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Electronic Theses and Dissertations Graduate School

2018

Leaping Behavior In Silver Carp (Hypophthalmichthys Molitrix): Leaping Behavior In Silver Carp (Hypophthalmichthys Molitrix):

Analysis Of Burst Swimming Speeds, Angle Of Escape, Height, Analysis Of Burst Swimming Speeds, Angle Of Escape, Height,

And Distance Of Leaps And Distance Of Leaps

Ehlana Stell University of Mississippi

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LEAPING BEHAVIOR IN SILVER CARP (HYPOPHTHALMICHTHYS MOLITRIX):

ANALYSIS OF BURST SWIMMING SPEEDS, ANGLE OF ESCAPE, HEIGHT, AND

DISTANCE OF LEAPS

A Thesis presented in partial fulfillment of requirements for the degree of Master of Science in the Department of Biology

The University of Mississippi

By

EHLANA G. STELL

May 2018

Copyright Ehlana G. Stell ALL RIGHTS RESERVED

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ABSTRACT

Silver Carp have rapidly expanded their range exploiting vulnerable habitats, disrupting

fisheries, and inflicting unknown ecological damage. These fish have continued to spread into

the Middle Mississippi River and the Tennessee River Valley and great effort is being expended

to prevent Silver Carp from entering the Great Lakes and expanding further into the Ohio,

Illinois, Missouri, and Tennessee Rivers. Using boat-mounted cameras, we recorded in situ video

of invasive Silver Carp, Hypopthalmichthys molitrix leaps to measure their horizontal distance,

height, and angle of escape as well as their burst speed. Video tapes of fish leaps were obtained

from populations of carp in Mississippi, Missouri, and Illinois. Additionally, morphometric and

environmental data were measured at each site. Carp reached mean leap heights of 124 cm with a

maximum of 276.08 cm in Ramsey Creek, Missouri. Maximum horizontal distance reached was

482.34 cm with a mean distance of 207.02 cm. Burst speeds varied across the three study sites

with significantly higher speeds in Missouri (628.4 + 99.9 cm/s (n=10)) compared to Mississippi

(471.2 + 77.2 cm/s (n=42)) and Illinois (551.7 + 95.7 cm/s (n=35)). Total lengths of Silver Carp

increased with decreasing latitude; 73.09 + 11.05 cm (n=113) from Mississippi, 60.86 + 4.1 cm

(n=30) from Missouri, and 54.79 + 9.3 cm (n=161) from Illinois. Our results documented the

burst speed of Silver Carp across a range of sizes and areas, revealed that the leaping abilities of

Silver Carp are greater than previously estimated, and demonstrated differences in leap

characteristics across populations of carp.

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DEDICATION

I would like to dedicate this thesis to my parents and other family for their constant

support and encouragement even as I pursued a degree they did not expect. I love you and could

not have done this without all of y’all!

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ACKNOWLEDGMENTS

There are several people who have helped me to accomplish this project, but first and

foremost I must thank Dr. Glenn Parsons for his taking a chance on me. I got to know you while

working on Bryan’s project and as a result you gave me many opportunities as an undergraduate

to experience research and start thinking beyond just topics covered in class. Your teaching and

guidance has helped me find a field where I am passionate about the work and truly enjoy my

job! You took a chance with me not having the highest GPA (but still good!) as an undergrad by

fully accepting me into your lab. If not for you, I don’t know where my life would have gone

beyond graduation. Thank you for all the support, reviews, and conversations over the last 5

years.

Along with Dr. Parsons, Dr. Jan Hoover also deserves much thanks! You were the first to

question me about what I wanted to do after graduation on a very cold winter day on Moon Lake

and pushed me to consider graduate school. Throughout my own research you have been very

present answering my many questions, coauthoring papers and being patient through my first

attempt at the whole publishing experience. Without your help, I am not sure this project would

have happened (quite literally in the funding department). With Jan came a host of others at

ERDC that have provided me help over the years: Alan Katzenmeyer, Jay Collins, Jack Killgore,

and Bill Lancaster (a better fisherman than I will ever be!). Between helping construct my net to

providing field assistance in Havana, IL, I can never express how grateful I am for your help and

express how highly I think of each of you.

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Dr. Buchholz, beyond being on my committee, you have dealt with me through every

course you teach and have influenced the way I approach questions in a way like no other.

Behavior is something I see left out of many projects in my field and I hope in the future I can

further incorporate what you have taught me to improve that area of fisheries and fish biology.

Behavior influences everything and thanks to you I now know how to recognize and study it.

A huge thanks goes to all those who volunteered to help me collect samples and be

bombarded by leaping fish, deal with boat breakdowns (thanks to said leaping fish!) and have

survived all the adventures associated with working in rural areas! A special thanks to Lauren

Fuller who was literally a target for these fish my first summer and who has aided me countless

times in writing, preparing presentations, and being a general soundboard for ideas. Bryan Cage,

I must thank you because without you and your project, I would never have had the opportunity

to begin doing research and would not have learned all the lessons about field work I did from

you!

To all my friends both in Shoemaker (Amy Hribar, Tim Colston, Lance Sullivan, Matt

Ward, Lauren Fuller (again), and Alan Katzenmeyer) and out (Riley Mitchell, April Steen, Casey

Caveness, George Humphry, Brian Taylor, Kristie Newland, and Trey Friend) who have kept me

sane and entertained throughout the years, thank you. You all have had to listen to my journey

for years now (and hopefully years to come!) and have provided much needed support and often

times distractions from all the hectic schedules and work.

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Finally, I want to thank my family. You all thought I was going to go into the medical

field but instead decided to play with fish as you say. Thanks for not calling me too crazy and

always being there when I needed you from moral support, to financial support, to taking care of

my pets when I had to run out of town for a week at a time! Mom, Dad, Rhiannon, and Amber, I

can never express how grateful I am to have all of you in my life and thank you for putting up

with my student career.

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TABLE OF CONTENTS

ABSTRACT………………………………………………………………………………………ii

DEDICATION……………………………………………………………………………………iii

ACKNOWLEDGEMENTS………………………………………………………………………iv

LIST OF FIGURES………………………………………………………………………………ix

CHAPTER 1: 1.1 GENERAL BACKGROUND..………………………………………………..1

1.2 MEASURING SPEED AND RESEARCH JUSTIFICATION……………………………….8

CHAPTER 2: MATERIALS AND METHODS..……………………………………………….12

2.1 STUDY ORGANISM: SILVER CARP HYPOPHTHALMICHTHYS MOLITRIX….12

2.2 NET CONSTRUCTION AND DESIGN…………………………………………….15

2.3 STUDY SITES.…..…………………………………………………………….……17

2.4 FIELD AND POST FIELD ANALYSIS………………….…………………………19

2.5 STATISTICAL ANALYSIS…………..………………………………………….…21

CHAPTER 3: ANALYZING LEAP CHARACTERISTICS AND BURST SPEEDS OF SILVER

CARP (HYPOPHTHALMICHTHYS MOLITRIX) USING IN-SITU VIDEO ANALYSIS……...22

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3.1 INTRODUCTION……….…………………………………………………………..22

3.2 RESULTS……………..……………………………………………………………..24

3.3 DISCUSSION……….….……………………………………………………………36

3.3.1 BURST SPEED……..………………………….………………………….36

3.3.2 LEAPING ABILITIES AND CHARACTERISTICS…………………….40

3.3.3. GEOGRAPHIC VARITION……………………………………………...44

CHAPTER 4: CONCLUSIONS/RECOMMENDATIONS…..…………………………………46

BIBLIOGRAPHY………………………………………………………………………………..48

VITA……………………………………………………………………………………………..58

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LIST OF FIGURES

Figure 1. Original surface level net for Silver Carp Sampling…………………………………..16

Figure 2. Elevated carp catching net design…………………………………………………….16

Figure 3. Sample locations for 2016-2017……………………………………………………….19

Figure 4. Total length (black) and weight (grey) of Silver Carp captured from each location.

Means and standard errors are shown………………………………………...………...………..25

Figure 5. Comparison of upper and lower caudal lobe lengths. Means and standard errors are shown. …………………………………………………………………………………………...26

Figure 6. Bar graphs of leap distance (a) (n=81) and leap height (b) by total length (n=81). 95%

CI are shown……………………………………………………………………………………..27

Figure 7. Regressions of burst speed on total length for each study site.………………….….…28

Figure 8. Scatterplot of total length (cm) and relative speed (body lengths*s-1).…………..…....29

Figure 9. Leap distance (a) (n=84) and leap height (b) (n=84) against burst speed for Silver Carp

from each study site. Scatterplot (c) of burst speed and height shows the significant correlation

found for Mississippi fish. Means and 95% confidence intervals are shown. Bars without

confidence intervals only had one sample…………………………………………………….…30

Figure 10. Leap distance, leap height, and burst swimming speed for all study sites. Means and

standard errors shown…………………………………………………………………..………..31

x

Figure 11. Average leap distances displayed against angle of escapes shows no differences.

Means and 95% confidence intervals are shown. Bars without confidence intervals only had one

sample……………………….…………………………………………………………………...32

Figure 12. Frequency distribution of leap angle for silver carp .………………………………...33

Figure 13. Plot of burst speed (cm/s) against leap distance (cm) with interaction of angle

(degrees)………………………………………………………………………………………….34

Figure 14. Path analysis for distance (cm) showing correlation values………………………….34

Figure 15. Correlation of height and angle of escape.……………………….…………………..36

Figure 16. Path analysis for height (cm) showing correlation coefficients ………………..……34

1

CHAPTER 1: GENERAL BACKGROUND

1.1 Types of Leaping

Any fisherman can attest to seeing fish of multiple species leap out of the water for no

apparent reason. Sometimes it is a solitary individual while other times it is entire schools of fish.

However, there is no single explanation for why fishes display this behavior and scientist have

developed several alternative hypotheses in an attempt to describe leaping or jumping behaviors.

A previous review by Saidel et al (2004) described only three hypotheses encompassing all

leaping behaviors in fish (predator avoidance, predation, and spawning). However, I have found

at least 7 hypotheses explaining leaping in fish and the diverse research that has taken place has

shown no single hypothesis is correct at all times. These hypotheses include:

1. Traversing Migrational Obstacles

Fish utilize jumping for multiple purposes even amongst conspecifics. One prime

example of jumping behavior is observed in salmon which use jumping to traverse waterways

and clear obstacles. The Kokanee salmon (Oncorhynchus nerka) has been documented

spontaneously jumping to scale waterfalls during their upstream migrations under specific

conditions (Lauritzen et al. 2010). Lauritzen et al. used video analysis to conduct field studies of

the salmon to reveal that fish use an S-start acceleration from beneath the surface immediately

followed by burst swimming until exiting the water and require a minimal depth to achieve

maximal acceleration. In the same study Lauritzen et al. described an alternative approach, the

2

C-start standing jump in which the fish, already near or at the surface, jump out of the water

without a running start (Lauritzen et al. 2010). This type of jumping behavior may result in only

3

partial breaching of the fish instead of a complete exit from the water. With this

information, scientists can improve the efficacy of fish ladders in order to facilitate the

movement of salmon. Conversely, similar data on Silver Carp will allow scientist to create

unfavorable passages and perhaps prevent their expansion.

The gobiid fish (Bathygobius soporator) likewise overcome obstacles by jumping from

one tidal pool to another even when the second pool is not visible (Aronson 1971, Warburton

1990). These fish arrive at high tide, remain in the tidal pools until conditions are no longer

favorable, and then escape into the next tidal pool with very accurate orientation and precision. It

is believed that the fish use previous high tides to learn the topography to maximize their

jumping ability (Aronson 1971).

1. Predator Avoidance

Aside from using jumping as a navigational tool, this behavior has also been documented

as a possible predator avoidance tactic. The ability to jump from the path of a predator is vital for

optimizing survival and can be presumed to be under intense selective pressures (Soares &

Beirman 2013). This behavior often results in anti-predatory movements in the horizontal

direction.

An interesting predator avoidance behavior was observed in two fully aquatic fishes

(Gambusia affinis and Danio rerio) that live at the water’s edge (Gibb et al. 2011). These fish

will intentionally strand themselves on land in an effort to avoid predators even though they

possess no morphological specializations for terrestrial locomotion. Instead, Gibb et al. (2011)

found they use a two-step tail flip jumping behavior to generate terrestrial jumping and return to

the water before they asphyxiate. The process begins as a fast start (S-start) aquatic type jump

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where a high-curvature lateral flexion of the body is followed by a flexion of the posterior body

section. This allows the fish to lift onto the peduncle that kicks outward propelling the fish

forward. By exiting the water, fish reduce the resistance and drag encountered during the escape

possibly gaining a few seconds of rapid distance gain.

3. Respiration

Several members of the Mugilidae family are well known for leaping out of the water.

One such member is the commercially important Striped Mullet (Mugil cephalus) (Hoese 1985).

Evidence shows mullet do not always leap in response to predation, however, Hoese (1985)

suggested that fish leap to expose their gills to atmospheric oxygen. The Striped Mullet and

menhaden (Brevoorita patronus) which also jump, both possess large gill surface areas relative

to body size. However, neither fish possess the common secondary respiratory structures that

allow for aerial respiration such as the respiratory tree in the suprabranchial cavities of Walking

Catfish (Clarias batrachus) (Moyle et al. 1986) or use the swim bladder as a respiratory organ

similar to gar (Lepisosteus spp.). Aside from jumping, mullet have also been documented

congregating in schools with their mouths, eyes, and upper opercular located above the water’s

surface. The amount exposed above the surface is enough to air pump which ventilates the large

pseudobranchia and 30% of the gill filaments in the pharyngobranchial organ (Hoese 1985). One

issue with using atmospheric oxygen in conjunction with gill filaments is that the gill filaments

collapse without the support of water leading to fish asphyxiation (Hoese 1985, Gibb et al.

2011). To reduce the occurrence of gill filament collapse, the mullet have stiffened filaments that

are capable of staying erect outside of water (Hoese 1985) which is somewhat similar to other air

breathing fishes. This respiration hypothesis arises from the mullets’ feeding habits as the

5

schools feed in low oxygen bottom waters. The fish likely deplete their oxygen supply and must

quickly replenish their oxygen by using high atmospheric concentrations.

4. Predation

Leaping behaviors are exhibited in multiple predatory species. Shih and Techet (2010)

documented Archer Fish (Toxotes microlepis) use leaping as a way to ambush their prey (insects)

resting on the water’s surface. They used high speed video recordings to document fish lining up

below spotted prey before orienting into a specific angle, β, and launching via an S-start jump up

to 2.5 body lengths in the air. Archer Fish can be seen as ambush predators in that they wait until

an insect is hovering in place before using their defined musculature to propel themselves out of

the free surface of the water (Shih & Techet 2010).

Another ambush jumping predatory fish is the osteoglossid Silver Arowana

(Osteoglossum bicirrhosum). These fish live in waters located along banks with low lying

branches where their prey congregate. It has been found that air attacks occur at speeds of 9.2

body lengths per second whereas water based attacks occur at 3 body lengths per second (Lowry

et al. 2005). Immediately before reaching their prey, the Silver Arowana conform into an S shape

posture to ensure they will impact their prey. Unlike the Archer Fish, the Silver Arowana only

reaches 1 body length out of the water limiting its striking range. Other studies completed on

jumping predatory fish include the Four-Eyed Fish (Anableps anableps), the Rivulus (Rivulus

hartii), the Atlantic Salmon (Salmo salar), and the Sea Trout (Salmo trutta) (Soares and Bieman

2013).

5. Communication

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Gulf Sturgeon (Acipenser oxyrinchus desotoi) are anadromous fish migrating from the

Gulf of Mexico to the freshwaters of the southeastern United States. Once these fish enter

freshwater for spawning, they no longer feed and will lose much of their body weight during

their journey. Because they do not feed in freshwater, scientist have been baffled by why these

large fish expend energy for jumping behaviors that peak during known spawning times. One

hypothesis that has gained support is that these fish use jumping for school communication and

cohesion in the muddy freshwaters of Florida. Sulak et al. (2002) found evidence using

hydrophone recordings that jumping by sturgeon emits specific sounds that the fish are most

likely able to detect. These sounds travel long distances in air and even further in water allowing

the entire school to receive the messages the jumps provide.

6. Parasite Shedding

Brunnshweiler (2006) and Ritter & Brunnschweiler (2002) describe how Blacktip Sharks

(Carcharhinus limbatus) may be induced by Sharksuckers (Echeneis naucrates) to jump from

the water in an attempt to shed the parasites from their ventral and dorsal surfaces as well as their

pectoral fin regions. Brunnshweiler (2006) observed that sharks (Blacktips and Bull Sharks,

Carcharhinus leucas) will increase speed when Sharksuckers attach. These parasites likely

increase drag and irritate the skin (Brunnschweiler 2002) resulting in the fish swimming faster to

shed the parasites. Jumping occurs when the sharks are swimming at these higher speeds and will

result in them attempting to dislodge the parasitic sharksuckers during the exit and reentry of the

water before they quickly swim away. Similar to these sharks, the Paddlefish in Prairie du Sac

are afflicted by Silver Lampreys (Ichthyomyzon unicuspis) that latch onto their scaleless sides.

Luckily, the Silver Lamprey is not well known for killing their host species, but pose enough of

an irritation that the Paddlefish attempt to dislodge the lampreys (Cochran et al. 2003). This is

7

accomplished by jumping out of the water and falling back to the surface. That surface impact

causes the lampreys to release and the Paddlefish continues unhindered.

7. Unknown/ Spontaneous

The aerial jumping exhibited by the Trinidadian Guppy (Poecilia retiuclata) analyzed by

Soares and Beirman (2013) is thought to be a spontaneous action. Fish were gathered from

highly predated populations in the mountains of Trinidad and raised in captivity. These fish were

recorded using high speed video cameras to determine the kinematics of their C-start jumping

behavior. However, over the course of the study, the fish jumped with no stimuli or cues.

Another fish that jumps for unknown reasons is the American Paddlefish (Polyodon spathula).

An important fish in the caviar industry, the Paddlefish is most recognizable by its large, blade

like rostrum composed of stellate bone. This projecting rostrum not only allows the fish to utilize

three sensory pathways (electromagnetism similar to sharks, light through a photophore, and

vibrations) but also acts as a hydroplane (Allison et al. 2014). No studies have concentrated on

the jumping behavior of this important fish leaving a large gap in our knowledge. Other fish

from sharks to manta rays to a myriad of fresh and salt water fish still need their behaviors

studied to determine the cause for their jumping.

While there are specific causes for jumping in fish, the same species of fish may jump for

multiple reasons. As previously stated, Mullet may jump for respiratory purposes. However,

Mullet may also jump as a predator avoidance tactic. The behavior of the jumps is noticeably

different with the later often seen as a whole school response. The normal jump for air

accumulation is slower, shorter and involves the fish turning on its side or upside down before

entering the water. Predator avoidance or parasite dislodgment involves a more rapid jump where

8

fish maintain upright posture entering and exiting the water cleanly (Hoese 1985). Fish utilize

jumping for multiple purposes demonstrating the complexity of this behavior and why all

jumping fish cannot be grouped together.

Leaping in Silver Carp also may fall into the predator avoidance and migrational

hypotheses. As the population of Silver Carp in North America has increased, reports of serious

boater injuries resulting from collisions with leaping carp have also increased (USFWS Region

3). These fish are capable of leaping up to 3 meters above the water’s surface (Region 3 U.S.

Fish & Wildlife Service) and pose a potentially lethal risk to unsuspecting skiers and boaters.

Because they generate high vertical leaps, carp may be able to use leaping to traverse vertical

barriers similar to salmon and thus utilize the migrational hypothesis. The leaping behavior of

Silver Carp is observable when outboard motors disturb a school of carp, demonstrating a

potential predator avoidance tactic in response to the sound and pressure wave generated by boat

motors. Silver Carp are predisposed for leaping and have morphological characteristics similar to

other jumping fish such as a larger lower caudal lobe. The lower caudal fin has been measured

by the United States Army Corps of Engineers (unpublished data) and found to be 1-2 cm longer

than the upper caudal lobe similar to the exaggerated fin lengths exhibited by marine flying

fishes. However, Silver Carp are much larger (50-100 cm) than marine flying fish,

(Exocoetidae), (< 10 cm total length) and freshwater hatchetfishes that also possess jumping

abilities (15-50 cm total length). Some jumping fish possess expansive wing-like pectoral fins

allowing for large amounts of thrust to be generated for breaching the surface (Davenport 1994).

Silver Carp do not possess such a feature but do have a large ventral keel that may allow the carp

to have better directionality control when they burst towards the surface and exit the water.

9

All leaping fish use one of two methods of muscle contraction to escape the water: S-start

and C-start methods. Of the two, the S-start response is the high-performance startle response

that maximizes a peak angular acceleration and angular velocity by bending the fish body into an

s curve (Hale 2002). The velocity generated by an S-start startle response is greater than the

velocity generated in typical undulatory swimming and higher than the C-start startle response.

C-start startles are identifiable by their simultaneous unilateral muscle contraction along the full

length of the body thus only curving the form into a C shape (Hale 2002). For exiting the water,

the S-shape startle behavior is most likely utilized to generate the most thrust to break free of the

water’s surface. This maneuver most likely results in the maximum burst speed of fish being

generated during the initiation of each jump. Previous work on leaping fish has focused on why

the fishes leap or the kinesiology of leaping, but have not used leaping as a way to passively

measure burst speeds, a notoriously difficult measurement to document.

1.2 MEASURING SPEED AND RESEARCH JUSTIFICATION

Of the fish studied for their characteristic jumping, none is more notorious than the Silver

Carp (Hypopthalmichthys molitrix) for the potential and on-going ecological damage they cause

and for their unique behavior. While these fish are not documented (to my knowledge) to use

leaping as a way to traverse barriers, the risk exists that if startled, they will leap near vertical or

hydrological barriers, in place to prevent their continuing range expansion, and thus generate

enough speed and height to clear the blockades. In order to prevent this potential barrier failure,

the burst speeds of Silver Carp (maximum speed a fish achieves lasting from 1-20 seconds

(Beamish 1978)) must be known and applied to barrier construction. Traditional methods of

measuring fish swimming speeds prove useful for measuring the sustained, prolonged, and

10

critical speeds of these fish; but these methods are not suitable for inducing burst swimming

speed conditions in all fish.

Swimming performance is an important part of the life history and behaviors of fishes.

For example, a fusiform fish is likely one that swims constantly and thus will be capable of

reaching and maintaining higher speeds than a non-fusiform fish. There are three main categories

to describe swimming performance: burst, prolonged, and sustained swimming speeds (Beamish

1972). Sustained swimming speeds are those speeds a fish can maintain for longer than 200

minutes while prolonged speeds are those maintained without fatigue between 20 seconds and

200 minutes. Both of these swimming speeds use aerobic respiration and are not expected to

result in lactic acid accumulation in tissues. However, burst swimming, the maximum speed

maintained for under 20 seconds, is anaerobic quickly resulting in fatigue and requires an

extended recovery. As a result, fish are not capable of performing multiple burst in rapid

succession without developing a large, possibly lethal, oxygen debt.

Prolonged and sustained swimming speeds have been measured in multiple species

through the use of swimming chambers (Adams et al. 2003, Parsons 1990, Hoover et al. 2011,

Hoover et al. 2012). However, burst swimming speeds are notoriously difficult to measure in a

swim tunnel due to individual variations in behavior and the method in which fish generate their

burst. Often times, a burst is the result of being startled or an attempt to capture prey. In some

cases, burst speeds have been measured by rapidly increasing the water velocity in order to force

the already swimming fish to speed up exponentially but this system precludes natural behavior.

Thus, the best way to ensure that accurate burst speeds are being recorded is to stimulate a

subject in-situ and record the speed and behavior. Two alternative methods involving video

recording and analysis have proven successful in measuring the swim speeds of dolphins (Rohr

11

et al .2002) and mackerel (Videler and Wardle 1991) and blacktip sharks (Brunnschweiler 2005).

However, while successful each of these studies suffered from low sample size because fish and

marine mammals in general do not jump in high frequency. Silver Carp are an exception.

Because this species is a schooling fish, it is common to find dozens to hundreds of fish jumping

in shallow waters during the warmer summer months. This unique behavior offers the

opportunity to measure burst swimming speeds of Silver Carp outside of the confines of a swim

tunnel or flume. In order to overcome the resistance of water and exit the water fully, I have

assumed the fish are generating close to their maximal speed.

Furthermore, because this is a startle response, I attempted to test several hypotheses in

this study. 1. Fish exit the water at an optimal angle to maximize horizontal distance traveled per

leap. 2.Fish total length impacts burst speed, leap height, and leap distance. 3. Temperature

determines the maximal burst speed of fish. 4. Water velocity trains fish to burst at different

speeds. 5. There are morphological differences in fish across a latitudinal gradient. I predicted

increasing temperature, total length, and water velocity will increase the burst speed generated

by fish in that area. I expect fish at lower latitudes to be larger than conspecifics at higher

altitudes with larger fish being younger in an oxbow system. Specific objectives included 1.)

determining burst speed, 2.) determining angle of escape and establishing if there is an optimal

angle of escape, 3.) determining height and horizontal distances of leaps, 4.) examining

relationships between leap characteristics and environmental variables and 5.) establishing

variation in fish leaps across latitude.

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CHAPTER 2: MATERIALS AND METHODS

2.1 Study Organisms: Silver Carp Hypophthalmichthys molitrix

Found within the largest family of freshwater fish, the Cyprinidae, (Nelson 1994), carp

have been introduced into new habitats all over the world. Beginning in the 1970’s, Silver Carp

were intentionally stocked in Arkansas waste water ponds for use in research and to clean

wastewater lagoons (Kolar et al .2005). Floodwaters during the time Silver Carp were stocked

allowed the fish to enter the Mississippi River Basin where they have rapidly exploited the

climate and resources to establish populations in all major water systems. While the Silver Carp

are not known to be cultured for marketing purposes in the United States, they are likely still

being spread through the use of live bait (they can easily be mistaken for other minnows and

shad when juvenile), livehaulers, in fish markets, and as prayer release animals (USFWS,

Arlington VA 2007) by some Asian cultures (Kolar et al. 2005).

The Silver Carps’ natural range is in eastern China where they are found in rivers, lakes,

and reservoirs (Kolar et al. 2005). Typical migratory patterns for Silver Carp include the

upstream movement of juveniles. The young will move continuously up stream utilizing oxbow

lakes and back water areas to feed. Once the fish reach reproductive age, they will move into

headwater areas (DeGrandchamp et al. 2008) to spawn using the fast-flowing river water to

disperse the eggs downstream (Jennings 1988). Silver Carp are easily dispersed by flood waters

and so will live in ponds and lakes that are only periodically connected to river systems (Kolar et

al. 2005, Freeze and Henderson 1982). However, they will not reproduce in these low flow areas

13

meaning that if they are to enter the Great Lake system, they would need to find a high

flow area to spawn. The movements of Silver Carp are similar to paddlefish Polyodon spathula

(Zigler et al. 2003) and pallid sturgeon Scaphirhynchus albus (Hurley et al. 2004) in large river

systems possibly leading to competition for food sources and reproductive habitats

(DeGrandchamp et al. 2008, Cooke et al. 2010).

Adult Silver Carp typically have gray-black dorsal coloration changing to silver ventrally.

The upper jaw has a notched appearance while there is a small tubercle on the lower jaw. Silver

Carp have cycloid scales with a lateral line count ranging from 85-108 and do not possess true

spines. The dorsal fin can have 3-7 branched fin rays typical of carps and can reach 1.2 m

(Kamilov and Salikhov 1996). According to the Minnesota Department of Natural Resources

they can weigh up to 28 kg. While there are many native North American cyprinids, the Silver

Carp can be easily distinguished from all but the juvenile golden shiner by a well-defined ventral

keel (Kolar et al. 2005). Confusion between the golden shiner and the juvenile Silver Carp has

led to the use of Silver Carp as baitfish and propagated their spread into new water systems

(Kolar et al. 2005). Further complicating identification of Silver Carp is their ability to hybridize

with another invasive cyprinid, the Bighead carp of the same genus Hypophthalmichthys (Lamer

et al. 2011). Recognition of hybrids is vital to any field work conducted in areas of Bighead and

Silver Carp populations.

The life cycle of Silver Carp is a primary cause for their rapid spread in new territories.

Males sexually mature 1 year before females with the maximal reproductive age at 5 (Krykhtin

and Gorback 1981). However, sexually maturity for these fish is dependent on temperature. For

example, fish in Guanxi, China live under 27.2°C conditions year-round and will reach sexual

maturity within 2 years for both sexes. While the ideal temperature for Silver Carp growth is 24-

14

31° C (Mahboob and Sheri 1997) they have been documented surviving near 0°C waters in

Alberta, Canada ponds (Alberta Department of Agriculture). This allows the Silver Carp to

survive year-round temperatures in the United States Mississippi River Basin and possibly the

harsh winters of the Great Lake system. For fish in the Mississippi River, fecundity has been

documented to range from 57,283 to 328,538 eggs per individual and as in other fishes, fecundity

increases as fish size increases. In common with Bighead carp, Silver Carp typically spawn when

water levels spike with spring rains which is considered an adaptive function as it decreases egg

mortality and increases the number of eggs that reach flood plain areas. This allows the young to

grow to maturity in the relative safety of back water oxbows and ponds before using flood waters

to reenter the main river system (Krykhtin and Gorback 1981).

Silver Carp are herbivorous fish that feed on phytoplankton and zooplankton similar to

many important game fishes (Pongruktham et al. 2010, Irons et al. 2007). Silver Carp use their

pharyngeal teeth and gill rakers to filter plankton and zooplankton. They feed by pump filtration

which differs from their normal respiration by being more rapid and vigorous (Smith 1989). The

fish will gulp water, close their mouths and force the water out over the opercula. Silver Carp

may be capable of outcompeting native filter feeders by using the more rapid pump filtration

method especially when they switch to zooplankton in times of low phytoplankton abundance

(Kolar et al. 2005). The abundance and structure of phytoplankton communities has been shown

to be affected by Silver Carp filter feeding behavior and usually results in a shift towards smaller

plankton species (Smith 1989).

Silver Carp are schooling fishes that spend the majority of their time in the upper surface

waters where the phytoplankton is most abundant (Kolar et al. 2005). But unlike the Bighead

15

carp, silvers are not often observed at the surface without being disturbed into motion often

resulting in their characteristic jumping behavior.

2.2 Net Construction and Design

The original idea for the capture of Silver Carp without altering their natural jumping

behavior was to use a 3 x 3 m surface level floating net with collection bag inserted (Figure 1).

However, when Silver Carp landed on this net, they were able to generate sufficient strength to

escape. Therefore, an elevated floating net was designed and constructed in partnership with the

Environmental Lab at ERDC in Vicksburg, Mississippi (Figure 2). Two-inch schedule 40 PVC

formed the frame of the net measuring 3.6 m long x 3.6 m wide x 0.75 m high. Nylon netting in

2.54 x 2.54 cm squares was sown around the top frame in two individual ‘baskets’ separated by

the middle support beams. These baskets hung above the bottom of the frame and thus did not

drag in the water during collection. Two pontoon-like floats were attached to each side of the

bottom frame via zip ties and used to elevate the frame out of the water during operation. Each of

the bottom frame pipes and vertical pipes were backfilled with foam to add rigidity to the

structure and reduce flexion. Rope was tied around each front corner of the bottom frame in

order to attach the net to the boat for towing. The floating net was designed to break into 3

individual pieces for storage and transport. Each piece was secured together using carriage bolts

and corresponding nuts.

16

Figure 1. Original surface level net for Silver Carp Sampling.

Figure 2. Elevated carp catching net designed with assistance from the engineering, research,

and development center.

17

2.3 Study Sites

Desoto Lake in Coahoma County, Mississippi (Figure 3) is a naturally occurring oxbow

of the Mississippi River which maintains one connection with the river and is completely

inundated during times of flooding and covers 14.5 km2. Depth is determined by the Mississippi

River and the banks are mostly unaltered and composed of willow trees, swamp privet, and

buttonbush (deltawildlife.org). Fish are free to move between the lake and the Mississippi River

year-round.

Lake Whittington is a man-made oxbow of the Mississippi River located along the

Mississippi/Arkansas border at Bolivar County, MS. The lake is connected directly to the

Mississippi River by a chute located on the southwestern end of the lake and the water level of is

directly related to the river. Once the river depth drops, constructed dikes cause water to drain

out of the system one way. During these periods of low water, three smaller ponds will separate

from the lake. During high water, the lake is roughly 9.3 km2 and banked by willow trees,

swamp privet, and rock banks.

Sampling within the Mississippi River took place near Cape Girardeau, MO over the

course of 3 days (July 2017). Areas sampled included the Castor River Diversion Channel,

Ramsey Creek Diversion Channel (a tributary of the Castor Diversion Channel which connects

to the Mississippi River), Apple Creek, and behind wing dikes between Apple Creek and Castor

River Diversion Channel. All areas were prone to downed trees and were directly connected to

the main Mississippi River channel. Both diversion channels and Apple Creek experienced algal

blooms during the late afternoon where carp were easily found (Figure 2).

18

Apple Creek is part of the Apple Creek Conservation Area along the Perry County-Cape

Girardeau Counties, MO. The stream is surrounded by steep hills and narrow valleys of upland

oak-hickory forest and some bottomland hardwoods. Treefalls and snags are common in the

stream providing ample fish habitat and areas of slack flow Silver Carp are likely using for

shelter.

Wing dikes along the Mississippi River have been seen to provide shelter for Silver Carp

from the extreme flow rates of the river and thus they can be found in large densities behind the

structures. There are a series of wing dikes stacked along the edge of the Mississippi River

passing through Perry County to Scott County MO channeling water into the shipping lane and

to prevent river bank erosion. Silver Carp were found immediately along the edge of the dikes

and the adjacent bank where water levels were shallow (0.8 m).

Three locations in Illinois were sampled during both summers (2016 and 2017): Bath

Chute, Illinois River at Havana, IL, and Spoon River. The Bath chute is a naturally occurring

backwater area of the Illinois River in Mason County, IL averaging 2.4 m and extending 21 km

forming the largest island on the Illinois River. This area is densely packed with Silver Carp and

is even home to an annual Silver Carp fishing tournament.

The Illinois River is a relatively slow moving and shallow river during the summer

regulated by a series of upstream locks and dams. The Spoon River is a slow flowing, winding

tributary of the Illinois river connecting at Havana, Illinois. The system is greatly unaltered with

agricultural prairie country along both banks. The water is deeper than in the Illinois River and

the system is more narrow.

19

Figure 3. Sample locations for 2016-2017.

2.4 Field and Post Field Analysis

To record fish leaps, two GoPro Hero 4 cameras were mounted on the stern of a 19’

Carolina Skiff. Cameras remained on and recording throughout the entirety of sampling. A 3.6 m

long x 3.6 m wide x 0.75 m high floating elevated net was used to capture leaping carp (Figure

2). The boat was maneuvered along the sampling location maintaining a speed between 8.05-

11.27 kph and fish either spontaneously leaped or were encouraged to leap by revving the boat

20

engine. Once a fish landed in the elevated floating net, the boat was stopped, water velocity was

measured (Flo-Mate 2000), temperature and dissolved oxygen at both the surface and bottom

were recorded (YSI 550A), depth of the water was recorded via a Hummingbird depth finder,

and an identification number was assigned to the fish and video recorded for later analysis. Fish

captured were then recovered from the net and total length, standard length, upper and lower

caudal lobe lengths, weight, and sex recorded. Each fish was dispatched via a quick, percussive

blow to the cranium for safety purposes and were disposed of in accordance with wildlife agency

regulations for each state. Silver Carp that landed in the boat but not captured on video were also

subjected to morphological measurements.

All video was analyzed using Avidemux 2.6 and Windows Movie Maker. The angle of

escape for every fish recorded was measured directly on the video using a protractor. A still

frame of those individuals captured in the net was used to measure body length, distance

traveled, and height traveled during each leap. Speed was calculated using the process described

in Parsons et al. (2016). Two GoPro Hero 4 cameras recording at 60 frames per second allowed a

measure of the time interval of when the tip of the nose just appeared at the waters’ surface and

when the tail fully exited the water. Absolute speed was then calculated by dividing fish length

by the time interval.

The number of frames required to exit the water 1 body length was then converted to

body lengths/second. For individuals positively identified, total length was multiplied by the

measured body lengths/second to determine the absolute speed (cm/s) achieved during the leap.

In some cases, the lengths of fish that were captured on video but not captured in the net were

estimated using known reference lengths provided by the frame of the capture net. This

methodology is a conservative estimate of burst speed and leap characteristics. In some cases the

21

caudle lobes exited the water between frames instead of at the end of a frame, yet whole frames

were counted. As a result, some time (up to 0.016 s) may have been added to some estimates.

Because the time was always added and never taken away, and due to the small amount of time,

the error was not considered high. This method becomes more efficient when higher frame rates

are used to record leaps or swimming performance.

2.5 Statistical Analysis

Descriptive statistics were used to compare morphological characteristics of fish across

the three study areas with one-way analysis of variance (ANOVA) used to test for significant

differences in height, weight, condition factor, sex, and caudal lobe lengths. T-tests were used to

test for differences between sex, size (estimated and observed), and caudal lobe lengths of the

entire population. ANOVA was also used to determine differences in leap characteristics across

study sites (distance, height, speed) while hierarchical regression was used to determine the

relationship between environmental and morphological characteristics with leap height, distance,

and burst speed each acting as a dependent variable. Analysis of covariance (ANCOVA) was

used to detect differences in speed across study sites. For burst speed, optimal angle and

distance/height analysis, total length of fish was eliminated as an effect by only analyzing fish

overlapping in size across all three study sites. Structural equation modeling was used as a

confirmatory method for the hierarchical regressions. The 0.05 probability level was used for all

significance testing. Tuckey Post Hoc analyses were used to determine differences signified by

ANOVAs. Data was inspected for normality and did not require transformation. All test were

completed using SPSS software by IBM.

22

CHAPTER 3: ANALYZING LEAP CHARACTERISTICS AND BURST SPEEDS OF

SILVER CARP (HYPOPHTHALMICHTHYS MOLITRIX) USING IN SITU VIDEO

ANALYSIS

3.1 Introduction

Silver carp Hypophthalmichthys molitrix, introduced into North America have rapidly

expanded their range exploiting vulnerable habitats, disrupting fisheries, and inflicting severe

ecological and economic damage (Kolar et al. 2005). Silver Carp have continued to spread into

the Middle Mississippi River and the Tennessee River Valley even as great effort is being

expended to prevent them from entering the Great Lakes and expanding further into the Ohio,

Illinois, Missouri, and Tennessee Rivers. High vagility, high fecundity (Williamson & Garvey

2005), and the ability to leap completely out of the water have aided in their rapid spread and

have complicated efforts to contain their movements.

In previous work, researchers have focused on how to facilitate fish passage past man-

made structures (Votapka 1991, Gowans et al. 1999, Cada 2011), but as invasive species have

continued to be introduced and spread, preventing fish movement past critical structures has

become important. For example, Sea Lamprey, Petromyzon marinus, were prevented from

entering the Great Lakes via an electrical barrier (Katopodis et al. 1994). In addition, hydrologic,

vertical drop, and water velocity barriers have been implemented in Great Lakes tributaries as

part of an integrated program to prevent spawning of Sea Lamprey (Heinrich et al. 2003, Siefkes

2017).

23

Currently there is an electric barrier in the Chicago Sanitary and Ship Canal preventing

entry of Silver Carp into Lake Michigan (McInerney et al. 2005) and other methods are being

investigated to prevent carp spread (Conover et al. 2007). However, concern exists that the

momentum of a rapidly swimming Silver Carp could carry the fish forward through the barrier

even if they are stunned (Parsons et al. 2016). Additionally, Silver Carp may be able to surmount

weirs or vertical drop barriers by leaping over them. Hydraulic barriers created by high velocity

outflows of locks and dams may prevent upstream movement. In constructing hydraulic barriers

Silver Carp maximum swimming speeds have been assumed to be similar to Pacific salmonids

due to their similar leaping behaviors, but no studies have definitively established their

maximum burst speeds (Hoover et al. 2016). Therefore, hydraulic barriers currently in use may

not prevent Silver Carp upstream movement but at the same time may prevent upstream

migration of native species (Stanley Consultants 2011). In addition, the leap characteristics of

Silver Carp, critically important for developing efficient physical and nonphysical barriers to

dispersal, have yet to be documented in-situ. In order to construct effective barriers for Silver

Carp without impeding waterway navigation or impeding native species, all aspects of their

behavior must be analyzed and, perhaps most importantly, their swimming abilities understood.

Previous studies relied on a variety of methods to determine the burst swimming speeds

of fish. These include high speed cameras (Wardle & He 1988, Videler & Wardle 1991), swim

tunnels (Parsons & Sylvester 1992, Hoover et al. 2012, Hoover et al. 2016), and hydraulic flumes

(Castro-Santos 2005). However, all cases involved removing the fish from the water and placing

them into a sealed system or transporting fish to a laboratory. Handling and confinement is

capable of stressing the fish leading to non-performers (Hoover et al. 2016) and precludes

leaping fish like Silver Carp from engaging in natural behaviors. Brunnschweiler (2005) used

24

high-speed video footage and projectile physics to measure at-sea, spontaneous blacktip shark,

Carcharinus limbatus, leaps, which eliminated the problem of capture stress. While this allowed

behaviors to be recorded and analyzed in-situ, it was greatly limited by the number of leaps

witnessed (3 leaps in 10 hrs.). However, Silver Carp are predisposed toward leaping and often

schools of a few to many thousands can be seen leaping simultaneously. Their behavior makes

them a perfect candidate for in-situ video recording and burst speed analysis.

This study aimed to document the in-situ leaping characteristics of Silver Carp across

multiple river systems and to examine variables that might affect those characteristics. Specific

objectives included: 1) determining burst speed; 2) determining angle of escape and establishing

if there is an optimal angle of escape; 3) determining height and horizontal distances of leaps; 4)

examining relationships between leap characteristics and environmental variables; and 5)

establishing variation in fish leaps across latitude.

3.2 Results

There was no significant difference in total length (t test, t32=2.04, p<0.339) between fish

whose lengths were measured directly and those estimated using known dimensions of the net

frame. Therefore, all fish were pooled for analysis involving total length. There was a significant

difference between total length and weight across the three study sites (ANOVA, F2,301=118.95,

p<0.001, F2,228=257.2, p<0.001 respectively) with the greatest difference between those in

Desoto Lake, Mississippi and the Illinois system (Tukey Post Hoc, 18.3 cm, p=0.005, 2.8 kg,

p<0.001 respectively). Fish in Mississippi (n=15) were significantly younger (1.5 years)

(Welch’s T-test, t51=-3.93, p<0.001) than fish in Illinois (n=38). Missouri fish were not aged.

There was no difference in length or weight between fish in the Missouri and Illinois systems

25

(Figure 4). Mean average total lengths among the three study sites were 54.79 + 9.3 cm (n=161)

for Illinois, 60.86 + 4.1 (n=30) for Missouri, and 73.09 + 11.05 cm (n=113) for Mississippi. The

largest fish measured was 92.3 cm long and weighed 6.21 kg. The largest fish estimated using

the net frame was 128 cm total length. The upper and lower caudal lobes were significantly

different in length (t409=-3.11, p<0.001) (Figure 5). There was a significant correlation between

total length and leap height (Pearson Cor84=0.294, p=0.007) as well as between total length and

distance (Pearson Cor81=0.363, p=0.001) (Figure 6). There were no differences between the

number of males and females captured across study sites. Condition factor did not significantly

vary across study sites.

Figure 4. Total length (black) and weight (grey) of Silver Carp captured from each location

(n=113, 30, 161 respectively). Means and standard errors are shown.

0

0.5

1

1.5

2

2.5

3

3.5

4

4.5

5

0

10

20

30

40

50

60

70

80

Mississippi Missouri Illinois

Wei

ght (

kg)

Tota

l Len

gth

(cm

)

26

Figure 5. Comparison of upper and lower caudal lobe lengths. Means and standard errors are shown.

1010.210.410.610.8

1111.211.411.611.8

Upper Lobe Lower Lobe

Leng

th (c

m)

a

27

Figure 6. Histograms of leap distance (a) (n=81) and leap height (b) by total length (n=81). 95%

CI are shown.

Burst speeds (cm/s) across study sites were significantly different (ANCOVA, F2, 84 =

18.964, p<0.001) with the fastest fish in Missouri (628.4 + 99.97, n=10, Tukey Post Hoc

p<0.01) followed by Mississippi (485.29 + 109.82, n=42) and Illinois (524.25 + 122.15, n=35)

(Figure 7). The maximum speed recorded of a captured fish was 771 cm*s-1 by a 77.1 cm fish

traveling at 10 body lengths per second whereas the maximum estimated speed was 1280 cm*s-

1by a 128 cm fish traveling 10 body lengths per second. The slowest burst speed calculated was

224.2 cm*s-1 by fish in Mississippi. Silver Carp of various total lengths exited the water at the

same relative speeds (Figure 8). Fish collected near Cape Girardeau, Missouri leaped greater

heights on average (ANOVA, F2,81=6.2012, p=0.003, Tukey Post Hoc p<0.005) with no

significant differences in leap height between Mississippi and Illinois fish (Tukey Post Hoc

b

Total Length (cm)

28

p=0.895) (Figure 9, Figure 10). Hierarchical regression was used to develop an equation for

predicting burst speed based on morphological and environmental variables. The only significant

variables were temperature, water velocity of the system and total length of the fish (adjusted R2

= 0.236);

𝐵𝐵𝐵𝐵𝐵𝐵𝐵𝐵𝐵𝐵 𝑆𝑆𝑆𝑆𝑆𝑆𝑆𝑆𝑆𝑆 = 139.282(𝑤𝑤𝑤𝑤𝐵𝐵𝑆𝑆𝐵𝐵 𝑣𝑣𝑆𝑆𝑣𝑣𝑣𝑣𝑣𝑣𝑣𝑣𝐵𝐵𝑣𝑣) + 4.377(𝐵𝐵𝑣𝑣𝐵𝐵𝑤𝑤𝑣𝑣 𝑣𝑣𝑆𝑆𝑙𝑙𝑙𝑙𝐵𝐵ℎ) +

16.117(𝐵𝐵𝑆𝑆𝑡𝑡𝑆𝑆𝑆𝑆𝐵𝐵𝑤𝑤𝐵𝐵𝐵𝐵𝐵𝐵𝑆𝑆) − 321.784.

Figure 7. Regressions of burst speed on total length for each study site.

R² = 0.3552

R² = 0.1407

R² = 0.193

0

100

200

300

400

500

600

700

800

900

20 40 60 80 100

Burs

t Spe

ed (c

m*s

-1)

Total Length (cm)

Mississippi

Missouri

Illinois

Linear (Mississippi)

Linear (Missouri)

Linear (Illinois)

29

Figure 8. Scatterplot of total length (cm) and relative speed (body lengths*s-1).

0

2

4

6

8

10

12

14

16

0 20 40 60 80 100

Rel

ativ

e Sp

eed

(bod

y le

ngth

s*s-1

)

Total Length (cm)

MississippiIllinoisMissouri

0

100

200

300

400

500

600

Leap

Dis

tanc

e (c

m)

Burst Speed (cm/s)

a

30

Figure 9. Leap distance (a) (n=84) and leap height (b) (n=84) against burst speed for Silver Carp

from each study site. Scatterplot (c) of burst speed and height shows the significant correlation

0

50

100

150

200

250

300

350Le

ap H

eigh

t (cm

)

Burst Speed (cm/s)

R² = 0.0349

R² = 0.0528R² = 0.3933

0

50

100

150

200

250

300

200 400 600 800

Leap

Hei

ght (

cm)

Burst Speed (cm*s-1)

IllinoisMissouriMississippiLinear (Illinois)Linear (Missouri)Linear (Mississippi)

c

b

31

found for Mississippi fish. Means and 95% confidence intervals are shown. Bars without

confidence intervals only had one sample.

Figure 10. Leap distance, leap height, and burst swimming speed for all study sites. Means and

standard errors shown.

To test for an optimal leap angle (the angle that maximizes horizontal distance traveled),

angles of escape were compared with the horizontal distance achieved by each fish (Figure 11).

However, there was no significant difference for leap distances at any given angle range. Leap

angles followed a normal distribution (Figure 12) with carp leaping at a mean angle of 55.3° and

a mode of 47°. The majority of angles used were between 46 to 65° while the least common

angles were below 23° and above 80°. A 2-way ANOVA showed a significant relationship

between speed and distance (F26,1=1.875, p=0.028) but no relationship between angle and

0

100

200

300

400

500

600

700

0

50

100

150

200

250

Mississippi Missouri Illinois

Burs

t Spe

ed (c

m*s

-1)

Tota

l Len

gth

(cm

)

Absolute DistanceAbsolute HeightBurst Speed

Distance Height Burst Speed

32

distance or an interaction between speed and angle (Figure 13). The longest distances were

achieved at speeds ranging from 524 to 643 cm*s-1 and angle of 37 to 76°. Burst speed, angle of

escape, and total length were used in the hierarchical regression equation;

Distance =2.382(Total Length) -0.035(Burst speed) – 1.639(Angle) + 174.693

and were significant (adjusted R2 = 0.165) predictors of H. molitrix leap distance though there. A

path analysis (R2 = 0.21) further explained the variation in distance and confirmed the

relationships between total length, speed, and angle with distance. Path analysis suggests that

total length of fish is the most important determinant of the total distance an individual can travel

(Figure 14).

Figure 11. Average leap distances against angle of escapes (n=78). Means and 95% confidence

intervals are shown. Bars without confidence intervals only had one sample. There was no

significant difference for distance at any angle range.

0

50

100

150

200

250

300

350

25-30 31-35 36-40 41-45 46-50 51-55 56-60 61-65 66-70 71-75 76-80

Leap

Dis

tanc

e (c

m)

Angle of Escape (degrees)

33

Figure 12. Frequency distribution of leap angle for Silver Carp. The majority of angles were

between 46-65° (66.1%) while <3% of fish exited at angles below 23° and above 80°.

34

Figure 13. Plot of burst speed (cm/s) against leap distance (cm) with the interaction of angle

(degree).

Figure 14. Path analysis for distance (cm) showing correlation values.

Angle (degrees)

Leap

Dis

tanc

e (c

m)

Burst Speed (cm/s)

35

Angle of escape and height were significantly correlated (Pearson Cor 85=0.247, p=0.02)

(Figure 16). The hierarchical regression;

𝐻𝐻𝑆𝑆𝑣𝑣𝑙𝑙ℎ𝐵𝐵 = 1.771(𝐴𝐴𝑙𝑙𝑙𝑙𝑣𝑣𝑆𝑆) + 1.125(𝑇𝑇𝑣𝑣𝐵𝐵𝑤𝑤𝑣𝑣 𝐿𝐿𝑆𝑆𝑙𝑙𝑙𝑙𝐵𝐵ℎ) + 0.050(𝐵𝐵𝐵𝐵𝐵𝐵𝐵𝐵𝐵𝐵 𝑆𝑆𝑆𝑆𝑆𝑆𝑆𝑆𝑆𝑆) − 84.451

was a significant (adjusted R2 = 0.321) predictor of Silver Carp leap height. A path analysis (R2 =

0.34) further explained the variation in height and confirmed the relationships between total

length, speed, and angle with height (Figure 16). In the case of height, angle of escape was the

most important factor contributing to height.

R² = 0.3515

R² = 0.0656

R² = 0.0767

0

50

100

150

200

250

300

20 40 60 80 100

Hei

ght (

cm)

Angle of escape

IllinoisMissouriMississippiLinear (Illinois)Linear (Missouri)Linear (Mississippi)

36

Figure 15. Correlation of height and angle of escape.

Figure 16. Path analysis for height (cm) showing correlation coefficients.

3.3 Discussion

Invasive species are often generalist capable of outperforming and outcompeting native

species (Cortes et al. 2016). Limiting the dispersal and potential environmental damage of

invasive species requires an understanding of their behavior, physiology and ecology. While

ecological and physiological studies on fish are common, behavioral studies have only recently

been implemented to address fish performance capabilities (Holway & Suarez 1999). Behavioral

analysis is a useful tool for designing in-situ studies on fishes swimming abilities and habitat use

and provides a pathway for examination of burst speeds.

3.3.1 Burst Speed:

37

While swimming performance studies have been completed on a variety of fish species,

burst swimming measurements are notoriously difficult to obtain and few studies have been

completed. In previous studies burst swimming speeds were determined using high speed

cameras (Wardle & He 1988, Videler & Wardle 1991), swim tunnels (Parsons & Sylvester 1992,

Hoover et al. 2012, Hoover et al. 2016), and hydraulic flumes (Castro-Santos 2005). However,

all cases involved removing the fish from the water and placing them into a sealed system or

transporting to a laboratory. Brunnschweiler (2005) suggested that observations of leaping fish

demonstrates a ‘standing jump’ (as opposed to porpoising) and provide for the best opportunity

of measuring maximum swimming speeds. Parsons et al. (2016) calculated burst speed using

projectile physics and video footage of leaping Silver Carp but further noted that video footage

of jumping fish can be used to determine speed directly when fish size can be obtained.

Furthermore, Parsons et al. (2016) showed a frames per second conversion approach provides

values on par with projectile physics estimates. However, field measurements still need to be

obtained to validate bust speed estimations.

Hoover et al. (2017) used a mobile swim tunnel to measure swimming speeds of Silver

Carp but did not stimulate fish to leap in the confined system. The Bighead, Hypophthalmichthys

noblis, and Silver Carp swam in their study showed swim speeds less than half that of Sockeye

Salmon, the assumed comparable fish for hydrological control. At 0.5 min Silver Carp averaging

80.1 cm were only able to maintain 2.57 body lengths per second. This is 2 to 3 times below our

average of 7.35 + 0.97 body lengths*s-1 for individuals ranging between 60 and 80 cm (Table 1).

However, Hoover et al. (2017) presented swim tunnel models which when extrapolated to < 1s

show burst speeds comparable to those in this study. This suggests that fish may be capable of

sustaining these high velocities only for a few seconds. Burst swimming speeds lead to

38

exhaustion within 30 s of maintained swimming. By leaping, Silver Carp may be capable of

bursting multiple times because the muscles can relax during the leap and prolong the time to

exhaustion. At present, no work has been conducted on the time to exhaustion from burst

swimming on Silver Carp.

Videler and Wardle (1991) predicted that adult fish are not capable of swimming more

than 10 body lengths* s-1. Indeed, multiple fish in this study reached more than 10 body

lengths*s-1 and demonstrated the need to analyze burst speeds under natural conditions. Multiple

fish obtained speeds of 14 body lengths*s-1 across a range of sizes showing these fish

overperform expectations. Other species known to break the 10 body lengths*s-1mark are

members of the family Scombridae which includes the mackerels and tunas. Tuna in particular

have been documented traveling at 20 body lenths*s-1 (Fierstine and Walters 1967) but their

superior performance may relate to their thermoregulatory ability. Sockeye Salmon,

Oncorhynchus nerka, generated burst speeds greater than 4.25 body lengths*s-1 (230 cm*s-1)

(Brett 1982, Hoover et al. 2016). However, this is still below the capabilities of Silver Carp

reported in this study. The data reported herein shows fish between 55 to 65 cm attaining the

highest burst speed between 500 and 700 cm*s-1. This suggests that using Sockeye salmon as a

standard baseline for performance when planning Silver Carp management will result in a

significant underestimation of carp performance abilities unless salmon are analyzed in-situ as

well. Indeed, Silver Carp attained speeds closer to that of a leaping blacktip shark than sockeye

salmon (Table 1). To my knowledge, the Silver Carp speeds reported herein are the highest of

any freshwater species and are similar to continuously swimming, migratory oceanic species.

Additionally, our study demonstrates the applicability of video-taped behavior for measuring

39

burst swimming speeds and suggests other methods may lead to significant underestimations of

performance.

40

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41

In general, stride length (amount moved during one tail thrust) and absolute speed

increases with fish total length (Cheong et al. 2006). However, this was not true for Silver Carp.

While the largest fish were captured in Mississippi, significantly higher burst speeds were

recorded for the smaller Missouri fish. One explanation for the burst speed difference is the good

athlete/bad athlete hypothesis (Reidy et al. 2000) which would suggest that genetic differences in

populations may lead to fish with greater performance abilities. However, the more parsimonious

explanation is the training effect. Fish living in the main Mississippi River or immediate adjacent

tributaries are exposed to higher water velocities year-round when compared to those in an

oxbow lake or the much slower Illinois River. As a result, the fish are forced to swim against

stronger, faster currents on a regular basis and are trained at higher speeds (Davison 1997,

Martinovic 1999). Martinovic 1999 also found that trained Grass Carp performed better than

untrained control fishes and recovered more quickly. Lactate in trained fish leveled off at 30 min

post swim trials whereas untrained fish lactate leveled off at 130 mins. This suggest river trained

fish may be capable of bursting in quick succession if sheltered areas (wing dikes, eddies, slack

flow fields) are present near high velocity flow fields.

3.3.2 Leaping Abilities and Characteristics

The leaping abilities of Silver Carp are well known although only one paper has been

published describing the behavior. Parsons et al. (2016) examined Silver Carp leaping ability

using public online videos. However, that study did not provide the in-situ measurements

documented here and was limited in sample size and location. Leaping behavior in fishes is

explained using multiple hypotheses; 1.) traversing migrational obstacles (Aronson 1971,

Warburton 1990, Lauritzen et al. 2010), 2.) respiration (Hoese 1985, Moyle et al. 1986), 3.)

predation (Lowry et al. 2005, Shih and Techet 2010, Soares and Bieman 2013), 4.)

42

communication (Sulak et al. 2002), 5.) parasite shedding (Cochran et al. 2003, Brunnshweiler

2006), and 6.) predator escape (Saidel et al. 2004, Gibb et al. 2011, Soares and Beirman 2013,

Parsons et al. 2016).The predator escape hypothesis may be more applicable to this study since

Silver Carp are often observed leaping in response to vessel created stimuli. Assuming a predator

escape hypothesis, I hypothesized that fish utilize an optimal angle of escape to maximize the

horizontal distance traversed and therefore put the greatest distance between themselves and a

potential predator. Optimality theory dictates that animals will choose behaviors which optimize

some benefit (Cody 1974) such as energy allocation, fitness, or in this case, overall survival. The

data presented here revealed that fish choosing to leap between angles of 37 to 46° and 67-76°

achieved the greatest horizontal distance. However, 66% of fish leaped at angles between 46-65°

and only 28.1% between 37 to 46 and 67 to 76°. Distance was significantly affected by burst

speed while burst speed and angle of escape had a non-significant interaction suggesting burst

speed and fish size were more important than choosing a specified angle to exit the water. A

parsimonious explanation for why Silver Carp do not optimize their leap distance with the best

angle of escape is Silver Carp are schooling fish and in a predator escape response may simply

choose an angle that allows them to prevent collision with conspecifics.

Both analytic models created in this study for predicting leaping distance agree that total

length of the fish, burst speed, and angle of escape were the most important determinates of

leaping distance. Burst speed in turn brings water velocity and water temperature forward as

prominent predictor variables. This combination demonstrates the importance of both fish

morphological and performance characteristics in addition to ecological variables. However, the

models only explained 21% of the variation, which suggests more exploration is needed to

accurately predict leap distance across the geographical range of Silver Carp.

43

Across all of our study sites, Silver Carp were capable of traversing, in a single leap,

great distances (482.34 cm) and heights (276.06). Silver Carp on average traveled 1.8 body

lengths in height comparable to the leaping abilities of other species (Table 2). However, the

variation in Silver Carp leaping ability is large across all sizes which indicates managers and

barrier designers should consider the maximal capabilities of Silver Carp instead of relying on

averages for performance and leaping. Furthermore, analysis of leap heights suggests a

combination of factors best explains leaping ability. This ability demonstrates their potential for

traversing navigational obstacles, surmounting hydraulic and physical barriers and thus

increasing their range. The maximum height (276.06 cm) observed in this study, is likely not the

greatest height that Silver Carp can achieve but should represent a baseline value for future

barrier design.

44

Table 2. Published leap heights for multiple species.

Species Mean Length (cm) Mean Height (cm) Reference

Silver Carp

62 113 This study

Aral Barbel 35-45 99.8 Lopez & Tolosa

2017

Goldfish 35-45 53.2 Lopez & Tolosa

2017

Brook Trout

15-20 73.5 Kondratieff &

Myrick 2005

While there were no significant differences between study sites in the horizontal distance

traveled by leaping carp, fish in Missouri reached significantly higher heights. Fish in Missouri

also went vertically higher than they did horizontally at these higher speeds. The values reported

herein may be important in designing hydraulic and physical barriers to prevent continued Silver

Carp dispersal. These characteristics are all interrelated with total length and burst speed

influencing both the distance and height for each leap. Because of this, wildlife officials cannot

focus on one characteristic without careful consideration of others. Furthermore, effects of the

environment were not fully explored in this study and may influence the development or training

these fish experience throughout their range.

45

3.3.3 Geographic Variation

Environmental conditions vary with latitudinal changes and can greatly impact the life

history of fishes even living in interconnected water systems. Many studies have focused on

morphological and physiological changes occurring within species as latitude and altitude

changes since Bergmann published his statements in 1847. Bergmann’s rule (1847) revised by

Rensch (1938) is an ecogeographical principle describing how homeotherms of the same

taxonomic group tend to increase in size with increasing latitude in order to energetically

compensate for declining temperature. However, ectothermic fish may follow the opposite trend

with size increasing as latitude decreases. Several species of fish show an increase in size-at-age

at lower latitudes (Parsons 1993, Lombardi-Carlson et al. 2003) similar to the findings in this

study. The lengthened growing season nearer the equator and increased phytoplankton and

zooplankton production may result in larger Silver Carp in the Lower Mississippi River Basin.

As a result, fish at lower latitudes may reach maturity before conspecifics at higher latitudes. For

example, there was a mean increase in age to maturity as latitude increased for Red Bandfish

(Cepola mcrophthalms) and Brown Trout (Salma trutta) (L’Abee-Lund et al. 1989, Stergiou

1999). Furthermore, the optimal temperature range for Silver Carp (24-31°) is present for the

majority of the year in the Lower Mississippi River. Fish in this study were largest in a

Mississippi oxbow which was the lowest latitude study site. Oxbow lakes in the Lower

Mississippi River experience large nutrient loads from runoff and have a greatly reduced rate of

flow explaining the larger average size of fish caught there. As a result, more energy is available

than is required to meet metabolic needs allowing southern fish to allocate more energy toward

initial growth. Garvey and Marschall (2003) documented such a trend in Largemouth bass

(Micropterus salmoides) and suggest older, reproductively mature fish had higher fecundity at

46

lower latitudes once maximum length was achieved. If this is the case, larger, younger fish at

lower latitudes should be considered in all management strategies and population estimates.

While this trend is common for freshwater teleosts, Parsons (1993) and Lombardi-Carlson et al.

(2003) documented that the Bonnethead Shark, Sphyrna tiburo grew faster at higher latitudes

along the Florida coast as a way to compensate for the shorter growing season in comparison

with those captured in southern Florida.

Most flying or leaping fish possess multiple unique morphological qualities including a

well-defined ventral keel and extended lower caudal lobe. While there was a significant

difference between caudal lobes, there were not enough fish with equal caudal lobe lengths

captured to investigate differences in speed, height, or distance from this feature. It is possible

this morphological feature may be selected for over time improving the leaping abilities of carp.

47

CHAPTER 4 CONCLUSIONS/RECOMMENDATIONS

The analysis of leaping behavior and burst speed in Silver Carp revealed that there are

significant differences in leap height and burst swimming speed of Silver Carp across the range

of this study. Faster fish were recorded in Missouri where water velocity was highest which

suggests there is an environmental impact on the leaping characteristics of Silver Carp. Burst

speed increased with total length of fish and conspecifics of the same size burst faster in this

study than in a previous swim tunnel-based study. Indeed, these may be the fastest speeds

recorded for freshwater fish and are more similar to those of oceanic species. Silver Carp did not

use an optimal angle to cover the greatest horizontal distance during their leaps, but angle did

play a significant role in determining the distance and height fish traversed. Speed and total

length were the most important factors in determining distance covered in each leap. Finally,

Silver Carp size increased with decreasing latitude across all study sites. To my knowledge these

are the highest burst speeds documented for freshwater fish and resemble those of oceanic

species.

Together the data presented here provided fundamental knowledge needed for the future

control of Silver Carp. Invasive species like carp are capable of superior performance in novel

habitats and take advantage of ecological gaps and anthropogenic alterations. This allows the

invaders to overtake the local biomass and increase their population in a short amount of time.

Managers must make decisions rapidly in order to prevent range expansion and establishment of

invasive species such as Silver Carp. For example, any hydrologic barriers must meet at least the

48

average burst speeds reported here to stop upstream migration of Silver Carp. Furthermore, a

comparison of Silver Carp burst speeds with native fishes of concern may allow for hydrologic

barriers to improve efficacy and aide conservation efforts. Silver Carp are an invasive species

that has permanently established itself in North America, but with the data herein and future

studies on the endurance of Silver Carp, control efforts may be successful. Overall, this study

shows Silver Carp performance is higher than initially thought and leaping characteristics are

explainable by an interaction of morphological and environmental factors.

49

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58

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59

VITA

Professional Preparation

Bachelor of Science University of Mississippi 2015 – Biology

Academic Appointments

2015-2017 Research Assistant, Army Corps of Engineers, Vicksburg, MS

2015-2017 Graduate Research Assistant, Department of Biology, University of Mississippi

2015-2018 Graduate Teaching Assistant, Department of Biology, University of Mississippi

2013-2015 Undergraduate Research Assistant, Department of Biology, University of

Mississippi

2014 Teaching Assistant, Department of Biology, University of Mississippi

Presentations – Oral

E. Stell, J. J. Hoover, G. R. Parsons, 2016. Why do fish jump? Mississippi Chapter of the

American Fisheries Society, Terra, MS.

E. Stell, J. J. Hoover, G. R. Parsons, 2016. Jumping characteristics of Silver Carp. Joint Meeting

of Ichthyologist and Herpetologist, New Orleans, LA.

60

E. Stell, J. J. Hoover, G. R. Parsons, 2018. Analyzing the leaping characteristics and burst

swimming speeds of Silver Carp using in-situ video analysis. Mississippi Chapter of the

American Fisheries Society, Oxford, MS. 2nd place presentation.

Publications

Parsons, G.R., E. Stell, and J.J. Hoover. 2015. Estimating burst swim speeds and jumping

characteristics of Silver Carp (Hypophthalmichthys molitrix) using video analyses and

principles of projectile physics. ANSRP Technical Notes Collection. ERDC/TN

ANSRP-xx-x. Vicksburg, MS: U.S. Army Engineer Research and Development Center.

Stell, E. G., J. J. Hoover, B. A. Cage, D. Hardesty, G. R. Parsons. 2017. Long distance

movements of four Paddlefish, Polyodon spathula, from a remote oxbow lake in the

Lower Mississippi River Basin. In publication.