AN ABSTRACT OF A THESIS

SPAWNING AND NURSERY HABITAT OF WILD MUSKELLUNGE

AND FATE OF STOCKED MUSKELLUNGE

IN MIDDLE TENNESSEE RIVERS

Lila Hammond Warren

Master of Science in Biology

Southern U.S. Muskellunge Esox masquinongy populations have not been well

studied; the most recently published study on river Muskellunge in Tennessee dates to

1959. Muskellunge were nearly extirpated from the state in the 1970s mainly due to

pollution by coal mining. A Muskellunge stocking program began in 1976 in the upper

Caney Fork River and its tributaries as part of a statewide effort by the Tennessee

Wildlife Resources Agency (TWRA) to preserve the species in the Cumberland River

drainage. The upper Caney Fork River system now supports a trophy fishery, with some

fish caught in recent years exceeding 1.3 m total length (TL). Objectives of this study

were to identify and describe spawning and nursery habitats of wild Muskellunge in the

upper Caney Fork River system, assess the extent of natural reproduction and growth of

wild juvenile Muskellunge in the upper Caney Fork River system, and describe mortality,

dispersal, and habitat use by recently stocked Muskellunge. Spawning areas of wild

Muskellunge were located using traditional ecological knowledge from anglers and

fishing guides as well as boat electrofishing surveys with TWRA biologists. To

determine whether natural reproduction was occurring and to locate nursery areas,

seining, backpack electrofishing, and boat electrofishing gears were used to sample age-0

fish. Post-stocking dispersal, mortality, and habitat use by stocked fingerling

Muskellunge were evaluated using radio telemetry. Vegetative cover and finer substrates

were important habitat characteristics in spawning and nursery habitats of wild

Muskellunge. Natural reproduction occurred in the main stem Caney Fork River and

three of four tributaries (Collins River, Calfkiller River, Cane Creek), but not in the

Rocky River. Wild age-0 Muskellunge reached maximum lengths of up to 399 mm TL

by October 2012. Stocked Muskellunge suffered high interval mortality (43.4-98.3%)

over 56 d, and the stocked cohort that dispersed the most suffered the highest mortality.

Predation by Great Blue Heron Ardea herodias and River Otter Lontra canadensis was a

source of mortality for stocked fish. Percent vegetation coverage was low in habitats used

by stocked Muskellunge compared to nursery areas of wild age-0 fish. Because natural

reproduction occurs in the system and post-stocking mortality was high, TWRA might be

justified in discontinuing Muskellunge stocking and continuing annual boat-

electrofishing surveys. Close monitoring of the Muskellunge fishery in the upper Caney

Fork River system should indicate within a few years whether it is self-sustaining.

SPAWNING AND NURSERY HABITAT OF WILD MUSKELLUNGE

AND FATE OF STOCKED MUSKELLUNGE

IN MIDDLE TENNESSEE RIVERS

____________________

A Thesis

Presented to

the Faculty of the Graduate School

Tennessee Technological University

by

Lila Hammond Warren

____________________

In Partial Fulfillment

of the Requirements of the Degree

MASTER OF SCIENCE

Biology

____________________

August 2013

ii

CERTIFICATE OF APPROVAL OF THESIS

SPAWNING AND NURSERY HABITAT OF MUSKELLUNGE

AND FATE OF STOCKED MUSKELLUNGE

IN MIDDLE TENNESSEE RIVERS

by

Lila Hammond Warren

Graduate Advisory Committee:

__________________________________ _______________

Phillip W. Bettoli, Chairperson Date

__________________________________ _______________

S. Bradford Cook Date

__________________________________ _______________

Michael J. Redding Date

Approved for the Faculty:

______________________________

Francis Otuonye

Associate Vice President

for Research and Graduate Studies

________________

Date

iii

ACKNOWLEDGEMENTS

Funding and support for this research was provided by the Tennessee Wildlife

Resources Agency (TWRA), the Center for the Management, Utilization, and Protection

of Water Resources at Tennessee Technological University, and the USGS Tennessee

Cooperative Fishery Research Unit at Tennessee Technological University.

I would like to thank my advisor, Dr. Phillip Bettoli, for his mentorship and

guidance. I appreciate the support of my graduate committee members, Dr. S. Bradford

Cook and Dr. J. Michael Redding. To Jack Swearingen, Jason Hennegar, Mark Thurman,

Scotty Webb, and Travis Scott from TWRA’s Region III, thank you for the adventures

and sense of humor in the field. Thank you to Mike “Stump” Smith, Ernie Poore, Bobby

Stooksbury, and Marilyn Davis at TWRA’s Eagle Bend Hatchery for taking care of the

Muskies before their journey to the Collins River.

Thank you to my fellow graduate students and coworkers at the Tennessee

Cooperative Fishery Research Unit for your help with field work as well as your

friendship, encouragement, and all the fun we had. Thanks to my professors in the

Biology department from whom I learned so much.

Mom and Dad, you raised me to be the passionate outdoorswoman and

conservationist I am today. I am so grateful for your support starting with trips to Secret

Pond and Goose Creek to catch bluegills with a cane pole all the way through to

completing my master’s degree. Thank you for your encouragement and positive attitudes

throughout.

iv

TABLE OF CONTENTS

Page

LIST OF TABLES ...............................................................................................................v

LIST OF FIGURES ........................................................................................................... vi

Chapter

1. INTRODUCTION ...................................................................................................1

2. STUDY AREA ........................................................................................................8

3. METHODS ............................................................................................................10

Spawning and Nursery Habitat of Wild Muskellunge .....................................10

Natural Reproduction and Growth of Wild Muskellunge................................11

Mortality, Dispersal, and Habitat Use of Stocked Muskellunge .....................13

4. RESULTS ..............................................................................................................18

Spawning and Nursery Habitat of Wild Muskellunge .......................................18

Natural Reproduction and Growth of Wild Muskellunge ..................................19

Mortality, Dispersal, and Habitat Use of Stocked Muskellunge .......................... 20

5. DISCUSSION ........................................................................................................22

Spawning and Nursery Habitat of Wild Muskellunge .......................................22

Natural Reproduction and Growth of Wild Muskellunge ..................................26

Mortality, Dispersal, and Habitat Use of Stocked Muskellunge .......................... 27

6. MANAGEMENT IMPLICATIONS .....................................................................33

REFERENCES ..................................................................................................................36

TABLES ............................................................................................................................45

FIGURES ...........................................................................................................................50

APPENDIX ........................................................................................................................58

Species and counts of fishes collected by seine haul during summer 2012

sampling in the upper Caney Fork River system. The number of seine hauls

in each river is in parentheses next to the river name ............................................59

VITA ..................................................................................................................................60

v

LIST OF TABLES

Table Page

1. Records of Muskellunge stocked into the upper Caney Fork River system,

Tennessee ....................................................................................................................46

2. Summary of habitat parameters measured at spawning and nursery areas for

wild and stocked Muskellunge in the upper Caney Fork River system in

Tennessee in 2012, including the number of sites (N), mean depth, mean

dissolved oxygen (DO), predominant macrohabitat unit (where BW indicates

backwater), mean percent cover by woody debris, vegetation, and detritus,

predominant substrate type (with the dominant substrate listed first) , and most

common genera of aquatic vegetation (listed in order of importance). Standard

errors are in parentheses. See text for a description of how each habitat

parameter was identified ..............................................................................................47

3. Date of capture, location, total length (TL), and weight (Wt) of wild

Muskellunge collected from the upper Caney Fork River and its tributaries

in 2012 using a bag seine and boat electrofishing gear ...............................................48

4. Summary of information for radio-tagged Muskellunge stocked into the Collins

River, Tennessee, 9 November 2012. Fate was determined after 56 d; TL = total

length. Tags recovered on the bank are indicated, as are the tags from two fish

consumed by a Great Blue Heron (see text for further details) ...................................49

vi

LIST OF FIGURES

Figure Page

1. Map of the upper Caney Fork River and its major tributaries above Great Falls

Dam, Tennessee ...........................................................................................................51

2. Map of sampling locations for age-0 Muskellunge in the upper Caney Fork

River system, Tennessee, using three different sampling gears: bag-seine,

backpack electrofisher, and boat electrofisher .............................................................52

3. Map of spawning areas located using traditional ecological knowledge, direct

observation, and boat-electrofishing in the Collins River, Tennessee .........................53

4. Map of locations where wild age-0 Muskellunge were collected, and nursery

habitat was quantified in the upper Caney Fork River system, Tennessee ..................54 5. Growth of age-0 Muskellunge collected in the upper Caney Fork River system,

Tennessee, in 2012. Two fish collected in the upper Collins River were

considered outliers and were excluded from the model...............................................55

6. Dispersal over 56 d of radio-tagged Muskellunge stocked at two sites on the

Collins River, Tennessee .............................................................................................56

7. Numbers of fingerling Muskellunge stocked into the upper Caney Fork River

system each year from 1976 to 2012 ...........................................................................57

1

CHAPTER 1

INTRODUCTION

Muskellunge Esox masquinongy is the largest member of the pike family

Esocidae and is a popular and economically valuable sportfish in the United States. At

one time there were thought to be three Muskellunge subspecies based on regional

differences in coloration: the Great Lakes subspecies E. m. masquinongy, the western or

northern subspecies E. m. immaculatus, and the Ohio subspecies E. m. ohioensis (Scott

and Crossman 1973; Crossman 1986; Crossman et al. 1986). Presently the scientific

name for Muskellunge is Esox masquinongy without a division into subspecies (Nelson et

al. 2004). Muskellunge populations have not been well studied in the southern U.S.

(Etnier and Starnes 1993; Brenden et al. 2006), and the most recently published study on

river-dwelling Muskellunge in Tennessee is more than 50 years old (Parsons 1959). In

Tennessee, the native range of Muskellunge includes tributaries to the Tennessee River

and Cumberland River (Parsons 1959; Etnier and Starnes 1993).

Loss or modification of habitat has been identified as the main cause of

population decline of Muskellunge in the United States (Dombeck et al. 1984). In

Tennessee during the early 20th

century, Muskellunge were so prevalent in their native

streams on the Cumberland Plateau that local anglers used nets, traps, gigs, trot lines, and

even shotguns to harvest fish (Parsons 1958). By the early 1930s, Muskellunge

populations were in decline in these streams, and 1950 was the last year local anglers

considered musky fishing to be “good” in the area. Angling effort and success for

Muskellunge declined significantly during the early 1950s and illegal harvesting of

2

Muskellunge was identified as a problem in the area at that time (Parsons 1958).

Muskellunge populations were described as “rapidly dwindling” in the 1950s and habitat

in more than 168 km of native Muskellunge streams in Tennessee had been destroyed by

acid mine drainage. A statewide 635 mm total length (TL) limit was placed on

Muskellunge in 1955, the only size restriction on any warm-water sportfish in Tennessee

at the time.

In 1955, 20 Muskellunge fingerlings (~180 mm TL) were transplanted from Rock

Creek, Tennessee, a Cumberland Plateau stream, to the upper Caney Fork River in a pilot

study to test the stocking efficacy of Muskellunge in that system. Great Falls, a large

waterfall on the Caney Fork River near Rock Island, Tennessee, prevented Muskellunge

and other large piscivore species such as Walleye Sander vitreus and Sauger Sander

canadensis from migrating upstream of the falls into the upper Caney Fork River and its

tributaries (Parsons 1958; Little et al. 1983). In 1957, five of the fish from the original

stocking were observed, indicating good survival (Parsons 1958). Based on this pilot

study, the upper Caney Fork River system was identified as a viable Muskellunge

restoration area.

Muskellunge populations were severely depleted throughout their native range in

Cumberland Plateau streams by the early 1970s (Riddle 1975). Muskellunge were

declared endangered in the state of Tennessee in 1975 by the Tennessee Wildlife

Resources Commission. Habitat destruction from coal mining was identified as the

primary cause of Muskellunge population decline; overfishing, poor logging practices,

and pollution by industrial and domestic sources also negatively impacted native

Muskellunge (Parsons 1952; Parsons 1958; Riddle 1975; Garavelli 1977; Little et al.

3

1983; TWRA 2011). The Tennessee Wildlife Resources Agency (TWRA) began stocking

Muskellunge into the upper Caney Fork River system above Great Falls Dam in 1976 as

part of a statewide effort to restore Muskellunge to the Cumberland River drainage

(Table 1). To that end, TWRA began propagating Muskellunge at Eagle Bend Hatchery

in Clinton, Tennessee in 1978 (Little et al. 1983). Although the upper Caney Fork River

system is not a documented part of the native range of Muskellunge in Tennessee,

TWRA chose it as a focal area for establishing a Muskellunge population because it is in

the Cumberland River drainage, it is similar in habitat to the species’ native streams, and

there was no threat of future habitat degradation by coal mining (Little et al. 1983).

Additionally, the pilot study in 1955-1957 (Parsons 1958) confirmed that Muskellunge

stocked into the upper Caney Fork River could grow and survive well.

Muskellunge have been stocked sporadically in the upper Caney Fork River and

its tributaries above Great Falls Dam since stocking began in 1976 (Table 1). The number

of fish stocked annually varies based on availability and in some years no fish are

stocked. The upper Caney Fork River system now supports a popular Muskellunge

fishery. After Muskellunge populations recovered sufficiently from their endangered

status, the fishery was opened in 1988 with a statewide 915 mm TL minimum length

limit and a creel limit of one fish per d (TWRA 2011). The goals of Muskellunge

management in Tennessee are to maintain the quality of the upper Caney Fork River

system fishery and to maintain the native range of Muskellunge in Tennessee (TWRA

2011). In recent years, some spawning and natural reproduction were thought to occur in

the upper Caney Fork watershed above Great Falls Dam (J. Swearingen, TWRA,

personal communication); however, it was not known with certainty whether natural

4

reproduction was occurring or whether recruitment was sufficient to maintain the fishery.

Soon after stocking of Muskellunge began in 1976 in the upper Caney Fork River system,

Little (1983) recognized the need to determine whether natural reproduction occurs there

and whether naturally reproduced Muskellunge grow well and reproduce. Despite this

recommendation, there has been no investigation of natural reproduction in the system

since then.

Etnier and Starnes (1993) reported that Muskellunge spawn in April and May in

Tennessee at water temperatures of 9.4-15° C (Etnier and Starnes 1993). Parsons (1959)

reported that Muskellunge spawned at 10° C in April in Tennessee. Muskellunge in rivers

moved upstream to spawn and used only 12% of total stream habitat for spawning

(Parsons 1958). There is some evidence that Muskellunge possess a homing instinct and

return to the same spawning grounds each year (Crossman 1990; Younk et al. 1996;

Jennings et al. 2011). Muskellunge spawning habitat in Tennessee has been described as

shallow pools with abundant organic material and aquatic vegetation in low-gradient

reaches with substrates dominated by silt (Parsons 1958, 1959). Spawning habitat in

midwestern lakes was shallow (< 1 m), near influent streams with warmer temperatures

than adjacent areas, and had muck or sand substrate with woody debris and vegetation

(Dombeck et al. 1984).

Muskellunge are broadcast spawners and high mortality is common in early life-

history stages of broadcast spawners that provide no parental care (Dahlberg 1979;

Dombeck et al. 1984; Zorn et al. 1998). Muskellunge eggs suffered high mortality in self-

sustaining lake populations in northwest Wisconsin and in a hatchery, indicating that

Muskellunge hatching success may always be poor, even under ideal conditions (Zorn et

5

al. 1998). Unlike northern pike Esox lucius eggs that adhere to vegetation, Muskellunge

eggs are not adhesive and fall to the substrate (Scott and Crossman 1973). Low dissolved

oxygen (DO) concentrations at the substrate:water interface can compromise

reproductive success (Scott and Crossman 1973; Dombeck et al. 1984). Organic carbon

levels, texture of spawning substrate, and water temperature were not associated with

reproductive success in Wisconsin lakes (Zorn et al. 1998). Dissolved oxygen

concentrations at the substrate:water interface were higher in systems with self-sustaining

Muskellunge populations than in systems maintained by stocking (Dombeck et al. 1984;

Zorn et al. 1998). Zorn et al. (1998) suggested that low DO concentrations at

Muskellunge spawning sites may limit reproduction. Dombeck et al. (1984) observed

high egg mortality on substrates with low DO concentrations (< 0.1 mg/l); egg survival

improved on substrates with DO concentrations of 3.8 to 4.1 mg/l.

There are four distinct life history stages for Muskellunge. Muskellunge are 9.5 to

10.3 mm TL at hatching and remain dormant in vegetation for approximately 10 d (Scott

and Crossman 1973). After the dormant larval stage, fish begin swimming and feeding

and are considered fingerlings (USFWS 1987). Fingerlings feed on zooplankton for one

to three weeks and switch to a piscivorous diet at approximately 40 mm TL (USFWS

1987). At approximately 75 mm TL, the fish is considered a juvenile until the onset of

sexual maturity (Buynak and Mohr 1979). Males in Tennessee streams matured at

approximately 560 mm TL and age 3; females matured at approximately 635 mm TL and

age 3 or age 4 (Parsons 1959).

Knowledge of the specific habitat requirements of Muskellunge at different life

stages was limited in the 1980s (Dombeck 1986) and remains so today. Larval fish

6

remain close to hatching locations; however, spawning habitat varies among systems and

there appears to be no specific combination of vegetation, depth, and substrate that is

critical for reproduction to occur (Dombeck et al. 1984). Fingerlings preferred shallow

habitats with aquatic vegetation or fallen tree trunks and branches for cover as their

nursery habitats in Canada and northern Wisconsin (Craig and Black 1986; Hanson and

Margenau 1992). Juvenile habitat use and the behavior of recently-stocked Muskellunge

are poorly understood, but a few studies have used radio-telemetry to study the ecology

of stocked Muskellunge. Hanson and Margenau (1992) studied post-stocking dispersal,

movements, and habitat selection of age-0 Muskellunge in two Wisconsin lakes. Wagner

and Wahl (2011) studied movement, home range size, and habitat selection by stocked

age-2 Muskellunge in an Illinois lake and Wagner and Wahl (2007) evaluated

temperature-selection among different stocks of age-2 Muskellunge.

Age-0 Muskellunge forage along shallow (< 3 m deep) shorelines and are

generally found near aquatic vegetation or submerged branches (Parsons 1959; Craig and

Black 1986; Hanson and Margenau 1992; Farrell and Werner 1999; Wagner and Wahl

2011). Density of age-0 Muskellunge varied inversely with water depth in nursery bays

of the upper St. Lawrence River (Farrell and Werner 1999; Murry and Farrell 2007).

Juvenile Muskellunge selected shallow littoral habitat in an Illinois lake and avoided

pelagic habitat in all seasons (Wagner and Wahl 2011). Characteristics of shallow-water

habitats preferred by juvenile Muskellunge included: sand substrate with submersed

vegetation, muck substrate without vegetation, downed trees (Hanson and Margenau

1992), coarse woody habitat with aquatic macrophytes (Wagner and Wahl 2011), dense

emergent vegetation (Craig and Black 1986), and submerged tree trunks with woody

7

debris (Kapuscinski et al. 2010). Age-0 Muskellunge habitat in bays of the upper St.

Lawrence River, New York, had 20-60% coverage by submerged macrophytes (Murry

and Farrell 2007).

The Muskellunge fishery in the upper Caney Fork River system above Great Falls

Dam is gaining popularity as a trophy fishery. In recent years, anglers have reported

catching fish longer than 1.3 m TL and large Muskellunge are routinely observed in

annual electrofishing surveys (J. Swearingen; TWRA, personal communication). Post-

stocking mortality, movement, and habitat use have not been documented for southern

U.S. Muskellunge (Wagner and Wahl, 2011). Studies of stocking efficacy and the extent

of natural recruitment will likely be important to the future management of Muskellunge

in this and other systems in the southeastern United States. The objectives of the study

were to 1) identify and describe spawning and nursery habitats for wild Muskellunge in

the upper Caney Fork River system; 2) assess the extent of natural reproduction and

growth of wild juvenile Muskellunge in the upper Caney Fork River system; and 3)

describe mortality, dispersal, and habitat use by recently stocked Muskellunge.

8

CHAPTER 2

STUDY AREA

This study was conducted on the upper Caney Fork River and its four major

tributaries (Calfkiller River, Rocky River, Collins River and Cane Creek) above Great

Falls Dam (Figure 1). The Caney Fork River is a tributary to the Cumberland River. The

Tennessee Electric Power Company constructed Great Falls Dam upstream of Great Falls

in 1916 for the purpose of power generation; it was sold to the Tennessee Valley

Authority (TVA) in 1939 (TVA 2011). The dam is 28 m high and 244 m wide and the

power plant has two generating units (TVA 2011). It is the only TVA dam located

outside of the Tennessee River watershed. Great Falls Lake is 35.4 km long and has a

surface area of approximately 740 ha at full pool. Seventy-six km of the Collins River, 20

km of the Calfkiller River, 16 km of the Rocky River, and 11 km of Cane Creek were

surveyed and sampled in this study. An additional 7-km reach of the Caney Fork River

near its confluence with Cane Creek was sampled.

The upper Caney Fork River Watershed spans eight counties in middle

Tennessee: Bledsoe, Cumberland, Grundy, Putnam, Sequatchie, Van Buren, Warren, and

White Counties; and three ecoregions: Cumberland Plateau, Plateau Escarpment, and

Eastern Highland Rim (TDEC 2003). The watershed of the entire Caney Fork River from

its headwaters to the Cumberland River covers approximately 458,687 ha (TDEC 2003).

Muskellunge spawning habitat was described only in the Collins River because

annual TWRA electrofishing surveys were restricted to that river during the spawning

period and the traditional ecological knowledge of anglers and fishing guides was most

9

readily available in that tributary. All tributaries and the mainstem Caney Fork River

were sampled to document nursery habitat and natural reproduction. The study of

mortality and dispersal of stocked fish was confined to the Collins River because it

received all Muskellunge stocked in the upper Caney Fork River system in 2012.

10

CHAPTER 3

METHODS

Spawning and Nursery Habitats of Wild Muskellunge

Spawning locations in the upper Caney Fork River tributaries were located by

direct observation and accessing traditional ecological knowledge from local fishing

guides and biologists (Brewer 1980; Craig and Black 1986; Price and Rulifson 2004;).

Although surveys of spawning areas in previous studies elsewhere were conducted at

night by boat using spotlights (Younk et al. 1996; Zorn et al. 1998; Rust et al. 2002),

those studies took place on lakes that were less dangerous to navigate at night than rivers.

In March and April 2012, I visually surveyed the upper Caney Fork River tributaries by

boat during the day to locate aggregations of Muskellunge presumably engaged in

spawning. I accompanied local fishing guides and TWRA biologists and noted locations

where they had observed Muskellunge spawning behavior in previous years. Similarly,

Dombeck et al. (1984) located spawning areas in lakes in the midwestern United States

by surveying fisheries biologists and Craig and Black (1986) located spawning areas in

Lake Huron, Ontario by surveying local fishing guides. I participated in TWRA’s annual

boat electrofishing survey of adult Muskellunge that took place on the Collins River

during spring 2012. I noted all locations where adult Muskellunge were seen or captured

in multiples, or captured and displayed signs of spawning activity (i.e., flowing eggs or

milt; exhibiting open lesions or torn fins).

11

Habitat parameters were recorded where spawning activity was observed (i.e.,

spawning habitat) or an age-0 Muskellunge was collected (i.e., nursery habitat). When

age-0 Muskellunge were captured in seine hauls, habitat parameters were measured at the

beginning and end of each 20-m haul. I took a GPS waypoint (using a Garmin GPSmap

60CSx receiver), recorded water temperature and DO concentrations in the water column

near the surface using a YSI 55 handheld DO meter, and measured depth using a meter

tape. Additionally, I characterized substrate, macrohabitat, percent cover, and vegetation.

Dominant and sub-dominant substrates were visually classified as predominantly

silt/clay, sand, gravel, pebble, cobble, or boulder, with sample codes of 0 through 5

respectively, based on the particle size ranges outlined by Bain (1999). Macrohabitat was

classified as pool, backwater, riffle, run, or littoral (Younk et al. 1996). Using a 1-m2

floating grid I visually estimated the percentage of woody debris, vegetation, and detritus

to quantify cover, and identified aquatic vegetation to genus level (Craig and Black 1986;

Zorn et al. 1998; Farrell and Werner 1999; Murry and Farrell 2007). Waypoints

identifying spawning sites and wild juvenile fish locations were mapped using ArcGIS

10.0 (ESRI, Redlands, CA).

Natural Reproduction and Growth of Wild Muskellunge

I used seining, backpack electrofishing, and boat electrofishing techniques to

determine the extent of natural reproduction and to describe growth of wild Muskellunge

in the four major tributaries and in the main stem of the upper Caney Fork River (Figure

2). I chose sampling sites for seining and backpack electrofishing based on wadeability,

12

accessibility by small motorboat or canoe, and suitability of habitat for juvenile

Muskellunge based on the literature. I used a bag seine (6 m x 1.8 m with a 1.8 m x 1.8 m

bag) to sample wild juvenile Muskellunge between 23 June and 22 August 2012. The

wings of the seine had 6.3 mm mesh and the bag had 4.7 mm mesh. The seine was

constructed of nylon delta knotless material and had a double lead-line and large floats. I

followed a modified version of the seining protocols of Farrell and Werner (1999), Murry

and Farrell (2007), and Kapuscinski et al. (2010, 2012). The seine was slowly dragged

parallel to shore for 20 m before being swung into shore. Sampling sites ranged from

about 20 to 100 m of shoreline and no more than three seine hauls were made at each site.

Each site sampled with the seine was revisited 24-48 h later and sampled using

backpack DC-electrofishing gear (Smith-Root model LR-24). A single zigzag pass was

made through each site by a two-person crew and the time was recorded.

I also sampled for age-0 Muskellunge beginning 23 August 2012 using a Smith-

Root 2.5 GPP electrofishing unit mounted in a 4.3-m johnboat powered by a jet-drive

outboard motor. Surveys targeted likely Muskellunge nursery habitat in reaches of all

four tributaries accessible by motorboat, as well as reaches of the upper Caney Fork

mainstem that could not be seined or electrofished using backpack electrofishing gear.

Boat electrofishing transects were not of a predetermined length of shoreline or time, but

total pedal-time at each site electrofished was recorded.

Each Muskellunge captured was weighed (g), measured (TL, mm), and scanned

for a microtag (using a handheld detector wand (Northwest Marine Technology, Inc.)

because all fish stocked the previous year (2011) were microtagged. Fish were also

scanned for a PIT tag using a Biomark 601 PIT-tag reader in case the individual was a

13

recapture from earlier in the 2012 field season. All untagged Muskellunge were PIT-

tagged in the left dorsal musculature before they were released. A GPS point was taken

each time an age-0 Muskellunge was collected.

Mortality, Dispersal, and Habitat Use of Stocked Muskellunge

I used radio telemetry to monitor mortality and dispersal of stocked fish. I

performed a pilot-study in December 2011 to evaluate radio tag retention and post-

surgical survival in age-0 Muskellunge using dummy radio tags (similar in size and

weight to the tags used to track fish). Surgeries were performed as described below on

eight age-0 Muskellunge (mean TL = 336.0 mm TL; SE = 7.9; mean weight = 172.2 g;

SE = 12.1) at Gallatin State Hatchery in Gallatin, Tennessee. Average fish surgery time

was 4.5 minutes. Eight fish (mean TL = 341.2 mm TL; SE = 6.7; mean weight = 173.6 g;

SE = 14.8) that were anesthetized but did not undergo surgery served as controls. All fish

implanted with dummy tags were alive and retained their tags after 32 d and all control

fish survived.

Forty-one age-0 Muskellunge were intensively reared in raceways on pellet feed

at North Carolina Wildlife Resources Commission’s Table Rock Fish Hatchery in

Morganton, North Carolina. When fish reached about 76 mm TL they were transported to

Eagle Bend Hatchery in Clinton, Tennessee, on 13 June 2012 where they were held in

indoor tanks and switched to a live minnow diet. Twenty-five fish (mean TL = 311.0 mm

TL; SE = 1.2; mean weight = 151.0 g; SE = 3.8) were surgically implanted with

Advanced Telemetry Systems model number F1530 radio tags on 5 November 2012.

14

Each tag broadcasted a signal on a unique frequency between 30.010 and 30.300 MHz.

Each tag weighed 1.7g in air (less than 2% of the weight of the smallest tagged fish;

Winter 1996), had a 20-cm long antenna, and a battery life of 71 d. Surgical tools were

autoclaved prior to the day of surgery. All tags were sanitized in chlorohexadrine

gulconate solution (2%) and rinsed with sterile saline prior to implantation (Mulcahy

2003). Surgeons wore sterile latex gloves. Each fish was anesthetized in a cooler using 50

ppm tricaine methanesulfonate (MS-222) (Hanson and Margenau 1992), measured (TL,

mm) and weighed (g). Fish were then placed ventral side up on a V-shaped wooden

trough covered with surgical draping. During surgery a 30ppm MS-222 solution was

pumped over the gills to maintain sedation. The mucous coating along the linea alba of

each fish was removed with a sterile cotton swab (Wildgoose 2000) before a 2 cm

incision was made anterior to the pelvic girdle (Lucas and Baras 2000). Using the

shielded needle technique (Ross and Kleiner 1982), the whip antennae was threaded

through the body cavity, over the pelvic girdle, and out between the pelvic fins using a

3/8 circle reverse triangular size 6 needle. Incisions were closed with two interrupted

sutures of size 3-0 Monocryl Plus (Ethicon, Inc.; Juarez, Mexico). Fish revived in a

cooler of untreated hatchery water before being transferred to a circular tank. Over the

course of implanting tags into those 25 Muskellunge, eight other fish (mean TL = 308.6

mm TL; SE = 3.4; mean weight = 143.2 g; SE = 5.0) were surgically implanted with

dummy tags (similar in size and weight to the functioning tags) and eight fish (mean TL

= 304.3 mm TL; SE = 4.2; mean weight = 142.4 g; SE = 6.6) that served as controls were

anesthetized but did not undergo surgery. Average fish surgery time was 3.5 minutes.

15

Implanted fish were allowed 4 d to recover (Hanson and Margenau 1992;

Oldenburg and Brown 2010) before they were loaded onto a hatchery truck and

transported to the Collins River for stocking on 9 November 2012. Fish implanted with

dummy tags and control fish were held in the hatchery for an additional 20 d to monitor

tag retention and post-surgical survival.

Radio-tagged fish were stocked at two boat ramps on the Collins River. Radio

tags of all fish were checked and were functioning at the time of stocking. Twelve fish

were stocked at the Veterans of Foreign Wars Lodge ramp on the lower Collins River

(hereafter the lower Collins River stocking site) and 13 fish were stocked at the Myers

Cove ramp on the upper Collins River (hereafter the upper Collins River stocking site).

The mean weights and total lengths of fish stocked at each site were similar ( 1 ; P

> 0.0752). The first tracking occasion took place 4 d after stocking to allow fish

additional time to adjust to the tags and recover from being transported to the river

(Clapp et al. 1990; Markham et al. 1991; Wagner and Wahl 2011).

Radio-tagged fish were tracked at least 2 d per week for the first 33 d post-

stocking and once more at 56 d post-stocking. Fish were located from a motorboat or

canoe using an ATS 4500S digital scanning receiver, a boat-mounted directional loop

antenna, and an omni-directional coaxial cable antenna. Fish were located from long

range using the loop antenna, and more precisely using the omni-directional coaxial cable

antenna (Niemela et al. 1993). A fish was considered dead when a tag was recovered or a

fish did not move over two or more consecutive tracking occasions. A fish was

considered missing if it was located on most tracking occasions but then disappeared and

could not be found again. Near the end of the 56 d study, I did a sweep of the Collins

16

River beginning 5 km upstream of the upper Collins River stocking site and ending at

Great Falls Lake (~ 56 km downstream) to try and locate missing fish.

Survival of radio-tagged fish over the 56 d study was estimated using a known-

fate model in program MARK (White and Burnham 1999; Ridgway 2002). Known-fate

models can be used when most individuals are located on each tracking occasion and few

individuals go missing over the course of the study. Fates can be live, missing, or dead at

the end of a tracking event. Survival estimates are based on the probability that an

individual will survive the interval between consecutive tracking occasions. Intervals of

varying length between tracking events can be accounted for in the model. Initial weight

and total length were tested as covariables and survival estimates were converted to a

daily instantaneous mortality rate (Miranda and Bettoli 2006).

Movements between tracking occasions and dispersal from stocking sites for each

individual were measured in ArcMap Version 10.0 (Bettinger and Bettoli 2002; Wagner

and Wahl 2011). Movements were measured as straight-line distances between locations

and were converted to a daily movement rate for each fish (daily movement rate =

straight line distance in the river channel / number of days between tracking occasions)

(Wagner and Wahl 2011; Petty et al. 2012). A mixed-model analysis of variance was

performed in Statistical Analysis Software Version 9.2 to test for differences between

stocking sites in fish movements (Rogers and White 2007). Total dispersal from each

stocking site was measured as the straight-line distance in the river channel between the

upstream-most fish and the downstream-most fish using the last locations where fish

were known to be alive.

17

To describe habitat use by stocked fish, habitat parameters were recorded each

time a radio-tagged fish was located. The same habitat parameters, except depth, were

measured for stocked fish as those described above for wild fish. Depth was not recorded

because radio telemetry methods were not accurate enough (~ 10 m) to determine the

exact location, and therefore depth occupied, of a fish.

18

CHAPTER 4

RESULTS

Spawning and Nursery Habitats of Wild Muskellunge

Reports of Muskellunge spawning activity by anglers were received as early as

the second week of March in 2012. Spawning locations were limited to the Collins River

(Figure 3) because (1) anglers did not identify spawning areas outside of that tributary,

and (2) TWRAs annual sampling during the spawning season was restricted to that

tributary. Nursery habitats for wild Muskellunge were identified in the upper Caney Fork

River, Cane Creek, the Calfkiller River, and the Collins River, but not in the Rocky River

(Figure 4).

Shallow depths (< 1m) and finer substrates (i.e., silt, sand, gravel) were an

important feature of spawning and nursery habitats of wild Muskellunge (Table 2).

Vegetative cover also appeared to be an important component in both habitat types,

particularly in spawning habitat (mean = 66.5%; SE = 5.58). The most common aquatic

vegetation genera in spawning habitats (in decreasing order of importance) were

Myriophyllum spp., Vallisneria spp., and Potamogeton spp.; Justicia spp., Myriophyllum

spp., and Polygonum spp. were the most common genera in nursery areas.

19

Natural Reproduction and Growth of Wild Muskellunge

Eighteen wild age-0 Muskellunge were observed in the Calfkiller River, Cane

Creek, the Collins River, and the upper Caney Fork River (Figure 4; Table 3); one fish

could not be netted but its location and habitat data were recorded. No wild age-0 fish

were collected in the Rocky River. The first wild fish was collected in a seine haul on 25

June 2012; the last was captured in a boat electrofishing sample on 26 October 2012.

None of those wild fish had a PIT-tag (i.e., none were recaptures from earlier in the

season). Of all three gears, boat electrofishing was the most efficient means of collecting

wild age-0 Muskellunge. Twelve fish were collected using boat electrofishing gear in 6.2

hours of pedal time in 9 field days by a two-person crew (1.9 fish per hour of pedal time).

Six fish were collected in 80 seine hauls in 11 field days by a two-person crew (mean

catch = 0.0750; SE = 0.0296). No wild Muskellunge were collected by backpack

electrofishing in 5.9 hours of sampling time in 9 field days by a two-person crew.

Age-0 Muskellunge in the upper Caney Fork River and its tributaries ranged in

length from 148 mm TL early in the growing season to 399 mm TL at the end of the

growing season in 2012 (Figure 5). Muskellunge grew in a linear fashion through 10

October 2012 (F = 216.11; df = 1, 13; P < 0.0001; R2 = 0.9433). The only two fish

collected from the upper Collins River were collected late in the season, were much

shorter than their counterparts, and were considered outliers. These two fish were

excluded from the regression model.

20

Mortality, Dispersal, and Habitat Use of Stocked Muskellunge

All tagged fish destined to be stocked were alive and retained their radio tags 4 d

after surgery. All fish implanted with dummy tags and held in the hatchery were alive and

retained their tags after 24 d.

Two stocked fish were omitted from the known-fates model. One fish (30.030)

was never located after stocking. Another fish (30.080) was tracked to a pool 9 km

downstream of the lower Collins River stocking site on the twelfth (and last) tracking

event. Its tag was in water about 1.5 m deep and could not be retrieved. That fish was

designated a mortality, but the interval in which it died was unknown. Parameter

estimates for the initial total length and weight covariables in the known-fates model

were considered imprecise because the 95% confidence intervals contained zero. Hence,

these covariables were not included in the final model that was used to estimate post-

stocking survival. The two cohorts of stocked fish experienced significantly different

mortality over the 56 d study. Mortality was 43.4% at the lower Collins River stocking

site (Zdaily = 0.01015), and 98.3% at the upper Collins River stocking site (Zdaily =

0.07278). Pooled mortality over 56 d was 78.6% (Zdaily = 0.02750). During one tracking

event, two individuals were tracked to a single Great Blue Heron Ardea Herodias (Table

4). Six tags were collected on the shoreline of the Collins River. Five of those tags

showed signs of chewing along the antenna and one was recovered outside a River Otter

Lontra canadensis den surrounded by scat and Otter tracks. Four fish that had been

located at least 11 times were missing by the end of the study.

21

The reach of river that encompassed the last known locations of live fish (i.e.

dispersal) was 660 m at the lower Collins River stocking site (Figure 6). This

measurement excludes one fish (30.100) found 746 m upstream on the third tracking

event that was dead when located, and another fish (30.080) found 7,100 m downstream

on the twelfth tracking event that was dead when located (Table 4). Dispersal at the upper

Collins River stocking site was 1,150 m. As with dispersal, movement rates (m/d) were

higher at the upper Collins River stocking site, where stocked fish experienced higher

mortality, than at the lower Collins River stocking site (P = 0.0019; F = 12.76; df = 1,

20).

Habitat use by stocked age-0 Muskellunge appeared to be different from that of

wild age-0 Muskellunge, particularly in terms of percent cover (Table 2). Vegetative

cover was low (mean = 5.8%; SE = 1.19) at sites occupied by stocked fish and the most

common genera were Myriophyllum spp., Justicia spp., and Vallisneria spp. Vegetative

cover was higher at sites where wild age-0 fish were collected and the prevalent genera at

those locations were Justicia spp., Myriophyllum spp., and Polygonum spp. Cover by

woody debris (mean = 28.1%; SE = 1.56) and detritus (mean = 28.5%; SE = 1.23)

appeared to be important components in nursery habitats utilized by stocked

Muskellunge. Substrates were similar (e.g., silt and sand) in the nursery habitats used by

stocked and wild age-0 fish.

22

CHAPTER 5

DISCUSSION

Spawning and Nursery Habitats of Wild Muskellunge

Spawning habitats in the Collins River were similar to those described in other

studies in which shallow depths, vegetative cover, and fine substrates were important

characteristics, and other attributes were variable. Spawning areas in other lakes and

rivers that supported self-sustaining Muskellunge populations were also shallow (< 100

cm, Dombeck 1979 and Zorn et al. 1998; mean depth = 74 cm, Dombeck et al. 1984;

mean depth = 81 cm, Younk et al. 1996). Bottom characteristics in spawning areas in

Midwestern lakes with self-sustaining Muskellunge populations were variable in terms of

percent cover, species of aquatic vegetation, and substrate type (Dombeck et al. 1984).

The most common genera of aquatic vegetation in spawning areas of the upper Caney

Fork system (e.g., Myriophyllum, Vallisneria, and Potamogeton) were similar to those

found in previous studies. For instance, Potamogeton spp. and Myriophyllum spp. were

common in Muskellunge spawning areas in Midwestern lakes (Dombeck et al. 1984).

Potamogeton spp., Lemna spp., and Chara spp., were dominant in spawning areas in the

St. Lawrence River and Farrell et al. (1996) concluded that dense vegetation in spawning

areas may prevent the nonadhesive eggs of Muskellunge from falling to the sediments

and dying, or being displaced by currents. Finer substrates (silt, sand, gravel) are

common in spawning areas in lakes and rivers with self-sustaining Muskellunge

populations and appear to yield higher DO concentrations at the substrate-water interface.

23

In a laboratory experiment that measured DO concentrations over different substrates,

DO was highest over sand, gravel, glass, and screen (3.8-4.1 mg/L), intermediate over

wood (1.3 mg/L), and lowest over leaves, plants, and silt (0.0-0.2 mg/L) (Dombeck et al.

1984). Similarly, two of five stockings of fertilized eggs over sand and gravel substrates

contributed to a measurable year-class of Muskellunge in Wisconsin lakes (Hanson et al.

1986). Spawning areas in the Mississippi River were in backwater habitats described as

finger channels, side channels, or island coves with low flow, muck, silt, or sand

substrates and decomposing vegetation (Younk et al. 1996). No differences in organic

carbon content, substrate texture, or water temperature were detected between self-

sustaining lakes and stocked lakes in Wisconsin and water velocity was low in all

spawning areas (Zorn et al. 1998). Muskellunge reproductive success (i.e., hatching

success) may always be low, even in lakes with self-sustaining populations (Zorn et al.

1998).

In the present study, DO at the substrate-water interface was not measured; rather

DO was measured in the water column near the surface. However, this may be an

important variable to measure in future studies because DO at the substrate-water

interface plays an important role in Muskellunge reproductive success in systems with

self-sustaining populations. Wisconsin lakes with self-sustaining Muskellunge

populations had more variable and often higher DO concentrations (0.5-9.6 mg/L) at the

substrate-water interface than lakes sustained by stocking (1.2-5.4 mg/L) (Zorn et al.

1998). Midwestern lakes with self-sustaining Muskellunge populations had higher DO

concentrations, (6.0-8.4 mg/L), at the substrate-water interface compared to lakes

24

sustained by stocking that had lower DO concentrations, (0.4-2.4 mg/L) (Dombeck et al.

1984).

Woody debris was not a common substrate characteristic of spawning areas in the

present study; however, other studies linked woody debris to increased Muskellunge egg

survival. In laboratory experiments, eggs survived better when held in suspension

(11.82%) or deposited on wood (0.93% survival) than placed directly on the substrate

(0.05% survival) (Zorn et al. 1998). A higher percentage of spawning areas in self-

sustaining lakes were covered by woody debris than in stocked lakes in northern

Wisconsin (Zorn et al. 1998). In laboratory experiments, eggs experienced high mortality

when incubated over leaves and macrophytes, moderate mortality over sand and glass,

lower mortality over polyethylene screen and wood, and lowest mortality over gravel and

silt (Dombeck et al. 1984). These findings indicate that woody debris may increase egg

survival in spawning areas by keeping eggs off the substrate where low DO

concentrations may kill Muskellunge eggs. Conversely, percent cover by woody debris

was low (1-4%) in spawning areas in northern Wisconsin lakes and did not differ

between self-sustaining and stocked systems (Rust et al. 2002).

Water level fluctuations may affect spawning success of Muskellunge, but this

relationship has not been quantified and conclusions from prior studies are confounding.

Rising spring-time water levels were an important predictor of Muskellunge spawning

success in Midwestern lakes (Dombeck et al. 1986). Zorn et al. (1998) caught more fry

during fall electrofishing surveys in a lake that had the greatest daily water level changes

during spawning and egg development in northwestern Wisconsin. Conversely, Rust et

al. (2002) observed water levels dropping after spawning in half of Wisconsin lakes with

25

self-sustaining Muskellunge populations and rising in 90% of lakes maintained by

stocking. Low spring discharge in Kentucky streams was associated with Muskellunge

reproductive success based on contribution to the age-1 year-class (Brewer 1980).

Nursery habitats of wild age-0 Muskellunge in the upper Caney Fork River

system were similar to those in other studies: shallow depths and moderate vegetative

cover were important characteristics. Muskellunge nursery habitats in other studies were

shallow (< 305 cm deep, Hanson and Margenau 1992; 25-50 cm deep, Zorn et al. 1998; <

150 cm deep, Farrell and Werner 1999; < 200 cm deep, Kapuscinski et al. 2012) and

associated with emergent and submerged macrophytes (Hanson and Margenau 1992;

Zorn et al. 1998; Murry and Farrell 2007; Kapuscinski et al. 2012). Age-0 Muskellunge

avoided open areas with depths greater than 305 cm in northern Wisconsin lakes (Hanson

and Margenau 1992). Catch per effort of age-0 Muskellunge declined with increasing

depth in upper St. Lawrence River nursery bays (Farrell and Werner 1999). Similarly,

there was an inverse relationship between age-0 Muskellunge abundance and water depth

in nursery bays of the St. Lawrence River (Murry and Farrell 2007). One of the most

common aquatic vegetation genera that occurred in nursery areas in the present study

(e.g., Myriophyllum) was also common in nursery bays of the upper St. Lawrence River

(Farrell and Werner 1999).

The availability of suitable prey species is an important characteristic of

Muskellunge nursery habitat. There was a direct relationship between age-0 Muskellunge

occurrence and prey availability, specifically Cyprinids Notropis spp., Banded Killifish

Fundulus diaphanous, and Tesselated Darters Etheostoma olmstedi in St. Lawrence River

nursery bays (Murry and Farrell 2007). When age-0 Muskellunge were collected in a

26

seine haul in the present study, other species collected were similar to species identified

as important prey of age-0 Muskellunge in northern U.S. rivers (Murry and Farrell 2007).

Banded Killifish, cyprinids, and darters Etheostoma spp. or Percina spp. were the most

important prey items in diets of age-0 Muskellunge in the Niagara and St. Lawrence

Rivers (Kapuscinski et al. 2012). Seine hauls in which age-0 Muskellunge were collected

in the present study commonly included Cyprinids (Telescope Shiners Notropis

telescopus, Striped Shiners Luxilus chrysocephalus, Scarlet Shiners Lythrurus fasciolaris,

and Central Stonerollers Campostoma anomalum), as well as Cherry Darters Etheostoma

etnieri, and Northern Studfish Fundulus catenatus. Muskellunge recruitment in Kentucky

streams was related to the relative abundance of small forage fishes including Cyprinids

(Brewer 1980). Kapuscinski et al. (2012) concluded that the availability of fusiform prey

species in a system should enhance survival of age-0 Muskellunge.

Natural Reproduction and Growth of Wild Muskellunge

Natural reproduction was documented in the upper Caney Fork River system in

four of five study rivers in the present study. Muskellunge recruitment in Kentucky was

higher in streams with a greater relative abundance of small forage fishes, especially

Cyprinids, and in streams that had relatively undeveloped watersheds (Brewer 1980).

High discharges, unseasonable temperatures, and high silt loads during the spawning

season were inversely related to recruitment in Kentucky streams. The present study

collected naturally reproduced age-0 Muskellunge over a single field season; however, it

should be noted that recruitment of native populations can be variable from year to year

27

(Brewer 1980). Whether 2012 was a good or poor year for recruitment in the upper

Caney Fork River system cannot be determined from the current study, but these data

may be used to compare to sampling results in future years.

Age-0 Muskellunge in the Caney Fork River system reached lengths of up to 399

mm TL by 9 October 2012. The large size achieved by the end of the growing season

might be attributed to a warm spring and early spawning in 2012 that began about 6

weeks earlier than is usual in Tennessee (Parsons 1959). Average total lengths for age-1

Muskellunge ranged from 216 mm to 338 mm TL in prior studies in Tennessee and five

other states (Schloemer 1936; Parsons 1959; Axon 1978; Belusz 1978; Miles 1978;

Brewer 1980; Larscheid et al. 1999). Naturally-reproduced Muskellunge in Wisconsin

Lakes reached approximately 250 mm TL by the end of their first growing season and

approximately 375 mm TL by the end of their second growing season (Jonas et al. 1996).

No age-0 Muskellunge were collected early in the season from the upper Collins

River to compare to the two age-0 fish that were collected from that reach late in the

season; those two fish were excluded from the growth model. Additional field

observations will be required to confirm that age-0 Muskellunge in the upper Collins

River grow slower than in the lower Collins River, and if so, to determine the factors

responsible for different growth.

Mortality, Dispersal, and Habitat Use of Stocked Muskellunge

This is only the second study anywhere to monitor mortality and dispersal of

stocked age-0 Muskellunge. Post-stocking mortality of Esocids has long been a concern

28

for fisheries managers and often occurs in the first few weeks after stocking (Carline et

al. 1986; Margenau 1992; Hanson and Margenau 1992; Wahl and Stein 1993; Szendry

and Wahl 1995; Szendry and Wahl 1996; Farrell and Werner, 1999). Fifty days after

stocking pellet-reared fingerling Tiger Muskellunge E. masquinongy x E. lucius, total

mortality ranged from 5 to 100% with an average of more than 70% across 16 trial

stockings into small impoundments in Ohio (Carline et al. 1986). Hanson and Margenau

(1992) observed 57% mortality of stocked radio-tagged age-0 Muskellunge after 34 d in

one lake, but only 15% mortality after 34 d in another lake in northern Wisconsin. In a

study of post-stocking fate of juvenile Northern Pike, Muskellunge, and Tiger

Muskellunge, survival through the first fall increased with stocking size and survival was

best when fish were longer than 175mm TL (Wahl and Stein 1993). However, most fish

in that study died within 2 months of stocking. Szendry and Wahl (1995) were unable to

consistently capture Muskellunge and Tiger Muskellunge one month post-stocking in

Illinois reservoirs, suggesting poor survival. Nearly all mortality of three sizes of stocked

Muskellunge occurred within 10-14 d post-stocking in northern Illinois reservoirs

(Szendry and Wahl 1996). Post-stocking survival of fall fingerling and spring yearling

Muskellunge was low and variable, with highest mortality occurring in the first few

months after stocking for both cohorts (Margenau 1992).

Factors that may affect post-stocking survival of Muskellunge are hatchery

rearing techniques, the size of fish at stocking, dispersal from stocking sites, and

movement rates. Rearing technique and length at stocking explained most (67%) of the

variation in survival to age-5 of stocked Muskellunge in Chautauqua Lake, New York

(McKeown et al. 1999). Muskellunge and Tiger Muskellunge reared on a live minnow

29

diet experienced higher post-stocking survival through fall than pellet-reared fish in

Illinois reservoirs, although prey-consumption, foraging behavior, avoidance behavior,

and habitat use were similar (Szendry and Wahl 1995). More pellet-reared Esocids were

lost to Largemouth Bass Micropterus salmoides predation, and therefore suffered higher

mortality, than Esocids reared on a live-minnow diet in Illinois reservoirs (Szendry and

Wahl 1995). Similarly, intensively-reared Tiger Muskellunge suffered greater predation

by Largemouth Bass than extensively-reared Muskellunge in Ohio reservoirs (Wahl and

Stein 1989). Post-stocking survival of Muskellunge raised on a live minnow diet was

highest for pond-reared Muskellunge fingerlings, intermediate for trough-reared and

“pond-finished” fingerlings, and lowest for fingerlings reared in troughs (McKeown et al.

1999). Because the stocked Muskellunge in the present study were reared intensively in

troughs and tanks, the post-stocking mortality estimates may be liberal. Although

survival has been linked to rearing technique, dispersal of stocked Esocids in Illinois

reservoirs was not affected by rearing technique (Szendry and Wahl 1995). Survival of

stocked Muskellunge in the upper Caney Fork River system might improve if extensively

reared Muskellunge were stocked. Generally, post-stocking survival of Muskellunge has

been inversely related to size at stocking, and stocking larger fingerlings has been more

cost-effective on a per survivor basis (Margenau 1992; Johnson and Margenau 1993;

Szendry and Wahl 1996; McKeown et al. 1999). However, stocked juvenile Muskellunge

did not suffer major losses to predation in those studies.

Predation played a major role in the post-stocking mortality of Muskellunge in the

present study. Twenty percent of confirmed mortalities resulted from mammal predation

and 8% from heron predation. Five of six radio-tags were recovered from the shoreline

30

and showed signs of chewing along the antennae, indicating predation by mammals. One

of these five tags was found outside a River Otter den with scat around it. Another

possible predator that resides along the Collins River is the Mink Neovison vison. Great

Blue Herons and River Otters were identified as predators of fingerling Muskellunge in

enclosures that were intended to monitor short-term post-stocking survival in a

Wisconsin lake (Margenau 1993). European Otters Lutra lutra readily preyed on

Northern Pike and Brown Trout Salmo trutta in European rivers (Sulkava 1996; Ludwig

2002; Jacobsen 2005; Aarestrup et al. 2005). European Otters and Great Blue Herons can

consume fish longer than the largest stocked Muskellunge in the present study. In a study

of post-stocking movement and mortality of Brown Trout in a Danish river, European

Otters and Great Blue Herons were identified as predators for age-1 (mean length = 190

mm TL) and age-2 fish (mean length = 273 mm TL) (Aarestrup et al. 2005). Jacobsen

(2005) documented European Otters preying upon stocked Brown Trout up to 390 mm

TL in a Danish River. Great Blue Herons can swallow Rainbow Trout Oncorhynchus

mykiss up to 390 mm TL (Hodgens et al. 2004). Great Blue Herons generally prey on fish

in shallow waters (Hodgens et al. 2004) and juvenile Muskellunge may be particularly

susceptible to avian predation because shallow depths are such an important habitat

characteristic for them. Although the role of Largemouth Bass predation on post-stocking

mortality of Esocids has been studied (e.g., Stein et al. 1981; Carline et al. 1986; Wahl

and Stein 1989; Szendry and Wahl 1995; Wahl et al. 2012), the role of predation by Great

Blue Herons and River Otters on post-stocking mortality of Muskellunge has not been

reported in the literature.

31

Dispersal from stocking sites and movement rates influence survival of stocked

fish. Post-stocking dispersal of Northern Pike fry was inversely related to size in a

Denmark lake (Skov et al. 2011). The two cohorts of Muskellunge stocked into the

Collins River in the present study were of similar length, but their dispersal differed. It

remains to be seen why stocked Muskellunge dispersed further from the upper Collins

River stocking site than the lower Collins River stocking site and whether a direct

relationship exists between dispersal and mortality in stocked Muskellunge. Other studies

have documented a direct relationship between dispersal (or movement) and post-

stocking mortality. Brown Trout that survived 5 weeks post-stocking in a Danish stream

moved less than fish that died or went missing by the end of the study (Aarestrup et al.

2005). Bettinger and Bettoli (2002) concluded that high dispersal rates of recently

stocked Rainbow Trout compared to resident Rainbow Trout in the Clinch River,

Tennessee, contributed to poor survival of stocked fish. Hanson and Margenau (1992)

monitored advanced Muskellunge fingerlings for 34 d post-stocking in two Wisconsin

lakes and daily movements were greater during the first 17 d of the study. These findings

that daily movements are greatest soon after stocking and fish that move more suffer

higher rates of mortality imply that mortality is highest soon after stocking.

In the present study, cover by aquatic vegetation was not a common habitat

characteristic in areas used by stocked fish, although it was common in nursery areas of

wild fish. Prior studies documented dense vegetation as a common characteristic of age-0

Muskellunge habitat (Hanson and Margenau 1992; Zorn et al. 1998; Murry and Farrell

2007; Kapuscinski et al. 2012). Conversely, percent cover in habitats occupied by

stocked age-0 Muskellunge in the Collins River consisted mainly of woody debris and

32

detritus; whereas, cover in habitats occupied by wild age-0 Muskellunge consisted mainly

of aquatic vegetation. In a study comparing the vulnerability of predator-acclimated and

naïve Muskellunge and Tiger Muskellunge to Largemouth Bass predation, survival rates

in laboratory, pond, and lake environments were similar for both species and both

treatments; however, experienced Esocids spent more time in vegetation than naïve

Esocids, and experienced fish moved less in the presence of Largemouth Bass (Wahl et

al. 2012). The finding that naïve fish moved more in the presence of a predator, in

combination with findings that fish that move more suffer higher mortality (e.g.,

Bettinger and Bettoli 2002; Aarestrup et al. 2005) suggests that the naiveté of the

intensively-reared fish in the present study may have contributed to their high mortality

rates. Wahl et al. (2012) suggested that rearing Esocids in the presence of cover and

natural predators may encourage them to use vegetation as a shield against predation

when they are stocked.

33

CHAPTER 6

MANAGEMENT IMPLICATIONS

High post-stocking mortality of fall fingerlings in the present study indicates the

need to re-evaluate TWRA’s annual Muskellunge stocking regime, especially when

considering the high costs of rearing fingerlings that are stocked in the fall. This study

documented that Muskellunge are reproducing in the upper Caney Fork River system and

producing fall recruits. Because stocking has been irregular since 1976 (Figure 7) and

annual boat electrofishing surveys did not begin until 2007, it is difficult to determine

whether the Muskellunge population is currently self-sustaining. If stocking is

discontinued, managers should be able to discern through annual boat-electrofishing

surveys whether the population is, in fact, self-sustaining or whether it requires

supplemental stocking to maintain a fishable stock.

If the TWRA decides to resume stocking Muskellunge in the future, post-stocking

mortality rates should be assessed for fry and for extensively-reared spring yearlings (as

alternatives to fall fingerlings) to determine which life-stage (if either) contributes to age-

1 abundance in the upper Caney Fork River system. Stocking Muskellunge fry and spring

yearlings have been effective strategies to augment year-class production in other

fisheries. In St. Lawrence River nursery bays, fry stocked in early July experienced lower

mean survival (~0.7% over ~1 month) than fingerlings stocked in early August (7.3-

18.0% over ~1 month) (Farrell and Werner 1999). Despite high mortality rates, fry in that

study contributed the most to age-0 abundance (~52%), whereas fingerlings contributed

21-25% and wild fish contributed 20-27% (Farrell and Werner 1999). Conversely,

34

stocking fry into Kentucky reservoirs yielded no age-1 Muskellunge (Brewer 1980).

Yearling Muskellunge were stocked in the spring in Wisconsin lakes before 1963, but

improved hatchery techniques produced fall fingerlings nearly as long as spring yearlings

and the yearling stocking program was discontinued. However, when post-stocking

survival to 18 months was compared, stocking spring fingerlings was one to four times

more cost-effective than stocking fall fingerlings (Margenau 1992). In Iowa lakes,

yearling fingerling Muskellunge raised on a live-minnow diet and stocked in the spring

survived better than fall-stocked fingerlings (Larscheid et al. 1999). It is not clear from

these studies which stocking strategy (fry or spring yearlings), if either, would be more

effective than current practices of stocking fall fingerlings in the upper Caney Fork River

system. However, because River Otters and Great Blue Herons were documented as a

major source of predation for large fall fingerlings in the present study, spring yearlings

that have been held in hatchery conditions even longer may be equally or more

susceptible to predation.

In the future, researchers should seek to quantify the role that native predators

such as Great Blue Herons and River Otters have on juvenile Muskellunge survival in the

upper Caney Fork River system and elsewhere. Wahl (1999) encouraged biologists to

examine the ecology of a river system when developing a Muskellunge stocking

program, emphasizing that predation, competition, and other abiotic factors play

important roles in the development of a fishery. Given that predation on stocked fish was

high (and may also be high for extensively-reared fish) and natural reproduction is known

to occur, the TWRA might be justified in discontinuing the Muskellunge stocking

program and continuing annual boat electrofishing surveys to monitor the fishery in the

35

upper Caney Fork River system. Close monitoring of the population should indicate

within a few years whether the Muskellunge fishery in the upper Caney Fork River

system is self-sustaining.

36

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45

TABLES

46

Table 1. Records of Muskellunge stocked into the upper Caney Fork River system,

Tennessee.

Year River Stocked Source Number

1976 Caney Fork Minor Clark, KY 62

Bee Creek Minor Clark, KY 40

1979 Caney Fork Eagle Bend, TN 404

Calfkiller Eagle Bend, TN 120

1980 Calfkiller Eagle Bend, TN 356

1982 Collins Eagle Bend, TN 664

1983 Collins Minor Clark, KY 66

1984 Caney Fork Minor Clark, KY 2,500

1991 Caney Fork Minor Clark, KY 302

1994 Calfkiller Minor Clark, KY 50

Collins Minor Clark, KY 70

Rocky Minor Clark, KY 45

1997 Calfkiller Minor Clark, KY 84

Collins Minor Clark, KY 118

2000 Caney Fork Minor Clark, KY 40

Collins Minor Clark, KY 100

2002 Caney Fork Minor Clark, KY 60

Collins Minor Clark, KY 288

2005 Caney Fork Minor Clark, KY 60

Collins Minor Clark, KY 100

Collins Eagle Bend, TN 56

2006 Caney Fork Minor Clark, KY 200

Caney Fork MO Department of Conservation 547

Collins Minor Clark, KY 210

Collins MO Department of Conservation 1,000

2007 Caney Fork Minor Clark, KY 250

Collins Minor Clark, KY 250

2008 Calfkiller Eagle Bend, TN 45

2010 Calfkiller Minor Clark, KY 388

Cane Creek Minor Clark, KY 100

Caney Fork Minor Clark, KY 100

Collins Minor Clark, KY 521

Collins Eagle Bend, TN 122

2011 Collins Minor Clark, KY 600

2012 Collins Table Rock, NC 25

47

Table 2. Summary of habitat parameters measured at spawning and nursery areas for wild and stocked Muskellunge in the

upper Caney Fork River system in Tennessee in 2012, including the number of sites (N), mean depth, mean dissolved oxygen

(DO), predominant macrohabitat unit (where BW indicates backwater), mean percent cover by woody debris, vegetation, and

detritus, predominant substrate type (with the dominant substrate listed first) , and most common genera of aquatic vegetation

(listed in order of importance). Standard errors are in parentheses. See text for a description of how each habitat parameter was

identified.

Habitat N

Mean

depth

(cm)

Mean

DO

(mg/L)

Macro

habitat

Cover (%)

Woody

Debris

Vegetation Detritus Substrate Vegetation

Genera

Spawning 34 92.0

(6.06)

8.1

(0.16) Run

3.5

(1.33)

66.5

(5.58) 0 Sand, Gravel

Myriophyllum,

Vallisneria,

Potamogeton

Nursery

(Wild) 18

71.4

(8.66)

9.7

(0.31) BW/Run

10.6

(4.75)

33.3

(9.39)

6.9

(2.59) Silt, Sand

Justicia,

Myriophyllum,

Polygonum

Nursery

(Stocked) 154

10.5

(0.06) Run

28.1

(1.56)

5.8

(1.19)

28.5

(1.23) Silt, Sand

Myriophyllum,

Justicia,

Vallisneria

48

Table 3. Date of capture, location, total length (TL), and weight (Wt) of wild

Muskellunge collected from the upper Caney Fork River and its tributaries in 2012 using

a bag seine and boat electrofishing gear.

Date River Latitude / Longitude Method TL

(mm)

Wt

(g)

6/25/2012 Calfkiller 35.907336 / -85.477889 Seine 150 13

7/2/2012 Collins 35.706794 / -85.725217 Seine 148 15

7/16/2012 Cane Creek 35.770831 / -85.400347 Seine 157 17

7/16/2012 Cane Creek 35.770832 / -85.401129 Seine 164 19

7/16/2012 Cane Creek 35.773094 / -85.404820 Seine 177 24

7/17/2012 Cane Creek 35.790905 / -85.405111 Seine 168 20

8/24/2012 Cane Creek 35.805081 / -85.440383 Electrofishing 253 82

8/29/2012 Collins 35.780986 / -85.682483 Electrofishing 276 102

9/14/2012 Collins 35.706725 / -85.725400 Electrofishing 314 138

9/14/2012 Collins 35.706864 / -85.725351 Electrofishing 362 238

10/9/2012 Collins 35.708785 / -85.731825 Electrofishing 362 247

10/9/2012 Collins 35.708762 / -85.737660 Electrofishing 391 302

10/9/2012 Collins 35.711685 / -85.740586 Electrofishing 399 320

10/9/2012 Collins 35.717833 / -85.741609 Electrofishing 386 309

10/10/2012 Caney Fork 35.818806 / -85.430172 Electrofishing 320 161

10/26/2012 Collins 35.629279 / -85.694686 Electrofishing 247 61

10/26/2012 Collins 35.625166 / -85.698671 Electrofishing 281 92

49

Table 4. Summary of information for radio-tagged Muskellunge stocked into the Collins

River, Tennessee, 9 November 2012. Fate was determined after 56 d; TL = total length.

Tags recovered on the bank are indicated, as are the tags from two fish consumed by a

Great Blue Heron (see text for further details).

Frequency Days

Observed

Total

Observations Fate

Initial

Weight

(g)

Initial

TL

(mm)

Comments

30.010 14 7 Dead 99 307

30.020 56 13 Live 135 308

30.030 1 1 Unknown1 143 306

30.040 27 11 Missing 138 310

30.050 7 4 Dead 130 300 Recovered

30.060 27 11 Missing 147 308

30.070 10 5 Dead 134 304

30.080 56 2 Dead1 153 313

30.090 19 8 Dead 168 317

30.100 7 2 Dead 171 318 Recovered

30.110 5 3 Dead 191 322 Recovered

30.120 56 13 Live 162 314

30.130 29 8 Dead 151 308

30.140 7 4 Dead 181 325 Recovered

30.150 5 3 Dead 154 309 Recovered

30.160 33 12 Dead 149 307 Heron

30.180 10 5 Dead 143 309

30.190 56 13 Live 170 318

30.201 19 8 Dead 164 316

30.250 56 13 Live 161 315

30.260 19 8 Dead 152 313 Recovered

30.270 33 12 Dead 161 313 Heron

30.280 10 5 Dead 137 301

30.290 27 11 Missing 148 311

30.300 33 12 Missing 133 304 1 Left out of known-fates model in Program MARK

50

FIGURES

51

Figure 1. Map of the upper Caney Fork River and its major tributaries above Great Falls

Dam, Tennessee.

52

Figure 2. Map of sampling locations for age-0 Muskellunge in the upper Caney Fork

River system, Tennessee, using three different sampling gears: bag-seine, backpack

electrofisher, and boat electrofisher.

53

Figure 3. Map of spawning areas located using traditional ecological knowledge, direct

observation, and boat-electrofishing in the Collins River, Tennessee.

54

Figure 4. Map of locations where wild age-0 Muskellunge were collected, and nursery

habitat was quantified in the upper Caney Fork River system, Tennessee.

55

Figure 5. Growth of age-0 Muskellunge collected in the upper Caney Fork River system,

Tennessee, in 2012. Two fish collected in the upper Collins River were considered

outliers and were excluded from the model.

0

50

100

150

200

250

300

350

400

450

To

tal

Len

gth

(m

m)

Other Rivers & Lower Collins River

Upper Collins River

June July August September October

Total Length = 70.72 + (2.34*Study Day)

R2 = 0.9433

P < 0.0001

F = 216.11

56

Figure 6. Dispersal over 56 d of radio-tagged Muskellunge stocked at two sites on the

Collins River, Tennessee.

57

Figure 7. Numbers of fingerling Muskellunge stocked into the upper Caney Fork River

system each year from 1976 to 2012.

0

100

200

300

400

500

600

700

1975 1985 1995 2005

Nu

mb

er

Year

1,231,957

2,500

58

APPENDIX

59

Species and counts of fishes collected by seine haul during summer 2012 sampling in the

upper Caney Fork River system. The number of seine hauls in each river is in parentheses

next to the river name.

Species

Collins

River

(34)

Rocky

River

(6)

Cane

Creek

(17)

Calfkiller

River

(23)

Muskellunge Esox masquinongy 1 . 4 1

Northern Studfish Fundulus catenatus 77 . 26 16

Western Mosquitofish Gambusia affinis 4 . . .

Banded Sculpin Cottus carolinae 2 . . .

Golden Redhorse Moxostoma erythrurum . . 1 7

Northern Hogsucker Hypentelium nigricans . . 3 28

Central Stoneroller Campostoma anomalum 201 9 21 115

Striped Shiner Lusilus chrysocephalus 369 209 119 59

Scarlet Shiner Lythrurus fasciolaris 47 10 143 40

Telescope Shiner Notropis telescopus 546 75 272 313

Highland Shiner Notropis micropteryx 7 31 . 69

Whitetail Shiner Cyprinella galactura 12 . . 26

Bigeye Chub Hybopsis amblops 56 . . 8

Common Carp Cyprinus carpio 1 . . .

Bluntnose Minnow Pimephales notatus 40 . 10 19

Smallmouth Bass Micropterus dolomieu 43 2 4 43

Largemouth Bass Micropterus salmoides 18 18 12 5

Bluegill Lepomis macrochirus 117 136 3 32

Rock Bass Ambloplites rupestris 338

1 10

Redear Sunfish Lepomis microplophus 1 8 . 18

Longear Sunfish Lepomis megalotis 10 5 2 6

Cherry Darter Etheostoma etnieri 9 . 4 1

Greenside Darter Etheostoma blennioides 4 . 2 8

Logperch Percina caprodes . . . 12

60

VITA

Lila Hammond Warren was born on April 29, 1987 and grew up in Paris,

Virginia. She graduated from Episcopal High School in Alexandria, Virginia in 2005.

She received her Bachelor of Arts degree from the University of Virginia in

Charlottesville, Virginia in 2009 with a double-major in Environmental Science and

Environmental Thought and Practice. She began coursework at Tennessee Technological

University in August 2011 and received a Master of Science degree in Biology with a

concentration in Fisheries Science in August 2013.