 E

574 E 575 E 576 ")E 577 582 E E 583 E 589 E 578 581 584 590 E E E E 588 E 585 579 580 E 591 E E E ID 29940 587 586 E kj Capture E F592 ") Acoustic Array 0 2 E Kilometers VR100 Miles Bearcat 0 2

139 84°30'0"W E E 1 E E E 7 3 6 E E E ") 4 E E 2 5 E E E E 1 E

567 5680 E EE

566 E 569 E 562 ") E 563 570 E E 564 565 571 E E ")E E 572 E 35°50'0"N kj 573 E

574 E 575 E 576 ")E 577 582 E E 583 E

578 581 584 E E E 588 E 585 579 580 E E E ID 29942 586 E kj Capture F ") Acoustic Array 0 2 Kilometers VR100 Miles Bearcat 0 2

140

Lake Sturgeon captured and tagged at Chickamauga Reservoir.

141 85°0'0"W 511 504 E E 505 ")E 503 510 E E 506 E 509 502 E E ") 507 508 !U E E

501 E

500 E 0 1 2 499E E E E") 498 3 E E 4 E 497 ")E 5 !UE

496 6 E E 7 E 8 E 9 495 E E 10 kj E 11 494 E E E 493 E 35°20'0"N 492 E ID 15954 kj Capture 491 F E ") Acoustic Array 0 2 VR100 Kilometers 490 Miles E Bearcat 0 2

142 85°0'0"W 511 504 E E 505 ")E 503 510 E E 506 E 509 502 E E ") 507 508 E E

501 E

500 E 0 1 2 499E E E E") 498 3 E E 4 E 497 ")E 5 E

496 6 E E 7 E 8 E 9 495 E E 10 E 11 494 E E E 493 E 35°20'0"N 492 E ID 15956 kj Capture 491 F E ") Acoustic Array 0 2 VR100 Kilometers 490 Miles E Bearcat 0 2

143 85°0'0"W 511 504 E E kj 505 ")E 503 510 E E 506 E 509 502 E E ") 507 508 E E

501 E

500 E 0 1 2 499E E E E") 498 3 E E 4 E 497 E 5 E

496 6 E E 7 E 8 E 9 495 E E 10 E 11 494 E E E 493 E 35°20'0"N 492 E ID 15957 kj Capture 491 F E ") Acoustic Array 0 2 VR100 Kilometers 490 Miles E Bearcat 0 2

144 85°0'0"W 511 504 E E 505 ")E 503 510 E E 506 E 509 502 E E ") 507 508 E E

501 E

500 E 0kj 1 2 499E E E E") 498 3 E E 4 E 497 ")E 5 E

496 6 E E 7 E 8 E 9 495 E E 10 E 11 494 E E E 493 E 35°20'0"N 492 E ID 24521 kj Capture 491 F E ") Acoustic Array 0 2 VR100 Kilometers 490 Miles E Bearcat 0 2

145 85°0'0"W E E504 511 505 ")E 503 510 E E 506 E 509 502 E E ") 507 508 E E

501 E

500 E 0 1 2 499E E E E") 498 3 E E 4 E 497 ")E 5 E

496 6 E E 7 E 8 E 495 E

494 E

493 E 35°20'0"N 492 E ID 24524 491 kj Capture E F ") Acoustic Array 0 2 490 Kilometers kjE VR100 Miles Bearcat 489 0 2 E

146 85°0'0"W 511 504 E E 505 E 503 510 E E 506 E 509 502 E E ") 507 508 kj E E 501 E

500 E 0 1 2 499E E E E

498 3 E E 4 E 497 E 5 ")E

496 6 E E 7 E 8 E 9 495 E E 10 E 11 494 E E E 493 E 35°20'0"N 492 E ID 29936 kj Capture 491 F E ") Acoustic Array 0 2 VR100 Kilometers 490 Miles E Bearcat 0 2

147 85°0'0"W 511 504 E E 505 ")E 503 510 E E 506 E 509 502 E E ") 507 508 kj E E

501 E

500 E 0 1 2 499E E E E") 498 3 E E 4 E 497 ")E 5 ")E

496 6 E E 7 E 8 E 9 495 E E 10 E 11 494 E E E 493 E 35°20'0"N 492 E ID 29937 kj Capture 491 F E ") Acoustic Array 0 2 VR100 Kilometers 490 Miles E Bearcat 0 2

148 Vita

Christina Grace Clickner was born June 2, 1980, in Hialeah, Florida. At age 9 she moved with her family to Lenoir City, Tennessee where she attended elementary school. During her sixth grade year her family moved to Kingston, Tennessee, where she attended middle and high school graduating from Roane County High School with honors. After several years of being a mother to four children with ages ranging from 3 to 16 years, she received her Bachelor of

Science degree in Forestry, Wildlife and Fisheries Management from the University of

Tennessee in 2013. Christina received her Masters of Science degree in Wildlife and Fisheries

Science from the University of Tennessee in December 2015.

149